Ll . : sn a Pa E s IS AA aa cn : : A . el EICDI PAR A , 7 h Ln E í Fire A Meri e o Mo E EPS : Daza ino Laa e ñ miei as > > z AAA TAR CAS SR IES tii As S .. a DA A el : + . A A NS ES | : É $ Fo ENE É A : | es lea A y | o EIN ; : : ae DR OS PEI Errar . : ns IT A EA A demore EN ra ARA da E Dali cre A Mt SED HARVARD UNIVERSITY 3 Library of the Museum of Comparative Zoology ÍS e O A 0 Es berus Vol. 15 (1) REVISTA DE LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA Oviedo, junio 1997 COMITÉ DE REDACCIÓN EDITOR Ángel Guerra Sierra EDITORES ADJUNTOS Eugenia M* Martínez Cueto-Felgueroso Francisco Javier Rocha Valdés Gonzalo Rodríguez Casero ComuTé EDITORIAL Kepa Altonaga Sustacha Eduardo Angulo Pinedo Thierry Backeljau Sigurd v. Boletzky Jose Castillejo Murillo Karl Edlinger José Carlos García Gómez Edmund Gittenberger Serge Gofas Ángel Antonio Luque del Villar María Yolanda Manga González Jordi Martinell Callico Iberus Revista de la SOCIEDAD ESPAÑOLA DE MALACOLOGÍA Instituto de Investigaciones Marinas, CSIC, Vigo, España Universidad de Oviedo, Oviedo, España Instituto de Investigaciones Marinas, CSIC, Vigo, España Universidad de Oviedo, Oviedo, España Universidad del País Vasco, Bilbao, España Universidad del País Vasco, Bilbao, España Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica Laboratoire Arago, Banyuls-surMer, Francia Universidad de Santiago de Compostela, Santiago de Compostela, España Naturhistorisches Museum Wien, Austria Universidad de Sevilla, Sevilla, España Notionaal Natuurhistorisch Museum, Leiden, Holanda Muséum National d'Histoire Naturelle, Paris, Francia Universidad Autónoma de Madrid, Madrid, España Estación Agrícola Experimental, CSIC, León, España Universidad de Barcelona, Barcelona, España Ron K. 0'Dor Dalhousie University, Halifax, Conada Marco Oliverio Universitá di Roma “La Sapienza”, Roma, Italia Pablo E. Penchaszadeh Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina Carlos Enrique Prieto Sierra Universidad del País Vasco, Bilbao, España María de los Ángeles Ramos Sánchez Museo Nacional de Ciencias Naturales, CSIC, Madrid, España Paul 6. Rodhouse British Antarctic Survey, Cambridge, Reino Unido Joandoménec Ros ¡ Aragones Universidad de Barcelona, Barcelona, España María del Carmen Salas Casanovas Universidad de Málaga, Málaga, España Gerhard Steiner Universitát Wien, Austria José Templado González Museo Nacional de Ciencias Naturales, CSIC, Madrid, España Victoriano Urgorri Carrasco Universidad de Santiago de Compostela, Santiago de Compostela, España Anders Warén Swedish Museum of Natural History, Estocolmo, Suecia 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: Dr. Ángel Guerra Sierra, Instituto de Investigaciones Marinas (CSIC), c/ Eduardo Cabello 6, 36208 Vigo, España. Los trabajos se entregarán por triplicado (original y dos copias). Se recomienda a los autores leer cuidadosa- mente 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 dirigirse 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 Electrónico: backhuysCeuronet.nl PORTADA DE Jberus Iberus gualterianus (Linnaeus, 1758), una especie emblemática de la península Ibérica, que da nombre a la revista. Dibujo realizado por Toza. Iberus REVISTA DE LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA Vol. 15 (1) Oviedo, junio 1997 Dep. Leg. B-43072-81 ISSN 0212-3010 Diseño y maquetación: Gonzalo Rodríguez Impresión: LOREDO, S. L. - Gijón O Sociedad Española de Malacología Iberus, 15 (1): 1-4, 1997 How many species of Candidula (Gastropoda: Hygromii- dae) in northern Portugal? ¿Cuantas especies de Candidula (Gastropoda: Hygromiidae) habitan en el norte de Portugal? Cristian R. ALTABA* Recibido el 6-11-1996. Aceptado el 24-IV-1996 ABSTRACT Two species of Candidula Kobelt, 1871 (Pulmonata: Hygromiidae) endemic to Portugal are reported from the northern part of the country. C. belemensis (Servain, 1880) is consi- dered a valid species similar to, but different from the widespread C. intersecta (Poiret, 1801). C. olisippensis (Servain, 1880) is distinctive, having a shell with very small, round umbilicus, a long spermathecal duct, and a short flagellum. RESUMEN Se citan para el norte de Portugal dos especies endémicas en este país pertenecientes al género Candidula Kobelt, 1871 (Pulmonata: Hygromiidae). C. belemensis (Servain, 1880) es considerada como una especie válida similar, pero diferente, a la ampliamente distribuidaC. intersecta (Poiret, 1801). C. olisippensis (Servain, 1880) es de fácil distin- ción, al tener una concha con un ombligo redondeado y pequeño, un largo conducto de la espermateca y un corto flagelo. KEY WORDS: Gastropoda, Hygromiidae, Candidula, Portugal, distribution, taxonomy. PALABRAS CLAVE: Gastropoda, Hygromiidae, Candidula, Portugal, distribución, taxonomía. INTRODUCTION The genus Candidula Kobelt, 1871 comprises several species of hygromiid land snails living in Western Europe. In the Iberian Peninsula this genus has un- dergone a remarkable diversification, particularly along the Atlantic drainages (ALTONAGA, GÓMEZ, MARTÍN, PRIETO, PUENTE AND RALLO, 1994). The taxo- nomy of this genus is complex because the diagnostic characters of the various species are not conspicuous, either in the internal anatomy or in the shells. The species of Candidula living in Portu- gal are still poorly known, yet this country has at least four endemics (AL- TIMIRA, 1969; GITTENBERGER, 1985, 1993). This note is a contribution towards a re- vision of the genus in the Iberian Penin- sula (Ondina and Altaba, in prep.). Two little-known, apparently rare species of Candidula are reported here, collected during a field trip in April *Institut Mediterrani d'Estudis Avancgats (CSIC-UIB). Ctra. de Valldemossa, Km 7. 5, 07071 Palma de Mallorca. Iber IIIO 1994 in northern Portugal. These species were described by SERVAIN (1880), and subsequently ignored for over a century (NOBRE, 1930, 1941; SErIxas, 1992). Con- RESULTS chological and anatomical data indicate that the species studied here are indeed distinct taxa. The specimens are kept in the author's malacological collection. Family HYGROMIIDAE Tryon, 1866 Genus Candidula Kobelt, 1871 Candidula belemensis (Servain, 1880) (Figs. 1, 2) C. belemensis was known up to date only from the original locality (Lisbon) and three other sites further south (GrT- TENBERGER, 1993). A single fresh shell was found at the base of the old city walls of Valenca do Minho, at the nort- hern border of Portugal (UTM 29T NG25; Figs. 1, 2). This locality represents a considerable extension of the species” known range. It is likely, however, that C. belemensis lives also further north, in Galicia, where specimens having a long flagellum have been reported as C. inter- secta by CASTILLEJO (1986). This species is probably closely rela- ted to C. intersecta (Poiret, 1801), which ranges discontinuously throughout Atlantic Europe, from southern Portugal to southern Sweden (KERNEY AND CA- MERON, 1979; GITTENBERGER, 1993; AL- TONAGA et al., 1994). C. belemensis differs conchologically from the latter in having a more depressed shell with a broader aperture, a wide and slightly eccentric umbilicus, and a much shallower sculp- ture of radial ribs and minute spiral striae (GITTENBERGER, 1993). Anatomically it differs from NW-Euro- pean C. intersecta in the relatively longer flagellum, which is about half the length of the epiphallus (GITTENBERGER, 1993). However, populations of C. intersecta from NW-Iberia have a flagellum shorter than half the length of the epiphallus, as is typical of that species (MANGA GONZÁ- LEZ, 1979). Thus, the available evidence suggests that C. belemensis is not conspe- cific with co-occurring C. intersecta. Candidula olisippensis (Servain, 1880) (Figs. 3-6) Two subadult specimens were col- lected under bolders in the vicinity of the Sanctuary of Sameiro, near Braga (UTM 29T NE59; Figs. 3, 4). Although the aperture edge of their shells was still tender and became damaged, it bears a conspicuous whitish internal rib. The radulae have one central, and 22 and 24 lateral teeth. These numbers fall within the range of other Candidula species from northwestern Iberia (MANGA GON- ZÁLEZ, 1979). The genitalia (still immature) of both specimens are shown in Figures 5 and 6. The flagellum is distinct, proximally na- rrow, and very short, measuring about 1/4 the length of the epiphallus, which is somewhat shorter and thinner than the penis. The bursa is small, oblong and in- distinctly united to its duct, which lies appressed to the spermoviduct throug- hout and is exceedingly long, measuring almost twice the length of the penis and epiphallus combined. There are no glan- dulae mucosae, their location being oc- cupied instead by a moderate swelling at the proximal end of the vagina. The dart sac appears partially developed, being only a large medial swelling of the vagina with no trace of a dart, but exhi- biting two large internal longitudinal folds adjacent to two shallow invagina- tions, the lower one bordered by a few papillae. This internal structure can be interpreted as an immature stage of that described for other species of Candidula ALTABA: How many species of Candidula in northern Portugal? Figures 1-4. Shells of Candidula from northern Portugal. 1, 2: C. belemensís from Valenga do Minho (CRA 4895); 3, 4: C. olisippensis from Sameiro, near Braga (CRA 4866-1). Scale bar 5 mm. Figuras 1-4. Conchas de Candidula del norte de Portugal. 1, 2: C. belemensis de Valenga do Minho (CRA 4895); 3, 4: C. olisippensis de Sameiro, cerca de Braga (CRA 4866-1). Escala 5 mm. Figures 5, 6. Proximal genitalia of two immature specimens of Candidula olisippensis. Abbreviations. A: atrium; E: epiphallus; F: flagellum; M: penial retractor muscle; P: penis; S: bursa copulatrix; SD: duct of the bursa; SO: spermoviduct; V: vagina; VD: vas deferens. Scale bar 2 mm. Figuras 5, 6. Genitalia proximal de dos especimenes inmaduros de Candidula olisippensis. Abreviaturas. A: atrio; E: epifalo; E: flagelo; M: músculo retractor peneal; P: pene; S: bursa copulatrix; SD: conducto de la bursa; SO: espermoviducto; V: vagina; VD: vaso deferente. Escala 2 mm. Iberus, 15 (1), 1997 (HAUSDORE, 1988, 1991), confirming the generic assignation by GITTENBERGER (1993), who examined a dried specimen in poor condition. Several nominal species described by LOCARD (1899) are probably sy- nonyms of Helix olisippensis Servain 1880. The general shape of the shell in this species (or species complex) is fairly variable, yet it is always fairly thin, with ACKNOWLEDGEMENTS My wife Catalina Ponsell helped in the field and provided the necessary conditions for the preparation of the manuscript. We are indebted to Or- lando Moreira and his family for in- comparable hospitality in Portugal and BIBLIOGRAPHY ALTIMIRA, C., 1969. Notas malacológicas. Pu- blicaciones del Instituto de Biología Aplicada, 46: 91-113. ALTONAGA, K.; GÓMEZ, B.; MARTÍN, R.; PRIETO, C.R.; PUENTE, A.I. AND RALLO, A., 1994. Es- tudio faunístico y biogeográfico de los moluscos terrestres del norte de la Península Ibérica. Eusko Legebiltzarra, Vitoria-Gasteiz, 505 pp. CASTILLEJO, J., 1986. Caracoles terrestres de Ga- licia. Familia Helicidae (Gastropoda, Pul- monata). Monografías de la Universidad de San- tiago de Compostela, 122: 1-65. GITTENBERGER, E., 1985. The taxonomic status of Xeroplexa Monterosato, 1892 (Pulmonata: Helicidae: Helicellinae), a surprise. Iberus, 5: 59-62. GITTENBERGER, E., 1993. Digging in the grave- yard of synonymy, in search of Portuguese species of Candidula Kobelt, 1871 (Mollusca: Gastropoda Pulmonata: Hygromiidae). Zo- ologische Meddedelingen, 67 (17): 283-293. HAUSDORE, B., 1988. Zur Kenntnis der syste- matischen Beziehungen einiger Taxa der He- licellinae Ihering 1909. Archiv fiir Mollusken- kunde, 119 (1/3): 9-37. HAUSDORE, B., 1991. Uber zwei Candidula-Ar- ten von der súdlichen Balkanhalbinsel (Gas- tropoda: Hygromiidae). Archiv fur Mollus- kenkunde, 120 (4/6): 119-129. rounded or slightly angular periphery, very narrow roundish umbilicus, spire formed by flattened whorls separated by an indented suture, radial sculpture moderately developed, and microspiral striae well marked (GITTENBERGER, 1993). Candidula olisippensis is appa- rently endemic to Portugal. The current finding represents the northern limit of its known range. Galicia. Paz Ondina, Carlos Prieto, Án- gel A. Luque, and two anonymous reviewers kindly provided helpful com- ments. This work was partially suppor- ted by Research Project PB93-0055 of DEE: KERNEY, M. P. AND CAMERON, R. A. D.,, 1979. A field guide to the land snails of Britain and north-west Europe. Collins, London, 288 pp. LOCARD, A., 1899. Conchyliologie Portugaise. Les coquilles terrestres, des eaux douces et saumátres. Archives du Muséum de Lyon, 7 (1): i-xi, 1-303. MANGA GONZALEZ, Y., 1979. Sobre las espe- cies del género Candidula Kobelt, 1871, (Gas- tropoda, Stylommatophora) en la provincia de León. Revista Ibérica de Parasitología, 79: 455- 465 + 1 pl. NOBRE, A,, 1930. Moluscos terrestres, fluviais e das aguas salobras de Portugal. Porto, 259 pp, 18 pls. NOBRE, A., 1941. Moluscos terrestres e fluviais. Fauna de Portugal, 2. Memórias e Estudos do Muséu de Zoología da Universidade da Coimbra, 124: 1-277, pls. i-iv, 1-30. SEIxAS, M., 1992. Gasterópodes terrestres da coleccao do Museu Bocage. Arquivos do Mu- séu Bocage, Nova Série, 2 (10): 155-255. SERVAIN, G., 1880. Étude sur les mollusques re- cueillis en Espagne et en Portugal. Saint Ger- main, Paris, 172 pp. O Sociedad Española de Malacología Iberus, 151% 9=-22, 1997 Nuevos datos sobre la distribución de la superfamilia Heli- coidea Rafinesque, 1815 (Gastropoda, Pulmonata, Stylom- matophora) en el oeste de Galicia New records about the distribution of the superfamily Helicoidea Rafinesque, 1815 (Gastropoda, Pulmonata, Stylommatophora) in the West of Galicia Paz ONDINA, Jesús HERMIDA y Adolfo OUTEIRO* Recibido el 21-IV-1996.Aceptado el 30-IV-1996 RESUMEN En este trabajo se realiza un estudio faunístico de las especies de la superfamilia Helicoi- dea Rafinesque, 1815, encontradas en el oeste de Galicia (provincias de A Coruña y Pontevedra). Para cada especie se incluyen citas previas y localidades de captura con las coordenadas U.T.M. 10x10 km. Teniendo en cuenta nuestros hallazgos y los de los auto- res consultados en la bibliografía se han elaborado los mapas de distribución correspon- dientes, en sistema U.T.M. de 10x10 km. Se cita por primera vez para el área de estudio Mengoana brigantina (da Silva Mengo, 1867). ABSTRACT A faunistic study of species of superfamily Helicoidea Rafinesque, 1815, in the west of Galicia (La Coruña and Pontevedra provinces) has been realized. For each species the previous records and the coordinates U.T.M. 10x10 km of the localities where the species have been found, are included. Taking our own findings into consideration, and the data from bibliography, a map showing the distribution of each species has been drawn up, using U.T.M. 10x10 km system. Mengoana brigantina [da Silva Mengo, 1867) is recor- ded for the first time in this area. PALABRAS CLAVE: Gastropoda, Pulmonata, Helicoidea, distribución, Galicia. KEY WORDS: Gastropoda, Pulmonata, Helicoidea, distribution, Galicia. INTRODUCCIÓN El conocimiento de la fauna de moluscos terrestres de la Península Ibérica presentaba, hasta no hace mucho tiempo, un gran retraso respecto a Europa. Fue a finales del siglo XIX y principios del XX cuando se desarrolló más intensamente la actividad en este campo, y cuando comienza a otorgár- sele validez taxonómica a ciertas estruc- turas anatómicas, especialmente el *Departamento de Biología Animal. Facultad de Biología. Universidad de Santiago de Compostela. 15706 Santiago de Compostela. La Coruña, España. Iberus, 15 (1), 1997 aparato genital, tras comprobar que especies con conchas muy parecidas albergaban animales de organización anatómica diferente. También en Galicia se inician, en este período, los estudios malacológicos con las aportaciones de autores como GRAELLS (1846), SEOANE (1866), MACHO VELADO (1871, 1878) e HIDALGO (1875). Posteriormente otros autores han reali- zado estudios en zonas geográficas con- cretas como ALTIMIRA (1969); SACCHI y VIOLANI (1977) y ROLÁN y OTERO (1988). Con mención específica a los helícidos cabe destacar el trabajo de CASTILLEJO (1981; 1986), que cubre gran parte del territorio gallego, recogiendo muestras en un total de 87 cuadrículas de 10x10 km U.T.M. A pesar de estos estudios existen dis- continuidades en la distribución de gran parte de las especies, motivo por el cual con el presente estudio queremos contri- buir a ampliar el conocimiento faunís- tico de los gasterópodos terrestres perte- necientes a la superfamilia Helicoidea Rafinesque, 1815, aportando nuevos datos de distribución para las provincias de A Coruña y Pontevedra. MATERIAL Y METODOS Durante el período 1986-1989 se ha recolectado material malacológico pro- cedente de las 176 cuadrículas U.T.M. de 10x10 km, en las que se han dividido las provincias de A Coruña y Pontevedra (Fig. la). En las distintas cuadrículas visitadas se han examinado detenida- mente, de día y de noche, los distintos hábitats donde suelen resguardarse, recogiéndose además, muestras de suelo y hojarasca de distintos biotopos, que posteriormente se lavaron y tamizaron para separar los ejemplares. Todos los gasterópodos capturados se sometieron al proceso habitual de muerte por anoxia sumergiéndolos en agua, para facilitar de este modo su disección, conservándose posterior- mente en alcohol de 70*. A partir de los datos obtenidos, se han elaborado los mapas de distribución de cada especie (Figs. 1b-g, Figs. 2a-1) en cuadrículas U.T.M. de 10x10 km, indi- cándose tanto las localidades aportadas en este trabajo (+) como las procedentes de la bibliografía (*), así como las locali- dades en las que se encontraron única- mente conchas vacías (O). La colección malacológica se encuen- tra depositada en el Departamento de Biología Animal (Facultad de Biología, Universidad de Santiago de Compos- tela). RESULTADOS Se han identificado un total de 6873 ejemplares pertenecientes a 18 especies. Para cada especie se incluyen los siguientes apartados: citas previas, material estudiado (en el que se indica el número de ejemplares capturados en cada cuadrícula y aquellas localidades donde se encontraron conchas vacías), un breve resumen de su distribución geográfica y algunas observaciones de interés para aquellas especies en que lo hemos considerado necesario. El listado de las localidades junto a su correspon- diente código, coordenadas U.T.M., Ayuntamiento y fecha de muestreo se pueden observar en la Tabla 1. Superfamilia HELICOIDEA Rafinesque, 1815 Familia XANTHONYCHIDAE Strebel y Pfeffer, 1880 Elona quimperiana (Férussac, 1821) (Fig. 1B) Citas previas: CAZIOT (1915) como H. quimperiana; CASTILLEJO (1986); OTERO y TRIGO (1989). Material examinado (107 ejemplares). 39: 1; 58: 1; 93: 9; 94: 3; 100: 7; 102: 1; 103: 8; 104: 12; 125: 9; 130: 1; 141: 3; 143: 2; 144: 1; 145: 5; 146: 15; 147: 14; 148: 4; 156: 6; 157: 5. ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia Tabla I. Listado de las localidades, provincia, U.T.M. en 10 x 10 Km y fecha de recogida de las muestras, junto a su correspondiente código. C: A Coruña; L: Lugo; O: Ourense; P: Pontevedra. Table I. List of the locations, provinces, U.I.M. 10x10 Km and sampling date, with its code. C: A Coruña; L: Lugo; O: Ourense; P: Pontevedra. Código O0XD0 NXNOo0hb0N — Localidad /Ayuntamiento Caldelas de Tui, Tui Cristelos, Tomiño Louredo, Mos Coruxo, Vigo Pintos, Pontevedra Dorrón, Sanxenxo A Xesta, A Lama O Covelo, O Covelo Caldas de Reis, Caldas de Reis Sobrido, Ribeira Noal, Porto do Son Sionlla, Santiago Negreira, Negreira Ponte Sarandón, Vedra Arca, A Estrada Ferreiros, Vila de Cruces Prado, Lalín Santa María, Rodeiro Amance, Golada Herbón, Padrón Budiño, Porriño Piñeiro, O Covelo Salvaterra, Salvaterra do Miño Arbo, Arbo loureza, Oia Baredo, Baiona Amoedo, Pazos de Borbén Magdalena, Cangas Leiro, Ribadumia San Vicente do Grove, O Grove Serrapio, Cerdedo Nigoi, A Estrada Vilatuxe, Lalín lagoa, Dozón Barrio, Vila de Cruces Ventosa, Golada Catoira, Catoira Boiro, Boiro Lapido, Ames Sergude, Boqueixón Costa de Mougás, Oia Camposancos, A Guarda Provincia ADA OO A A A A AS AAA A A SAO ASISTAN O O) A A a a) e) A) a) A) a) Coord. UTM 29TNG35 29TNG15 29TNG37 29TNG17 29TNG39 29TNG19 29TNG59 29TNG57 29TNH21 29TNHO1 29TNHO3 29TNHA5 29TNH25 29TNH43 29TNH41 29TNHÓ64 29TNH62 29TNH82 29TNH84 29TNH23 29TNG36 29TNG58 29TNGA45 29TNG56 29TNG14 29TNG16 29TNG38 29TNG18 29TNH20 29TNHOO 29TNH40 29TNH42 29TNH61 29TNH81 29TNH63 29TNH83 29TNH22 29TNHO2 29TNH24 29TNH44 29TNGO5 29TNG13 Fecha 26-09-86 26-09-86 27-09-86 27-09-86 28-09-86 28-09-86 29-09-86 29-09-86 30-09-86 01-10-86 01-10-86 02-10-86 02-10-86 03-10-86 03-10-86 04-10-86 04-10-86 05-10-86 05-10-86 06-10-86 04-12-86 04-12-86 05-12-86 05-12-86 06-12-86 06-12-86 07-12-86 07-12-86 08-12-86 08-12-86 09-12-86 09-12-86 10-12-86 10-12-86 11-12-86 11-12-86 12-12-86 12-12-86 13-12-86 13-12-86 10-04-87 11-04-87 Iberus, 15 (1), 1997 Tabla I. Continuación. Table I. Continuation. Código Localidad/Ayuntamiento Provincia Coord. UTM 44 Mondariz, Mondariz P 29TNG47 A5 Lavadores, Vigo P 291NG27 46 Pontecaldelas, Pontecaldelas p 29TNG49 A7 Postemirón, Vilaboa P 29TNG29 48 Triñnáns, Boiro (E 29TNH11 49 Oleiros, Ribeira S 29TMH91 50 Portobravo, Lousame a 29TNH13 51 César, Caldas de Reis P 29TNH31 2 Carcacía, Padrón E 29TNH33 53 Carballeda, Lalín P 29TNH72 54 Baíña, A Golada P 29TNH74 DS) Quintás, Touro E 29TNH54 56 Rellas, Silleda Cc 291NH52 7 Louro, Muros € 29TMH93 58 Ordoeste, A Baña E 29TNH15 DY A Peregrina, Santiago € 291NH35 60 Illas Cíes, Vigo P 29TNGO7 61 Barrantes, Ribadumia P 29TNH10 62 Fontáns, Barro Pp 29TNH30 63 Forcarei, Forcarei P 29TNH5 1 64 Lamela, Silleda P 29TNH53 65 Muimenta, Lalín P 29TNH73 66 A Xesta, Lalín P 29TNH7 1 67 Baroña, Porto do Son a 29TMH92 68 Pontenafonso, Noia € 29TNH14 69 Carreira, Ribeira E 29TMH9O 70 Bures, Catoira € 29TNH 12 71 Santa Mariña de Barcala, A Estrada. P 291TNH32 Ez Oitavén, Fornelos de Montes P 29TNG48 US Ponteareas, Ponteareas P 29TNG46 7A Moaña, Moaña P 29TNG28 75 Mosende, Porriño P 29TNG26 76 Tomiño, Tomiño P 29TNG24 7 Chavella, Oia P 29TNGO4 78 Cabo Silleiro, Baiona P 29TNGO06 79 Santiago, Santiago € 29TNH34 80 Esmorode, Santa Comba E 29TNH16 81 Serra de Outes, Serra de Outes E 29TNHOA4 82 Quilmas, Carnota e 291MH84 83 Vilela, Muxía E 29TMH76 84 Berdoias, Vimianzo e 29TMH96 85 Nande, Laxe € 29TMH98 86 As Tarandeiras, Coristanco E 29TNH18 Fecha 12-04-87 12-04-87 13-04-87 13-04-87 14-04-87 15-04-87 15-04-87 16-04-87 16-04-87 17-04-87 17-04-87 18-04-87 18-04-87 19-04-87 20-04-87 20-04-87 12-05-87 27-06-87 27-06-87 28-06-87 28-06-87 29-06-87 29-06-87 01-07-87 01-07-87 02-07-87 03-07-87 03-07-87 04-07-87 04-07-87 05-07-87 06-07-87 06-07-87 07-07-87 07-07-87 08-07-87 09-10-87 09-10-87 10-10-87 10-10-87 11-10-87 11-10-87 12-10-87 ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia Tabla I. Continuación. Table I. Continuation. Código Localidad/Ayuntamiento 88 Rial, Trazo 89 Zas de Reis, Melide 90 Padreiro, Curtis 91 Xestal, Irixoa 92 Saa, As Pontes 93 Freires, Ortigueira 94 Regoa, Cedeira 95 Esmelle, Ferrol 96 Bouzarredonda, Neda 97 Sigrás, Cambre 98 Abellá, Frades 99 Pedrouzo, O Pino 100 Viñas, Paderne 101 Neda, Neda 102 Ponte de Mera, Ortigueira 103 Mañón, Mañón 104 Recemel, Somozas 105 Boliqueiras, As Pontes 106 Salto do Conexo, Monfero 107 Monte do Arco, Curtis 108 Rexidoira, Cesuras 109 Arzúa, Arzúa 110 Furelos, Melide ME Vila da Igrexa, Cerceda 112 Parada, Ordes 118 Pedrafigueira, Carnota 114 Cabo Fisterra, Fisterra MS Ozón, Muxía 116 Areosa, Vimianzo IN Ponteceso, Ponteceso 118 Cances, Carballo 119 Bembibre, Val do Dubra 120 Sobrado dos Monxes, Sobrado 121 A Castellana, Aranga 1122 A Capela, A Capela 123 San Sadurniño, San Sadurniño 124 O Ermo, Ortigueira 125 Sismundi, Ortigueira 126 Raxón, Ferrol 127 Mabegondo, Abegondo 128 Carnoedo, Sada 129 Poulo, Ordes 130 Río, Cerceda Provincia DKOOOOOOoOS0OoOO0O0o0o00 0000 O0O0oO0O0o0 0000000000000 000000O00O Coord. UTM 29TNH36 29TNH75 291NH77 291NH79 29TNJ91 29T1NJ93 29 T1NJ73 29TNJ51 29 1INJ7 1 29TNH59 29TNH57 29TNH55 29TNHÓ9 29TNJÓ 1 29T1NJ83 29TPJO3 29TNJ8 1 29TNJ90 29TNH89 291NH87 29TNH67 29TNHÓ5 29TNH85 29TNH48 29TNH46 291MH94 291MH74 291MH86 29TNHO6 29TNHO8 291NH28 29TNH26 29TNH7Ó 291NH78 29T1NJ7O 29TNJ72 29TNJ92 291NJ94 29TNJ52 29TNH58 29TNJ50 291NH5Ó 29TNH37 Fecha 13-10-87 19-10-87 19-10-87 20-10-87 20-10-87 21-10-87 21-10-87 22-10-87 22-10-87 23-10-87 23-10-87 24-10-87 18-01-88 18-01-88 19-01-88 19-01-88 20-01-88 20-01-88 21-01-88 22-01-88 22-01-88 23-01-88 23-01-88 24-01-88 24-01-88 26-01-88 26-01-88 27-01-88 27-01-88 28-01-88 28-01-88 29-01-88 25-03-88 25-03-88 26-03-88 26-03-88 27-03-88 28-03-88 29-03-88 30-03-88 30-03-88 31-03-88 01-04-88 Iberus, 15 (1), 1997 Tabla I. Continuación. Table I. Continuation. Código 132 188 134 139 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 159 156 157% 158 159 160 161 162 163 164 165 166 167 168 169 170 171 172 17 174 173 Localidad/Ayuntamiento Provincia Agualada, Coristanco Buño, Malpica Santa Mariña, Camariñas Ponte do Porto, Camariñas Fisterra, Fisterra Logoso, Dumbría Arceo, Boimorto Lagoa de Sobrado, Sobrado Aranga, Aranga Burricios, Oza dos Ríos Faeira, As Pontes As Somozas, Somozas Grañas, Mañón Loiba, Ortigueira Serra da Capelada, Ortigueira Sedes, Narón laraxe, Cabanas ledoño, Culleredo Leira, Ordes Razo, Carballo Canosa, Carballo Corme, Ponteceso Baio, Zas Moraime, Muxía lobelos, Cée A Picota, Mazaricos Setados, As Neves Crecente, Crecente Paredes, A Cañiza Pardesoa, Forcarei Lira, Carnota Couto, Rodeiro Vilafrío, Rodeiro San Pedro de Visma, A Coruña Sucadío, As Pontes Punta Candelaria, Cedeira Illa de Ons, Bueu Momán, Xermade Vilaverde, Ribadavia Codesas, Forcarei Cabo da Voutra, Muxía Laxe, Chantada Punta Frouxeira, Valdoviño Illa de Ons (Sur), Bueu UR OAOR LO O AOS LIO ADO OOoOOO.OOOOOOOOA Coord. UTM 29TNH17 29TNH19 291MH88 291MH97 29TMH75 29TMH95 29TNHÓ66 291NH8ó 29TNH88 29TNHÓ8 291NJ80 291NJ82 29TPJO2 29TPJO4 29T1NJ84 29TNJÓ2 29TNJ6O 291NH49 291NH47 291NH29 29TNH27 29TNHO9 29TNHO7 291MH87 291MH85 29TNHO5 29TNG55 29TNG66 291NGÓ67 29TNH50 291MH83 29TNH93 291NH92 29TNJ40 29TPJO1 29T1NJ74 29TNGO9 291NH99 29TNG68 29TNHÓO 291MH77 29TNH91 29TNJÓ3 29TNGO9 10 Fecha 02-04-88 02-04-88 03-04-88 03-04-88 04-04-88 05-04-88 30-06-88 30-06-88 01-07-88 01-07-88 02-07-88 02-07-88 03-07-88 03-07-88 04-07-88 05-07-88 05-07-88 06-07-88 06-07-88 07-07-88 07-07-88 08-07-88 08-07-88 09-07-88 09-07-88 10-07-88 17-03-89 17-03-89 18-03-89 19-03-89 22-04-89 23-05-89 23-05-89 30-05-89 08-06-89 28-03-88 31-05-89 08-06-89 18-03-89 19-03-89 22-04-89 23-05-89 10-05-89 31-05-89 ONDINA £7 AL.: Distribución de la superfamilia Helicoidea en el oeste de Galicia Distribución geográfica: Se trata de una especie de distribución atlántico-eu- ropea (KERNEY, CAMERON Y JUNGBLUTH, 1983). Fuera de la Península Ibérica su área de distribución se restringe a la Bre-. taña francesa y al País Vasco francés (GERMAIN, 1930). En la Península se limita a la costa sep- tentrional. Está citada en el norte desde Galicia hasta el País Vasco (HIDALGO, 1875; GITTENBERGER, 1979; LARRAZ y JORDANA, 1984; CASTILLEJO, 1986; HERMIDA, OUTEIRO Y RODRÍGUEZ, 1992; PUENTE y PRIETO, 1992; LARRAZ y EQUISOAIN, 1993). Familia HYGROMIIDAE Tryon, 1866 Candidula intersecta (Poiret, 1801) (Fig. 1C) Citas previas: MACHO VELADO (1871) como H. caperata; HIDALGO (1875, 1890) como H. caperata; SACCHI y VIOLANI (1977) como Helicella caperata; CASTILLEJO (1986). Material examinado (93 ejemplares). 6: 1; 11: 3; 19: 1; 20: 1; 30: 1; 35: 10; 37: 1; 38: 1; 42: 3; 46: 5; 47: 2; 48: 4; 49: 6; 57: 1; 60: 6; 69: 3; 74: 3; 77: 2; 78: 5; 79: 4; 82: 1; 83: 8; 93: 6; 113: 3; 114: 4; 136: 2; 137: 1; 151: 3; 156: 2; 171: 1. Localidades con conchas vacías: 153. Distribución geográfica: Su área de distribución se enmarca en el oeste de Europa (KERNEY ET AL., 1983). En la Península se extiende por dos áreas, una en el sector norte oriental, el País Vasco y Navarra, y la otra en el tercio oeste, desde Galicia y centro de León hasta el Algarve, con algunas citas hacia el centro peninsular como Segovia (HIDALGO, 1875; NOBRE, 1941; RAMOS y APARICIO, 1985a; (CASTILLEJO, 1986; HERMIDA ET AL., 1992). En Asturias, desde la cita de ALTIMIRA (1969), no ha vuelto a ser encontrada (OJEA y ANADÓN, 1983; HERMIDA ET AL., 1992), y tampoco se tiene constancia de ella en Cantabria. Cernuella (Cernuella) virgata (da Costa, 1778) (Fig. 1D) Citas previas: SACCHI y VIOLANI (1977); CASTILLEJO (1986). Material examinado (3 ejemplares). 6: 3. Distribución geográfica: Especie mediterránea, ampliamente distribuida por el sur y oeste de Europa (KERNEY ET AL., 1983). Siendo común en el área peninsular y en las islas Baleares, es menos frecuente en la franja norte que comprende desde el valle del Tajo hasta el País Vasco, donde se convierte en una especie litoral (NOBRE, 1941; ORTIZ DE ZÁRATE y ORTIZ DE ZARATE, 1949; Seixas, 1976; PAUL, 1982; MANGA, 1983; LARRAZ y JORDANA, 1984; RAMOS y APARICIO, 1985b; CASTI- LLEJO, 1986; APARICIO, 1986; ROBLES, ISSO PEUENTESyARRIELO Ugg L9O92: HERMIDA ET AL., 1992; LARRAZ y EQUISO- AIN, 1993; ALTONAGA, GÓMEZ, MARTÍN, PRIETO, PUENTE Y RALLO, 1994). Helicella (Helicella) itala (Linneo, 1758) (Fig. 1E) Citas previas: MACHO VELADO (1871) como H. ericetorum; HIDALGO (1875) como H. ericetorum; CASTILLEJO (1986); OTERO y TRIGO (1989). Material examinado (32 ejemplares). 134: 10; 151: 20; 173: 1; 174: 1. Distribución geográfica: Se trata de una especie distribuida por el oeste de Europa (KERNEY ET AL., 1983). En la Península Ibérica está presente en la mitad norte, siendo sus citas más es- casas en Galicia (HIDALGO, 1875; HAAS, 11 Maira IMA 1929; ORTIZ DE ZÁRATE y ORTIZ DE ZÁ- RATE, 1949; MANGA, 1983, APARICIO, 1986; CASTILLEJO, 1986; HERMIDA ET AL., 1992; LARRAZ y EQUISOAIN, 1993; ALTONAGA ET AL., 1994). En el área de estudio la hemos hallado únicamente en zonas del litoral. Xerotricha apicina (Lamarck, 1822) (Fig. 1F) Citas previas: MACHO VELADO (1871) como H. apicina; HIDALGO (1875, 1890) como H. apicina; ALTIMIRA (1969); SACCHI y VIOLANI (1977) como Helicella apicina; CASTILLEJO (1986). Material examinado (3 ejemplares). 26: 3. Distribución geográfica: Es una especie con un área de distribución mediterránea (KERNEY ET AL., 1983). En la Península presenta una distri- bución costera bastante fragmentada, encontrándose en el norte unicamente en algunas localidades del litoral de Galicia (ALTIMIRA, 1969; SACCHI y VIOLANI, 1977; CASTILLEJO, 1986), Can- tabria y Asturias (PUENTE y PRIETO, 1992). En el litoral mediterráneo está citada en Cataluña (Haas, 1929; ALTI- MIRA, 1969), Valencia (ROBLES, 1990); Málaga (ALTONAGA ET AL., 1994), conti- nuando desde ahí, de una forma más regular, por el cuadrante suroccidental hasta Aveiro en Portugal (SERVAIN, 1880; LOCARD, 1899; HIDALGO, 1875; NOBRE, 1941; RAMOS y APARICIO, 1985a) y hacia el interior hasta Badajoz. GASULL (1965) también la cita abun- dantemente en las Islas Baleares. Cochlicella acuta (Muller, 1774) (Fig. 1G) Citas previas: MACHO VELADO (1871) como Bulimus acutus; HIDALGO (1875) como B. acutus; HIDALGO (1890) como H. acuta; ALTIMIRA (1969); SACCHI y VIOLANI (1977); CASTILLEJO (1986); OTERO y TRIGO (1989). Material examinado (99 ejemplares). 4: 8; 6: 8; 11: 1; 26: 9; 49: 3; 57: 4; 60: 2; 69: 4; 82: 3; 83: 8; 85: 12; 93: 5; 113: 6; 136: 10; 151: 6; 156: 4; 168: 3; 173: 3. Distribución geográfica: Distribuida por la zona mediterránea y atlántica de Europa (BOATO, BODON Y GIUSTI, 1982). Aparece practicamente en toda la costa ibérica e Islas Baleares (NOBRE, 1941; ORTIZ DE ZARATE y ORTIZ DE BEcH, 1990; MARTÍNEZ-ORTÍ, MARTÍNEZ- LÓPEZ, ROBLES Y RODRÍGUEZ BABÍO, 1990) penetrando en ocasiones hacia el interior siguiendo los valles fluviales. Está citada en puntos del interior como Salamanca (HERMIDA ET AL., 1992), ZARATE, 1949; SACCcHI, 1954; GASULL, 1965; ALTIMIRA, 1969; CASTILLEJO, 1986; Huesca y Zaragoza (PUENTE y PRIETO, 1992). Cochlicella barbara (Linneo, 1758) (Fig. 2A) Citas previas: MACHO VELADO (1871) como B. ventrosus; HIDALGO (1875) como B. ventrosus; ALTI- MIRA (1969) como C. ventricosa; SACCHI y VIOLANI (1977) como C. ventricosa; CASTILLEJO (1986) como C. ventricosa; OTERO y TRIGO (1989) como C. ventricosa. Material examinado (704 ejemplares). 1: 3; 2: 1; 3: 2; 4: 6; 5: 4; 6: 17; 9: 16; 11: 12; 21: 1; 24: 3; 25: 7; 26: 218; 27: 5; 29: 2; 30: 1; 37: 1; 38: 6; 43: 2; 44: 1; 47: 8; 48: 20; 49: 82; 50: 1; 51: 1; 52: 4; 57: 8; 60: 4; 61: 5; 68: 1; 69: 73; 70: 2; 73: 15; 74: 12; 76: 16; 81: 3; 82: 3; 83: 2; 85: 8; 86: 2; 93: 6; 94: 17; 95: 1; 97: 22; 113: 19; 117: 1; 118: 2; 126: 1; 128: 1; 132: 6; 133: 7; 135: 1; 136: 6; 148: 5; 149: 7; 151: 6; 156: 4; 168: 12; 174: 2. Localidades con conchas vacías: 127. Distribución geográfica: Especie con un amplio rango de distribución en el área mediterránea y a lo largo de las costas del oeste de Europa (BACKHUYS, 1975). 112 ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia 2 Es ll 16 | ana Ñ aan OÍ Ñ ru he) Figura 1. A: Area de estudio en la Península Ibérica. B-G: Mapas de distribución. B: Elona quimpe- riana, C: Candidula intersecta, D: Cernuella virgata; E: Helicella itala, E: Xerotricha apicina; G: Cochlicella acuta. Localidades citadas en este trabajo (+); procedentes de la bibliografía (*); única- mente encontradas conchas vacías (O). Figure 1: A: The study area in the Iberian Peninsula. B-G: Distribution maps. B: Elona quimperiana; C: Candidula intersecta; D: Cernuella virgata; £: Helicella itala; F: Xerotricha apicina; G: Cochli- cella acuta. Localities reported in this paper (+); bibliographic records (*); only shells found (O). 13 Iberus, 15 (1), 1997 En la Península presenta una distri- bución bastante amplia, y aunque aparece con mayor frecuencia en el litoral, citándose en casi todas las pro- vincias estudiadas (BOFILL y HAAS, 1920a; NOBRE, 1941; ALTIMIRA, 1969; OJEA y ANADÓN, 1983; RAMOS y APARI- CIO, 1985b, CASTILLEJO, 1986; MARTÍNEZ- ORTÍ ET AL., 1990; PUENTE y PRIETO, 1991, 1992; HERMIDA ET AL., 1992; LARRAZ y EQUISOAIN, 1993; PAREJO ET AL., 1993b). También se ha encontrado en las islas Baleares (GASULL, 1965; PAUL, 1982). En la zona de estudio está ligada a la franja litoral, aunque penetra más hacia el inte- rior que C. acuta y C. conotdea. Cochlicella conoidea (Draparnaud, 1801) (Fig. 2B) Citas previas: MACHO VELADO (1871) como B. pring1; HIDALGO (1875) como B. pring1; HIDALGO (1890) como H. conoidea; SACCHI y VIOLANI (1977); OTERO y TRIGO (1989). Material examinado (79 ejemplares). 4: 9; 6: 8; 49: 3; 57: 4; 60: 1; 82: 4, 83: 20; 113: 4; 136: 20; 156: 6. Distribución geográfica: Especie común a lo largo de la costa mediterrá- nea, desde los Pirineos orientales hasta los Alpes marítimos (KERNEY ET AL., 1983) En la Península se encuentra en el li- toral mediterráneo, aunque también está citada en la costa atlántica portuguesa y gallega, y aisladamente y con escasos ejemplares en el interior (LOCARD, 1899; HAas, 1929; NOBRE, 1941; SaccHi, 1954; GASULL, 1965; ALTIMIRA, 1969; GASULL, 1975; SACCHI y VIOLANI, 1977; RAMOS y APARICIO, 1985b; CASTILLEJO, 1986; MAR- TÍNEZ-ORTÍ ET AL., 1990). La única cita existente en el litoral cantábrico dada por ORTIZ DE ZARATE y ORTIZ DE ZÁ- RATE (1949) ha sido asignada por distin- tos autores a C. barbara (PUENTE y PRIETO, 1992). Ashfordia granulata (Alder, 1830) (Fig. 2C) Citas previas: MACHO VELADO (1871) como H. sericea; HIDALGO (1875) como H. sericea; CASTI- LLEJO (1986) como Monacha (Ashfordia) granulata. Material examinado (151 ejemplares). 6: 1; 35: 6; 53: 1; 69: 3; 79: 24; 83: 1; 90: 5; 93: 5; 94: 23; 95: 13; 96: 1; 97: 14; 102: 11; 115: 2; 117: 3; 118: 1; 120: 3; 135: 22; 143: 1; 145: 1; 149: 2; 153: 2; 155: 3; 165: 3. Distribución geográfica: Se distri- buye por el oeste de Europa (KERNEY ET AL., 1983). En la Península Ibérica su área de distribución comprende desde Galicia hasta el extremo occidental de Vizcaya, penetrando hacia León (MANGA, 1983; ANADÓN y OJEA, 1984; HOLYOAK y SED- DON, 1985; CASTILLEJO, 1986; HERMIDA ET AL., 1992; PUENTE y PRIETO, 1992). Zenobiella subrufescens (Miller, 1822) (Fig. 2D) Citas previas: CASTILLEJO (1986) como Monacha (Zenobiella) subrufescens. Material examinado (34 ejemplares). 39: 1; 40: 1; 50: 4; 68: 1; 79: 1; 87: 5; 90: 2; 93: 3; 94: 2; 97: 1; 102: 2; 108: 1; 111: 1; 116: 1; 117: 5; 118: 1; 128: 1; 162: 1. Localidades con conchas vacías: 13, 58. Distribución geográfica: Se extiende por el oeste europeo (KERNEY ET AL., 1983). Según GERMAIN (1930) no se aleja de las regiones de influencia marítima. En la Península Ibérica está citada unicamente en la franja norte que com- 14 prende desde Galicia hasta el País Vasco y Navarra (OJEAa y ANADÓN, 1983; CASTILLEJO, 1986, HERMIDA ET AL., 1992; PUENTE y PRIETO, 1992; LARRAZ y EQUISOAIN, 1993; ALTONAGA ET AL., 1994). ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia Portugala inchoata (Morelet, 1845) (Fig. 2E) Citas previas: MACHO VELADO (1871) como H. inchoata; HIDALGO (1875, 1890) como H. inchoata; GITTENBERGER (1980); SAccHI (1981) como Monachoides inchoatus; CASTILLEJO (1986); OTERO y TRIGO (1989). Material examinado (928 ejemplares). 1: 11; 2: 17; 3: 13: 1; 14: 2; 16: 16; 17: 3; 18: 2; 19: 5; 20: 4; 21: 19; 22: 30: 3; 31: 4; 32: 3; 33: 1; 35: 10; 36: 3; 37: 1; 38: 8; 39: 16; 47: 9; 48: 17; 49: 3; 50: 12; 51: 1; 52: 8; 53: 2; do 6; 55: 5; 63: 1; 64: 6; 65: 5; 67: 8; 68: 4; 69: 6; 70: 9; 71: 2; 72: 5; 73: 89: 1 ;4: 7; 5: 7; 6: 20; 8: 5; 9: 11; 10: 2; 11: 7; 12: 8; o : 4; 41: 14; 42: 12; 43: 5; 44: 4; 45: 7; 46: 7; : 4; 57: 3; 58: 2; 59: 2; 60: 21; 61: 10; 62: 4; 71: 4: e 75: 17; 76: 15; 77: 7; 78: 5; 79: 9; 80: 10; 81: 2; 82: 3; 83: 10; 84: 2; 85: 5; 86: 14; 87: 1; 88: 1; 1: 2; 93: 4; 94: 5; 95: 8; 96: 2; 97: 9; 98: 1; 99: 6; 100: 5; 101: 7; 103: 1; 104: 2; 105: 1; 106: : 1; 109: 3; 110: 4; 111: 11; 112: 15; 113: 3; 114: 3; 115: 5; 116: 4; 117: 12; 118: 7; 119: 3; 121: ; 124: 1; 125: 13; 127: 1; 128: 4; 129: 1; 130: 2; 131: 4; 132: 2; 133: 4; 135: 4; 136: 21; 137: 8; 138: 2; 139: 2: 140: 1; 141: 2; 142: 4; 143: 7; 145: 6; 146: 5; 147: 2; 148: 3; 149: 4; 150: 2; 151: 8; 152: 5; 153: 13; 154: 1; 155: 8; 156: 8; 157: 4; 158: 5; 160: 1; 161: 2; 162: 4; 164: 2; 165: 2; 167: 2; 168: 3; 170: 1; 171: 8; 172: 2. Localidades con conchas vacías: 15. , Distribución geográfica: Esta especie es endémica del oeste de la Península Ibérica (NOBRE, 1941; GITTENBERGER, 1980). Está citada en el norte, centro y sur de Portugal por diversos autores (MORE- LET, 1877; SERVAIN, 1880; LOCARD, 1899; NOBRE, 1941; RAMOS y APARICIO, 1985a) y en toda la franja oeste española desde Asturias a Huelva (MANGA, 1983; HER- MIDA ET AL., 1992; PUENTE y PRIETO, 1992). En Galicia es uno de los elementos más característicos de su malacofauna. Ponentina subvirescens (Bellamy, 1839) (Fig. 2F) Citas previas: MACHO VELADO (1871) como H. occidentalis; HIDALGO (1875) como H. occidentalis; SACCHI y VIOLANI (1977) como Trichia occidentalis; CASTILLEJO (1986) como Ponentina ponentina. Material examinado (321 ejemplares). 1: 2; 2: 2; 4: 1; 19: 3; 21: 5; 26: 6; 30: 1; 32: 1; 36: 3; 38: 1; 41: 4; 42: 14; 43: 10; 44: 3; 45: 2; 47: 7; 48: 3; 49: 4; 50: 10; 51: 1; 52: 8; 54: 2; 57: 5; 59: 1; 60: 6; 61: 1; 70: 1; 76: 2; 77: 2; 79: 1; 86: 1; 89: 1; 91: 2; 97: 2; 99: 115: 2; 117: 1; 119: 119; 122: 7; 132: 3; 13 Localidades con conchas vacías: 3; 6; 131; 142; 148; 151; 159; 160. 3 9: Distribución geográfica: Especie atlántica que se distribuye por el suroeste de Europa (KERNEY ET AL., 1983), pertene- ciente a la fauna atlantico-lusitánica. En la Península se distribuye a lo largo de todo su tercio oeste (LOCARD, 5; 103: 3; 107: 2; 109: 1; 110: 1; 111: 3; 112: 1; 113: 1; 114: 9; :2; 134: 22; 135: 1; 137: 2; 138: 1; 146: 7; 153: 8; 169: 2. 22; 23; 24; 25; 28; 29; 34; 39; 53; 73; 83; 88; 120; 121; 127; 1899; NOBRE, 1941; MANGA, 1983; Cas- TILLEJO, 1986; HERMIDA ET AL., 1992; PUENTE y PRIETO, 1992). Los puntos más orientales corresponden a La Rioja, Burgos y Alava (PUENTE y PRIETO, 1992). Mengoana brigantina (da Silva Mengo, 1867) (Fig. 2G) Material examinado (3 ejemplares). 9: 1; 67: 1; 128: 1. Distribución geográfica: Esta espe- cie es un endemismo ibérico restringido al noroeste de la Península. Su área de distribución se extiende desde el extremo nororiental de Portu- gal, hasta el oeste de Vizcaya (HIDALGO, 1875; ORTIZ DE ZARATE, 1949; RAMOS y APARICIO, 1985a; MANGA, 1983; HER- MIDA ET AL., 1992; PUENTE y PRIETO, 1992). Hasta este momento su límite oc- cidental se encontraba en el este de Gali- cia (CASTILLEJO, 1986). 15 Iberus, 15 (1), 1997 Observaciones: Se trata de la cita más occidental de las conocidas y de la primera para el área de estudio, encon- trándose en zonas costeras. Su carácter calcícola puede ser la causa de que hayamos recogido un escaso número de ejemplares en nuestra área de estudio, predominantemente granítica. Este taxon presenta una cierta contro- versia dado que el holotipo de esta espe- cie, citada como Helix brigantina (da Silva Mengo, 1867) y recogido en Braganca (Portugal), se ha extraviado, conserván- dose únicamente su descripción, sin nin- gún dibujo o representanción y sin que haya vuelto a encontrarse ningún ejem- plar en la localidad tipo. ROSSMASSLER (1879) figura un ejemplar de esta especie, aportando otros datos a su descripción y señalando una nueva localidad en el norte de la Península (La Liébana, Can- tabria). NOBRE (1941) no encuentra esta especie en ningún punto de Portugal y presupone que la especie descrita por DA SILVA MENGO (1867) podría tratarse de una variedad minor de Helix inchoata (Morelet, 1879). Este problema persiste hoy en día y sigue siendo necesario com- probar si el genital de los individuos en- contrados en los alrededores de Bra- ganca coincide con el representado por ORTIZ DE ZARATE (1949). De no ser así, lo que ahora se conoce como Mengoana bri- gantina en el norte de la Península debe- ría ser considerada como Mengoana jes- chaul (ORTIZ DE ZARATE, 1949). Oestophora (Oestophora) barbula (Rossmássler, 1838) (Fig. 2h) Citas previas: GRAELLS (1846) como H. holosericea; MACHO VELADO (1871) como H. barbula; HIDALGO (1875, 1890) como H. barbula; ORTIZ DE ZARATE (1962); SACCHI y VIOLANI (1977); CASTI- LLEJO (1984); OTERO y TRIGO (1989). Material examinado (661 ejemplares). 1: 10; 2: 4; 3: 7; 4: 8; 5: 2; 2; 19: 2; 20: 4; 21: 2; 22: 2; 23: 4; 24: 5; 25: 10; 26: 14; 27: 6 15; 41: 17; 42: 26; 43: 7; 45: 2; 46: 3; 47: 1; 48: 11; 49: 60: 7; 61: 1; 62: 1; 67: 2; 68: 4; 69: 33; 70: 1; 74: 2; 77: 6; 88: 2; 89: 4; 94: 2; 95: 2; 96: 1; 97: 8; 98: 2; 99: 3; 116: 2; 117: 8; 118: 3; 119: 5; 120: 6; 122: 1; 123: 4; 127: 8; 1 136: 7; 137: 4; 138: 8; 139: 1; 143: 1; 145: 2; 149: 10; 150: 1; 151: 157: 1; 158: 8; 161: 6; 168: 7; 171: 14. Localidades con conchas vacías: 15. Distribución geográfica: Esta espe- cie es un endemismo ibérico pertene- ciente a la fauna atlántico lusitánica. Está presente en todo el oeste penin- sular, desde Galicia y Asturias hasta el sur de Portugal y Huelva, adentrándose en algunos puntos del centro peninsular ; 6: 3;7: 1; 9: 2; 10: 1; 11: 1; 14: 7; 17: ; 28: 9; 29: 20; 30: 7; 32: 2; 35: 10; 37: 15; 39: ; 51: 6;52: 5;53: 1; 54: 1; 56: 2; 57: 6; 58: 8; 59: 3; ;79: de 81: 4; 82: 2; 83: 10; 84: 3; 85: 1; 86: 14; 87: : 4; 109: 6; 110: 7; 112: 1; 113: 23; 114: 17; 115: 4; sl 5; 129: 5; 130: 1; 132: 2; 134: 1; 135: 34; 3; 152: 8; 153: 15; 154: 2; 155: 7; 156: 8; (NOBRE, 1941; ORTIZ DE ZÁRATE, 1962; ALTIMIRA, 1969; MANGA, 1983; RAMOS y APARICIO, 1985a; CASTILLEJO, 1984; HERMIDA ET AL., 1992). Alejada de este área se ha citado también en sureste ibérico (HIDALGO, 1875; GASULL, 1975) y Pirineos. Oestophora (Oestophora) silvae Ortiz de Zárate López, 1962 (Fig. 21) Citas previas: HIDALGO (1875, 1890) como O. lusitanica; ORTIZ DE ZARATE (1962); CASTILLEJO (1984) como O. lusitanica var. minor; OTERO y TRIGO (1989). Material examinado (474 ejemplares). 1: 16; 3: 1; 4: 2; 5: 5; 7: 3; 8: 2; 13: 1; 1 o ec ads 35: 4 8; 43: 10; 44: 7; 45: 2; 46: 9; 47: 9; 50: 13; 51: 3; 53: 15; 54: 2; 77: 7; 82: 1; 84: 3; 88: 1; 90: 2; 91: 2; 99: 18; 100: 10; 106: 3; 121: 16; 130: 1; 137: 4; 138: 1; 141: 6; 154: 1; 155: 1; 158: 1; Localidades con conchas vacías: 16; 87; 93. 16 4: 4; 18: 2; 19: 1; 21: 8; 3: 13; ; 37:29; 39: 7; 40: 34; 41: 39; 42: 55: 2; 56: 1; 58: 5; 62: 1; 63: 3; 71: 2; 75: 1; 108: 7; 109: 7; 110: 10; 111: 1; 115: 9; 120: 1; 159: 6; 160: 3; 166: 2; 170: 4. ONDINA £7 AZz.: Distribución de la superfamilia Helicoidea en el oeste de Galicia ¡cleje! fejele| ¡ojoje| ¡[eloje| | 300 AJ000220:1 “GOPsog00bn ”O0D900095 ¡mole olela!_| ¡eo ¡qloeje! || ge: ECU [ol6je] E ¡des ¡Je NW POJE PABuBI ¡elejel AE NA A OSEA E a EE ?CÍGUOGA EAS ae ES CENA se |ejejejel ¡ejojojeje] ! lol eje] | Tele] e ¡ejojele] ejeje] je xjejalolelolal O ejejejaja! ABRAN PEOR 230 pt] Da m8 AÑO 107 O”SgO 1 Oe AQDOBOO cOn 95 jojo|ojo 4 y ea Ira J: AS 2 DO: el 0003 AS DECO 5 ¡588 jé1 7] Sl 13439000 eló| [ojala K L [ojejejsíate] Figura 2. Mapas de distribución. A: Cochicella barbara; B: Cochlicella conoidea; C: Ashfordia granu- lata; D: Zenobiella subrufescens, E: Portugala inchoata, E: Ponentina subvirescens, G: Mengoana bri- gantina; H: Oestophora barbula; 1: Oestophora silvae, J: Theba pisana; K: Cepaea nemoralis, L: Helix aspersa. Localidades citadas en este trabajo (*); procedentes de la bibliografía (*); únicamente encontradas conchas vacías (O). Figure 2: Distribution maps. A: Cochlicella barbara; B: Cochlicella conoidea; C: Ashfordia granu- lata; D: Zenobiella subrufescens; E: Portugala inchoata; F: Ponentina subvirescens; G: Mengoana brigantina; A: Oestophora barbula; /: Oestophora silvae; /: Theba pisana; K: Cepaea nemoralis; L: Helix aspersa. Localities reported in this paper (e); bibliographic records (*); only shells found (O). 17 Iberus, 15 (1), 1997 Distribución geográfica: Es un en- demismo ibérico restringido al norte de la Península, desde Galicia hasta el norte de Alava (CASTILLEJO, 1986; OJEA y ANADÓN, 1983; PUENTE y PRIETO, 1992; HERMIDA, ET AL., 1992). Observaciones: ORTIZ DE ZARATE (1962) al describir O. silvae expuso las dife- rencias que existían entre esta especie y Oestophora lusitanica (Pfeiffer, 1841). Señala que O. silvae es más pequeña, tiene el borde superior de la abertura más corto, un engrosamiento mayor del peristoma en toda su extensión, presenta una callosi- dad blanca interna en el extremo del borde superior de la última vuelta con la ante- penúltima y carece del estriado espiral en la cara inferior, cerca de la abertura, que caracteriza a O. lusitanica. Respecto al aparato genital destaca la distinta inserción del músculo retractor del pene, que se sitúa hacia la mitad de la vaina en O. lusi- tanica y cerca del extremo en O. silvae, y la distinta longitud de las glándulas multí- fidas y del oviducto libre, mucho más cortos en O. silvae. Según ORTIZ DE ZÁRATE (1962) O. silvae es la misma especie que fue descrita por da Silva en 1871 como O. lusitanica var. minor. Así mismo asigna a esta especie todas las citas de O. lusitanica, dadas por HIDALGO (1875), para el norte de la Penín- sula y Galicia. Sin embargo CASTILLEJO (1984) encontró, en diversos puntos de Galicia, ejemplares que no coincidían com- pletamente con la descripción de O. silvae, siendo más similar a la de O. lusitanica var. minor, a la que asignó esos ejemplares. Nosotros hemos podido estudiar indi- viduos de O. lusitanica de Portugal, y nin- guno de nuestros ejemplares gallegos se ajusta a las características de esta especie. Respecto a O. silvae, hemos observado que los datos de la descripción de ORTIZ DE ZARATE (1962) deberían ser revisados, ya que existen ejemplares de mayor diá- metro que el señalado por él, y una obser- vación detallada de la concha húmeda pone de manifiesto, en todos los ejempla- res, las estrías espirales en la cara inferior que, en un principio, sólo parecía presen- tar O. lusitanica. En lo referente al aparato genital, las glándulas multífidas pueden ser más largas que las descritas y el mús- culo retractor del pene presenta cierta variabilidad en su inserción. Observadas estas variaciones y una vez estudiados los ejemplares de nuestra colección y la del Dr. Castillejo, hemos seguido a Ortiz de Zárate considerando las citas gallegas de O. lusitanica var. minor como O. silvae. Familia HELICIDAE Rafinesque, 1815 Theba pisana (Muller, 1774) (Fig. 2) Citas previas: MACHO VELADO (1871) como H. pisana; HIDALGO (1875, 1890) como H. pisana; SACCHI y VIOLANI (1977) como Euparipha pisana; CASTILLEJO (1986); OTERO y TRIGO (1989). Material examinado (373 ejemplares). 4: 20; 6: 3; 11: 3; 26: 10; 30: 2; 48: 1; 49: 1; 57: 14; 60: 2; 67: 5; 69: 50; 82: 1; 83: 4; 85: 12; 86: 5; 87: 10; 91: 10; 93: 2; 94: 17; 95: 1; 97: 3; 100: 3; 113: 7; 117: 1; 118: 12; 126: 9; 128: 1; 131: 33; 133: 12; 136: 15; 149: 35; 151: 25; 153: 10; 156: 3; 165: 2; 168: 5; 173: 24. Distribución geográfica: Esta especie presenta una distribución circunmedite- rránea que remonta las costas atlánticas hasta Inglaterra (KERNEY ET AL., 1983). En la Península se extiende a lo largo de toda la costa, aunque es capaz de colo- nizar áreas del interior cuando se dan con- diciones adecuadas, tales como disponi- bilidad de sales solubles (Prieto, com. pers.). Como citas más recientes en el interior se pueden señalar las dadas por HERMIDA ET AL. (1992) y PAREJO ET AL. (1993b). Cepaea (Cepaea) nemoralis (Linneo, 1758) (Fig. 2K) Citas previas: MACHO VELADO (1871) como H. nemoralis; HIDALGO (1875, 1890) como H. nemoralis; SACCHI y VIOLANI (1977); SACCHI (1981); CASTILLEJO (1986); OTERO y TRIGO (1989). * 18 ONDINA £7 4Z.: Distribución de la superfamilia Helicoidea en el oeste de Galicia Material examinado (1229 ejemplares). 1: 32; 2: 13; 3: 10; 4: 7; 5: 3; 6: 11;7: 2; 8: 4; 9: 4,10: 1; 11: 4; 12: 4; 13: 4; 14: 7; 16: 8; 17: 3; 18: 4; 19: 4; 20: 2; 21: 5; 22: 9; 23: 3; 24: 9; 25: 17; 26: 8; 27: 9; 28: 10; 29: 4; 30: 15; 31: 4; 32: 10; 33: 4; 35: 1; 36: 12; 37: 4; 38: 12; 39: 12; 40: 5; 41: 28; 42: 2; 43: 8; 44: 5; 45: 12; 46: 12; 47: 15; 48: 13; 49: 7; 50: 7; 51: 9; 52: 3; 53: 1; 54: 9; 55: 5; 56: 4; 57: 7; 58: 2; 59: 6; 61: 9; 62: 2; 63: 9; 64: 11; 65: 1; 66: 1; 67: 12; 68: 2; 69: 1; 70: 5; 71: 7; 72: 10; 73: 2; 74: 4; 75: 10; 76: 6; 77: 20; 79: 6; 80: 7; 81: 8; 82: 4; 83: 12; 84: 11; 85: 3; 86: 10; 87: 11; 88: 6; 89: 7; 90: 8; 91: 2; 92: 6; 93: 17; 94: 3; 95: 11; 96: 12; 97: 10; 98: 4; 99: 14; 100: 14; 101: 8; 102: 20; 103: 2; 104: 9; 105: 2; 106: 4; 107: 2; 108: 10; 109: 15; 110: 23; 111: 8; 112: 14; 113: 16; 114: 7; 115: 12; 116: 13; 117: 26; 118: 8; 119: 8; 120: 12; 121: 11; 122: 7; 123: 11; 125: 7; 126: 1; 127: 10; 128: 5; 129: 8; 130: 3; 131: 11; 132: 12; 133: 14; 134: 3; 135: 6; 136: 4; 137: 5; 138: 7; 139: 6; 140: 1; 141: 10; 142: 8; 143: 8; 144: 4; 145: 5; 146: 20; 147: 12; 148: 7; 149: 8; 150: 5; 151: 4; 152: 7; 153: 12; 154: 2; 155: 3; 156: 7; 157: 3; 158: 1; 159: 2; 160: 1; 161: 3; 162: 2; 163: 2; 164: 1; 165: 2; 166: 1; 170: 2; 171: 3. Distribución geográfica: Presenta una distribución centro-occidental euro- pea (BOATO ET AL., 1982). En la Península Ibérica se extiende por la franja portuguesa (HIDALGO, 1875; LOCARD, 1899; NOBRE, 1941) y por toda la mitad norte (BOFILL y HAAs, 1919, 1920a, 1920b; ORTIZ DE ZARATE y ORTIZ DE ZARATE, 1949; MANGA, 1983; RAMOS, 1985; APARICIO, 1986; HERMIDA ET AL., 1992; LARRAZ y EQUISOAIN, 1993; PAREJO ET AL., 1993b; ALTONAGA ET AL., 1994). En la mitad sur peninsular es más frecuente en Portugal, pero en los úl- timos años varios autores han ampliado su distribución en esta zona (MARTÍNEZ- ORTI y ROBLES, 1993; PAREJO, ET AL., 1993a). Helix (Cornu) aspersa (Múller, 1774) (Fig. 2L) Citas previas: MACHO VELADO (1871); HIDALGO (1875, 1890); SAccHI y VIOLANI (1977) como Cryptomphalus aspersus; CASTILLEJO (1986); OTERO y TRIGO (1989). Material examinado (1579 ejemplares). 1: 1; 2: 1; 3: 5; 4: 32; 5: 3; 6: 12; 8: 4; 9: 6; 10: 1; 11: 3; 12: 5; 13: 12; 14: 3; 16: 2; 17: 3; 19: 5; 20: 7; 21: 30; 22: 4; 23: 4; 24: 3; 25: 6; 26: 16; 27: 6; 28: 54; 29: 15; 30: 9; 31: 3; 32: 12; 35: 18; 36: 8; 37: 20; 38: 17; 39: 15; 40: 13; 41: 28; 42: 22; 43: 2; 44: 12; 45: 17; 46: 11; 47: 4; 48: 26; 49: 18; 50: 11; 51: 20; 52: 22; 53: 20; 54: 11; 55: 10; 56: 7; 57: 19; 58: 1; 59: 12; 60: 11; 61: 21; 62: 9; 63: 1; 64: 27; 65: 2; 67: 23; 68: 16; 69: 12; 70: 14; 71: 11; 73: 6; 75: 13; 76: 4; 77: 7; 78: 1; 79: 6; 80: 2; 81: 10; 82: 10; 83: 5; 84: 5; 85: 8; 86: 10; 87: 10; 88: 10; 90: 2; 91: 17; 92: 11; 93: 16; 94: 16; 95: 14; 96: 11; 97: 9; 98: 9; 100: 14; 101: 42; 102: 48; 103: 21; 104: 5; 106: 10; 109: 10; 110: 7; 111: 4; 112: 6; 113: 17; 114: 22; 115: 14; 116: 2; 117: 36; 118: 8; 119: 15; 120: 16; 121: 11; 122: 11; 123: 36; 124: 3; 125: 9; 126: 8; 127: 13; 128: 9; 129: 12; 130: 2; 131: 11; 132: 16; 133: 8; 134: 22; 135: 5; 136: 10; 137: 12; 138: 2; 139: 1; 141: 3; 143: 4; 144: 1; 145: 8; 146: 16; 147: 4; 148: 6; 149: 6; 150: 1; 151: 3; 152: 2; 153: 9; 154: 2; 155: 5; 156: 2; 157: 1; 158: 2; 159: 1; 160: 4; 162: 3; 165: 3; 167: 1; 168: 1; 171: 3. Distribución geográfica: Especie distribuida por el Mediterráneo y oeste de Europa. (BOATO ET AL., 1982). En la Península Ibérica se ha citado repetidamente en todas las regiones DISCUSIÓN En un principio y dadas las condicio- nes climáticas gallegas favorables para la vida de los gasterópodos terrestres, podrí- amos pensar que es un área de abundante fauna malacológica. Pero comparándola con otras zonas de la Península Ibérica enclavadas en el área atlántica podemos comprobar que la diversidad y abundan- (ALTONAGA ET AL., 1994) y en las Islas Baleares, y aún faltando zonas por estu- diar, dado su carácter ubiquista y sinan- trópico, puede asegurarse que está pre- sente en todo el territorio peninsular. cia de helícidos es especialmente escasa en Galicia. Probablemente esto es debido, principalmente, a la escasez de sustrato calizo, necesario para la formación de la concha de todos los gasterópodos en general y, de la conchas gruesas y duras de los helícidos en particular (BOYcorr, 1934; KERNEY y CAMERON, 1983). 19 Iberus, 15 (1), 1997 Con este trabajo se ha ampliado el área de distribución conocida de la ma- yor parte de las especies, pudiendo seña- lar la notable presencia en la zona de es- tudio de especies como H. aspersa, C. ne- moralis y P. inchoata, que aparecen en más del 80% de las cuadrículas visitadas. A diferencia de ésto, existen otras especies que han presentado distribuciones res- tringidas, especialmente aquellas origi- nariamente mediterráneas, intimamente ligadas al litoral como T. pisana, C. acuta, C. barbara, y C. conoidea, destacando es- pecialmente esta última, que al igual que sucede en el resto de la Península, no as- ciende por la costa cantábrica. En contra- posición a ésto, H. itala, también con un comportamiento litoral en el área de es- tudio, no desciende del norte de Galicia, siendo su cita más al sur, en todo este te- rritorio, la Sierra de O Courel, zona ca- liza con una altitud superior a los 900 m. También hemos de señalar que la mayor parte de las especies han sido capturadas en gran variedad de hábitats (bajo troncos, muros, linderas de culti- vos...), pudiendo destacar únicamente la preferencia de E. quimperiana por los diferentes tipos de arbolados, en mayor medida pinares y robledales, y de P. inchoata y P. subvirescens en los prados y zonas carentes de vegetación arborea. BIBLIOGRAFÍA ALTIMIRA, C., 1969. Notas Malacológicas. VIH Moluscos del Delta del Llobregat. IX Nuevas aportaciones y datos a la fauna malacoló- gica catalana. XI Moluscos terrestres y de agua dulce recogidos en la provincia de Lugo y en Asturias. Publicaciones del Instituto de Biología Aplicada, 46: 91-113. ALTONAGA, K.; GÓMEZ, B.; MARTÍN, R.; PRIETO, C.; PUENTE, A. y RALLO, A., 1994. Estudio fau- nístico y biogeográfico de los moluscos terrestres del norte de la Península Ibérica. 505 pp. Ed. Eusko Legebiltzarra, Parlamento Vasco. ANADÓN, N. y OJEA, M., 1984. Gasterópodos te- rrestres del Monte Naranco (Oviedo, Astu- rias). Distribución, diversidad y afinidades faunísticas. Revista de Biología de la Universi- dad de Oviedo, 2: 121-129. APARICIO, M. T., 1986. The geographic distri- bution of the Family Helicidae in Central Spain. Proceedings of the 8* International Ma- lacological Congress, Budapest. 20 Por último queremos mencionar que hemos encontrado la mayoría de las especies citadas anteriormente a noso- tros en el área de estudio, y las que no lo han sido podemos atribuirlo, no tanto a errores de muestreo, como a la posibili- dad de que se encuentren poco repre- sentadas. Este es el caso de Helicigona lapicida (Linneo, 1758) citada por ROLÁN y OTERO (1988) en una única localidad de la provincia de A Coruña. Así mismo, hay que señalar Otala punctata (Múller, 1771) citada como Otala lactea (Múller, 1771) por CASTILLEJO (1986) en la esta- ción biológica de Lourizán (Ponteve- dra). Esta especie de ámbito mediterrá- neo fue, con seguridad, introducida con un cultivo experimental en la estación agrícola, y la población, muy abundante en el momento de su cita (el autor recoge unos 300 ejemplares) probable- mente no prosperó, puesto que hemos realizado repetidas visitas a esa zona, sin encontrar ningún ejemplar. AGRADECIMIENTOS Este trabajo forma parte del Proyecto “Invertebrados terrestres de Galicia. 1. Provincias de La Coruña y Pontevedra” financiado por la Xunta de Galicia. BACKHUYS, W., 1975. Land and fresh-water mo- lluscs of the Azores. Backhuys y Meesters. Amsterdam. 350 pp. BEcH, M., 1990. Fauna malacologica de Cata- lunya. Molluscs terrestres i d'aigua dolca. Treballs de l'Institució catalana d'Históoria Na- tural, 12: 1-229. BOATO, A., BODON, M. y Giusti, F., 1982. Mo- lluschi terrestri e d'acqua dolce delle Alpi Li- guri. Lavori della societa Italiana di Biogeografia, Nuova serie, 9: 237-371. BOFILL, A. y Hass, F., 1919. Molluscos rellits en Asturias, en 1918 per en Josep Maluque pre- cedits de consideracions bibliografiques so- bre la Malacología asturiana. Butlletí de la Ins- titució Catalana d'Historia Natural, 12: 25-34. BOFILL, A. y Haas, F., 1920a. Estudi sobre la malacologia de les Valls Pirenaiques. Vall del Noguera Pallaresa. Treballs del Museu de Ciences Naturals de Barcelona, 3(10): 100- 220. ONDINA E7 42.: Distribución de la superfamilia Helicoidea en el oeste de Galicia BOFILL, A. y Haas, F., 1920b. Estudi sobre la ma- lacologia de les Valls Pirenaiques. Vall del Se- gre i Andorra. Treballs del Museu de Ciences Naturals de Barcelona, 3(12): 225-375. CASTILLEJO, J., 1984. Caracoles terrestres de Ga- licia, [. Género Oestophora Hesse, 1907. (Pul- monata, Helicidae). Iberus, 4: 125-133. CASTILLEJO, J., 1986. Caracoles terrestres de Gali- cia. Familia Helicidae (Gastropoda, Pulmonata). Servicio de Publicaciones de la Universidad de Santiago, 122, 66 pp. Cazior, E., 1915. La fauna terrestre Lusita- nienne. Bulletin de la Société Linnéenne de Lyon, 62: 43-65. DA SILVA MENGO, J., 1867. Descripcao de um “Helix” novo de Portugal. Journal de Sciences Mathematicas, Physicas e Naturales, 1: 170-171. GASULL, L., 1965. Algunos moluscos terrestres y de agua dulce de Baleares. Bolletí de la So- cietat d'Historia Natural de les Balears, 9: 1-80. GASULL, L., 1975. Fauna malacológica terrestre del sudeste ibérico. Bolletí de la Societat d'His- toria Natural de les Balears, 20: 5-155 + 4 pl. GERMAIN, L., 1930. Faune de France, 21. Mollus- ques terrestres et fluviatiles. 1-2. París. 897 pp. GITTENBERGER, E., 1979. On Elona (Pulmonata, Elonidae fam. nov). 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O Sociedad Española de Malacología Iberus, 15 (1): 23-29, 1997 Una nueva especie de Lirularía (Gastropoda: Trochidae) de las islas de Sáo Tomé y Príncipe, África Occidental A new species of Lirularia (Gastropoda: Trochidae) from Sao Tomé y Príncipe Islands, West Africa Federico RUBIO* y Emilio ROLÁN** Recibido el 18-111-1996. Aceptado el 8-X-1996 RESUMEN El estudio del material obtenido durante dos expediciones realizadas a las Islas de Sáo Tomé y Príncipe en los años 1989 y 1990, ha proporcionado una nueva especie de tró- quido con apariencia de Solariella, que por sus características morfológicas y radulares pertenece a la subfamilia Lirulariinae. Se describe esta especie, nueva para la ciencia, comentándose las características morfológicas de la concha, partes blandas y rádula, e incluyéndola en el género Lirularia. Se discute la inclusión en dicho género de otras dos especies de la costa occidental africana. ABSTRACT The study of the material obtained in Sáo Tomé and Principe Islands during two expedi- tions in the years 1989 and 1990, have yielded a new trochid species with Solariella, which for its morphological and radular characters belongs to the subfamily Lirulariinae. This species is described as new for science within the genus Lirularia and its morphologi- cal characteristics of shell, soft parts and radula are commented on. The assignment to this genus of other two species of the West African coast is discussed. PALABRAS CLAVE: Gastropoda, Trochidae, Lirularia, nueva especie, Islas de Siío Tomé y Príncipe. KEY WORDS: Gastropoda, Trochidae, Lirularia, new species, Sio Tomé and Príncipe Islands. INTRODUCCIÓN Se han citado varias especies de Sola- riella s. 1. de la costa africana en diversos trabajos. SMITH (1871) describe la primera, Solariella canaliculata Smith, 1871, de Whydah (Dahomey), actualmente Repú- blica de Benin. Esta misma especie y Sola- riella dereimsí Dollfus, 1911 son citadas para Angola por GOFAs, PINTO AFONSO Y BRANDAO (1985); NICKLÉS (1950) menciona Solariella monodi Fischer y Nicklés, 1946 para Guinea Francesa (Guinea Conakry) y S. dereimsi para Mauritania y Senegal. Alguna otra especie, como S. valida Daut- zenberg y Fischer, 1906, descrita para el archipiélago de Cabo Verde, procede de aguas profundas. En BERNARD (1984) aparece repre- sentada una concha con la denomina- *Dpto. de Zoología. Facultad de Ciencias Biológicas, Universidad de Valencia. Dr. Moliner, 50, 46100 Burjasot (Valencia). **Cánovas del Castillo, 22-59 F, 36202 Vigo. 23 Iberus, 15 (1), 1997 ción Solariella sp., pero en realidad se trata de Cyclostremiscus calameli (Jousse- aume, 1872) que es en realidad un Torni- dae (Gofas, com. pers.). No hay citas de especies de Solariella para el archipiélago de Sáo Tomé y Prín- cipe, aunque en el último listado de es- pecies de FERNANDES Y ROLÁN (1993), se menciona una única especie de este gé- nero, como Solariella sp. HERBERT (1987) demostró que muchas de las especies de África del Sur, consi- deradas tradicionalmente del género Sola- riella no lo son, sino que pertenecen a la subfamilia Umboniinae en lugar de Sola- riellinae. HICKMAN Y MCLEAN (1990) pu- blicaron una revisión de la superfamilia Trochoidea y, atendiendo a la morfología de sus partes blandas y rádula, agrupa- ron las “Solariellas” en un grupo informal formado por las subfamilias Trochinae Ra- finesque, 1815, Stomatellidae Gray, 1840, Calliostomatinae Thiele, 1924 y Solarielli- nae Powell, 1951; agrupando las especies de Umboniinae en otro grupo informal formado por Lirulariinae Hickman y McLean, 1990, Halistylinae Keen, 1958 y Umboniinae Adams y Adams, 1854. Wa- RÉN (1993), siguiendo dicha ordenación, considera que “Solariella” canaliculata y “Solariella” dereimsi pertenecen a la subfa- milia Umboniinae. Durante las campañas de recolección de moluscos efectuadas por el segundo RESULTADOS autor en los años 1989 y 1990 en el archi- piélago de Sáo Tomé y Príncipe, Golfo de Guinea, se recolectaron ejemplares y conchas de una “Solariella” aparente- mente diferente de las especies previa- mente conocidas. Esta misma especie, referida como Solariella sp. por FERNAN- DES Y ROLÁN (1993) y WARÉN (1993), es ahora descrita, aunque en un género diferente. Al mismo tiempo, las especies “Sola- riella” canaliculata y “Solariella” dereimsi, recolectadas y observadas en las expedi- ciones a Angola y Mauritania de 1989 y 1996 respectivamente, son transferidas a la subfamilia Lirulariinae, género Lirula- ria, basándonos en la similitud morfoló- gica de sus partes blandas y rádula. MATERIAL Y MÉTODOS Se han estudiado 8 ejemplares y 82 conchas, procedentes de sedimentos obtenidos en distintas localidades del archipiélago, mediante buceo a pulmón libre, a profundidades comprendidas entre 5 y 15 metros. Tras la observación de su comportamiento, algunos indivi- duos se relajaron y posteriormente se fijaron en solución tamponada de for- maldehído al 5%. Para la observación conquiológica y radular se ha utilizado la microscopía electrónica de barrido. Superfamilia TROCHACEA Rafinesque, 1815 Familia TROCHIDAE Rafinesque, 1815 Subfamilia LIRULARINAE Hickman y McLean, 1990 Género Lirularia Dall, 1909 Lirularia antoniae spec. nov. (Figs. 1A-B, 2 - 6) Material estudiado: Isla de Sáo Tomé: Praia das conchas: 8 ejemplares y 16 conchas a -15 m; Ciudad Sáo Tomé: 36 conchas. Isla de Príncipe: Bahía de Santo Antonio: 24 conchas a -10 my; Bahía das Agulhas: 6 conchas. Material tipo: Holotipo (Fig. 1B) y dos paratipos procedentes de la localidad tipo, depositados en el Museo Nacional de Ciencias Naturales de Madrid, con el n* 15.05/23749, dos paratipos proce- dentes de Praia das Conchas (Sáo Tomé) en el Muséum National d Histoire Naturelle de París, dos paratipos procedentes de Ciudad de Sáo Tomé en el American Museum of Natural History de Nueva York y en The Natural History Museum de Londres y 25 en cada una de las colecciones de los autores. Localidad tipo: Praia das Conchas, Sáo Tomé. Etimología: La especie está dedicada a Antonia Hueso, esposa del primer autor. 24 RUBIO Y ROLÁN: Nueva especie de Lirularia de Sáo “Tomé y Príncipe UN AN Figura 1. Lirularia antoniae spec. nov. Holotipo. Praia das Conchas, Sáo Tomé. A: animal en movi- miento. B: concha (MNCN). Escalas 1 mm. Abreviaturas. o: ojo; op: opérculo; p: pie pd: proceso digitiforme sobre el extremo del morro; s: sifón; tc: tentáculo cefálico; te: tentáculo epipodial. Figure 1. Lirularia antoniae spec. nov. Holotype. Praia das Conchas, Sáo Tomé. A: crawling animal. B: shell (MNCN). Scale bars 1 mm. Abbreviations. o: eye; op: operculum; p: foot; pd: digitiform processes on the tip of'snout; s: siphon; tc: cep- halic tentacle; te: epipodial tentacle. Descripción: Concha sólida, bri- llante, nacarada, de perfil cónico y espira algo elevada, compuesta por unas 5 vueltas convexas, que están sepa- radas por una sutura ancha y acanalada. Protoconcha (Fig. 5) con apenas una vuelta de espira, lisa, con el núcleo de- formado y de unas 200 ym. Ornamenta- ción formada por cordones espirales y costillas transversales muy numerosas, que al entrecruzarse forman pequeños nódulos; se observan, además, sutiles líneas de crecimiento que se extienden paralelas a las costillas. Última vuelta con 10 cordones espirales, de los que el primero, subsutural, y el décimo, periumbilical, son los más prominentes por ser nodulosos y angulan la concha. Los restantes cordones son poco marca- dos, sobre todo en la parte media de la periferia, donde apenas son percepti- bles. Las costillas transversales están menos marcadas en la primera y última vuelta de la teloconcha, aunque en ésta última son numerosísimas; su curso es prosoclino y atraviesan la totalidad de la vuelta. Ombligo ancho y profundo, bor- deado por el décimo cordón espiral; en su interior se observan otros cuatro cor- dones espirales más. Abertura subcircu- lar, prosoclina; labio externo fino, angu- lado por la presencia de los cordones espirales; labio interno, ligeramente arqueado, reflejado hacia el exterior, pero sin llegar a ocluir el ombligo. Coloración muy variable, de blanco- amarillento a rosa pálido, con manchas pardo rojizas o pardo oscuras y cierta iridiscencia. Respecto a sus dimensiones, el holo- tipo (Fig. 1B) mide 1,78 mm de altura y 1,98 mm de anchura. El animal (Fig. 1A) es de color blan- quecino excepto el sifón y una franja de color negro situada en la parte distal del morro. La cabeza tiene un par de tentá- culos cefálicos muy largos y con micro- papilas, ojos negros situados sobre cortos pedúnculos y carece de membra- nas cefálicas. El morro está muy depri- 23) Tberus, 15 (1), 1997 mido distalmente y sus extremos se pro- longan transversalmente; hay un proceso digitiforme con forma de peine sobre su extremo. A cada lado de la cabeza se observa un lóbulo cervical modificado; el izquierdo, subdividido en apéndices tentaculiformes y, el derecho, que es plano, en el animal vivo, se enrolla para formar una estructura tubular con aspecto de sifón, moteado con manchas negro-opacas y de un tamaño similar a los tentáculos cefálicos. Epipodio con cuatro pares de tentácu- los, carentes de macropapilas sensoria- les en su base, el primer par en posición anterior y los tres pares restantes alrede- dor del lóbulo opercular. El pie es muy móvil, su extremo anterior es bilobulado y se prolonga lateralmente y luego se afila progresivamente hacia su extremo posterior, para acabar en punta. Rádula (Fig. 6) formula N. 4. 1. 4. N. El diente central y los laterales son muy similares y están reducidos a láminas basales con una pequeña cúspide cada uno. El diente central está cubierto en parte por las láminas laterales más inter- nas. Dientes marginales con cúspides relativamente largas y anchas, con den- tículos romos y aserrados. Distribución: Sólo conocida del archipiélago de Sáo Tomé y Príncipe (Golfo de Guinea). No se han encon- trado ejemplares de esta especie en zonas continentales próximas al archi- piélago, por lo que, probablemente, se trata de un endemismo insular. Hábitat: Especie infralitoral que vive sobre fondos de arena en zonas de aguas claras, entre -5 y -15 metros. Discusión: Tanto la estructura plana, enrollada en forma de sifón, como los procesos digitiformes visibles a cada lado de la cabeza en Lirularia antoniae, al igual que en otras especies de África oc- cidental (“Solariella” canaliculata y “Sola- riella” dereimsi1) (Gofás com. pers.), son probablemente lóbulos cervicales modi- ficados, homólogos a los de otros Trocoi- deos. Su función, para el lado inhalante, se puede suponer que es la de actuar 26 como un filtro para evitar la entrada de las partículas en la cavidad paleal. El ló- bulo derecho tiene una función exha- lante. Dichas estructuras surgen como consecuencia de la adaptación de las es- pecies de Solariellinae, Umboniinae y Lirulariinae de África occidental a los fondos blandos sobre los que habitan, a diferencia de otros trocoideos (Clancu- lus, Collonia, Gibbula, Tricolia, etc.) cuyo hábitat está limitado a fondos rocosos. Aspectos de esta adaptación coinciden- tes con estas características anatómicas pueden verse en FRETTER (1975) y HICk- MAN (1985). Siguiendo la clasificación de la fami- lia Trochidae propuesta por HICKMAN Y MCLEAN (1990), y atendiendo a los ca- racteres morfológicos y radulares que diferencian las tres subfamilias que per- tenecen al grupo informal Halistylinae + Umboniinae + Lirulariinae, hemos si- tuado la nueva especie en la subfamilia Lirulariinae, género Lirularia. Las espe- cies “Solariella” canaliculata (Fig. 7) y “Solariella” dereimsil (Figs. 8-9) son con- genéricas entre sí, y se diferencian ana- tómicamente de Lirularia antontae tan solo por la distribución de las papillas del morro; sin embargo, comparten ca- racteres comunes como: e protoconchas de pequeño tamaño, no superior a 200 um, con el núcleo deformado; e ombligo amplio, no ocluido ni total ni parcialmente; e radulas similares, muy parecidas a su vez a las de Umbonium; e lóbulo cervical izquierdo subdivi- dido en un proceso tentaculiforme y lóbulo cervical derecho plano, enrollado para formar una especie de sifón. Todo esto nos hace considerarlas pertenecientes a la subfamilia Lirularii- nae, género Lirularia, en lugar de Sola- riellinae o Umboniinae. Eirularia antoniae, L. canaliculata y L. dereímsii, se diferencian de las especies pertenecientes a Solariellinae por tener protoconcha pequeña con el núcleo de- formado, la rádula con su zona central simplificada, el diente central y los dien- RUBIO Y ROLÁN: Nueva especie de Lirularia de Sao “Tomé y Príncipe Figuras 2-6. Lirularia antoniae spec. nov., Praia das Conchas, Sáo Tomé. 2: individuo juvenil, vista apical (col. E Rubio); 3: individuo juvenil, vista dorsal; 4: paratipo (col. E. Rubio); 5: protoconcha; 6: rádula. Escalas, 2-4: 0,5 mm; 5: 100 um; 6: 12 pm. Figures 2-6. Lirularia antoniae spec. nov., Praia das Conchas, Sáo Tomé. 2: young specimen, apical view (E Rubio coll.). 3: young specimen, dorsal view. 4: paratype (E Rubio coll.); 5: protoconch; 6: radula. Scale bars, 2-4: 0,5 mm; 5: 100 ym; 6: 12 um. 27 Iberus, 15 (1), 1997 Figuras 7-9. Animal de otras especies de Lirularia de Africa. 7: L. dereimsiz (tomado de GOFAS, PINTO AFONSO Y BRANDAO, 1985); 8-9: L. canaliculata (dibujo de S. Gofas). Figures 7-9. Animal of other species of Lirularia from Africa. 7: L. dereimsii (after GOFAS, PINTO AÁFONSO AND BRANDAO, 1985); 8-9: L. canaliculata (drawing from S. Gofas). tes laterales reducidos, los tentáculos epi- podiales carentes de macropapilas senso- riales en su base y las papilas situadas en el extremo anterior del morro y no alre- dedor del disco oral. De las especies per- 28 tenecientes a Umboniinae, tribu Um- boniini se diferencian porque estas últi- mas poseen tentáculos cefálicos dimórfi- cos y el lóbulo cervical izquierdo en- vuelve el tentáculo cefálico izquierdo y RUBIO Y ROLÁN: Nueva especie de Lirularia de Sao Tomé y Príncipe pedúnculo ocular. De las especies perte- necientes a la tribu Monileini se diferen- cian porque presentan un ombligo cerrado, total o parcialmente, por un callo; por po- seer pedúnculos oculares prominentes, AGRADECIMIENTOS Al Servicio de Microscopía Electró- nica de la Universidad de Valencia, por la ayuda prestada en la realización de foto- grafías al Microscopio Electrónico de Ba- rrido. A Francisco Fernandes por su cola- BIBLIOGRAFÍA BERNARD, P. A., 1984. Coquillages du Gabon. Li- breville. 140 pp., 75 láms. FERNANDES, F. Y ROLÁN, E., 1993. Moluscos ma- rinos de Sao Tomé y Principe: actualización bibliográfica y nuevas aportaciones. Iberus, 11 (1): 31-47. FRETTER, V., 1975. Umbonium vestiatium, a filter- feeding trochid. Journal of Conchology, 177: 541-552. GOFAS, S., PINTO AFONSO, J. Y BRANDAO, M., 1985. Conchas e moluscos de Angola. Universi- dad de Agostinho Neto / Elf Aquitaine. An- gola. 139 pp. HERBERT, D. G., 1987. Revision of the Solarie- llinae (Mollusca: Prosobranchia: Trochidae) in South Africa. Annals of the Natal Museum, 28: 283-382. HICKMAN, C. S., 1985. Comparative morphology and ecology of free living suspension-fee- ding gastropods from Hong Kong. En Mor- ton, B. y Dudgeon D. (Eds.): Proceedings of the second International Workshop on the Malaco- fauna of Hong kong and Southern China, Hong kong University Press.: 217-234. con anchos ojos y bases desarrolladas; y porque su lóbulo cervical izquierdo (in- halante) está subdividido terminalmente en un proceso tentaculiforme dimórfico, orientado regular y alternativamente. boración en las expediciones a Sáo Tomé y Príncipe. A Serge Gofas por la lectura crítica del manuscrito, por su informa- ción y aportación de dibujos, y por sus sugerencias. HICKMAN, C. S. Y MCLEAN, J. H., 1990. Syste- matic revision and suprageneric classification of trochacean gastropods. Natural History Mu- seum of Los Angeles County, Science Se- ries, 35. 169 pp. NICKLÉS, M., 1950. Mollusques testacés marins de la cóte occidentale d'Afrique. Lechevalier, Pa- ris, 269 pp., 434 figs. SmITH, E. A., 1871. A list of species of shells from West Africa, with descriptions of those hit- herto undescribed. Proceedings of the Zoological Society of London, 1871: 727-739. WARÉN, A. S., 1993. New and little know Mo- llusca from Iceland and Scandinavia. Part 2. Sarsia, 78: 159-201. 2D, pe AS 2 AA a) yr turn hala E Acaban ii Ñd óí a 8d besa bl hos pe ep ienecion tes TS ropnpilaa Je Era Al E e bl . O Sociedad Española de Malacología Iberus, 15 (1): 31-34, 1997 Una nueva especie de Anticlimax (Gastropoda: Vitrinelli- dae) de Cuba A new species of Anticlimax (Gastropoda: Vitrinellidae) from Cuba Emilio ROLÁN” Raúl FERNÁNDEZ-GARCÉS” y Federico RUBIO” Recibido el 18-111-1996. Aceptado el 8-X-1996 RESUMEN Se describe una nueva especie del género Anticlimax (Gastropoda: Vitrinellidae). Se dis- cute su asignación genérica y se compara con las especies más próximas. ABSTRACT A new species of the genus Anticlimax (Gastropoda: Vitrinellidae) is described. lts generic assignment is discussed and a comparison is made with allied species. PALABRAS CLAVE: Gastropoda, Vitrinellidae, Anticlimax, especie nueva, Cuba. KEY WORDS: Gastropoda, Vitrinellidae, Anticlimax, new species, Cuba. INTRODUCCIÓN DALL (1903) menciona por primera vez el subgénero Climacia en la descrip- ción de Teinostoma (Climacia) calliglyp- tum, sin hacer descripción alguna de las características del subgénero ni de las de la especie típica, siendo ambos definidos por figuras (PILSBRY Y MCGINTY, 1946a). PILSBRY Y MCGINTY (1946a) dan a Climacia categoría genérica y describen y comentan las características de T. calliglyptum como las de la especie típica (por monotipia); al tiempo describen varias especies nuevas que incluyen en Climacia. AGUAYO Y BORRO (1946) sustituyen por Climacina el previamente ocupado nombre genérico de Climacia Dall, 1903 non M'Lachlan, 1869 (Neuroptera). Poco " Cánovas del Castillo 22, 36202 Vigo, España. “Calle 41, 6607, Cienfuegos 55100, Cuba. tiempo después, PiLsBRY Y MCGINTY (1946b) sustituyen Climacina Aguayo y Borro, 1946 non Gemmellaro, 1878 (Mollusca), por un nuevo nombre: Anti- climax. PILSBRY Y OLSSON (1950) hacen una revisión del género describiendo algunas especies nuevas del Terciario americano, y lo subdividen en dos sub- géneros Anticlimax y Subclimax subgen. nov. En el material recolectado en Cuba durante los muestreos realizados por los dos primeros autores en los últimos años, se han encontrado conchas de una especie que parece pertenecer a este género, y que es descrita en el presente trabajo. Departamento de Zoología, Universidad de Valencia, Dr. Moliner 50, 41600 Burjasot, España. 31 Iberus, 15 (1), 1997 RESULTADOS Género Anticlimax Pilsbry y McGinty, 1946 Especie tipo, por monotipia: Teinostoma (Climacia) calliglyptum Dall, 1903 Anticlimax decorata spec. nov. Material tipo: Holotipo, de Rancho Luna, Cienfuegos, Cuba (Fig. 2), con 1,3 mm de dimensión máxima, Museo Nacional de Ciencias Naturales de Madrid n* 15.05/27420. De la misma locali- dad, un paratipo en las colecciones del American Museum of Natural History de Nueva York (Fig. 1) y The Natural History Museum de Londres y dos en la de R. Fernández Garcés. Zona del Hotel Comodoro, La Habana, Cuba: un paratipo en la colección del Instituto de Ecología y Siste- mática de La Habana. Jibacoa, Distrito de La Habana, Cuba: un paratipo en la colección de F. Rubio y otro en la de E. Rolán. Localidad tipo: Rancho Luna, Cienfuegos, Cuba. Etimología: El nombre específico deriva de su elaborada escultura. Descripción: Concha (Figs. 1 y 2) discoidal algo ovalada, aplanada en la base y con el ápice apenas saliente, espira con elevación uniforme, suave- mente convexa y no muy pronunciada. Color blanquecino. Protoconcha lisa con 1*/4 vueltas de espira. Termina bruscamente y de forma bien delimitada con la teloconcha. Teloconcha formada por 1*/4 vuel- tas. La microescultura es compleja: al final de la última vuelta hay tres cordo- nes espirales en posición subsutural, de los cuales, los dos primeros se inician de forma progresiva, mientras el tercero se origina por una división del segundo. El resto de la superficie presenta surcos bastante uniformes constituidos, al prin- cipio, por perforaciones y, posterior- mente, por hundimientos de forma más alargada. Esta escultura desaparece casi por completo en el último cuarto de vuelta de forma muy constante. Hacia la parte inferior de la última vuelta la concha se angula y se hace prominente por medio de un grueso cordón que sobresale del perfil de la concha a modo de quilla y sobre el cual aparecen varios cordoncillos en zigzag. Este cordón, visto desde la parte superior de la concha, es más visible por un lado que por el otro, lo que aumenta el perfil ovoide de la concha. La parte de la última vuelta que está por debajo de la angulación apenas sobrepasa al cordón SZ periférico y es cóncava en la proximidad del mismo. En su parte central hay un ombligo profundo. Toda la base pre- senta cordoncillos espirales; los que bordean el ombligo son gruesos y zigza- gueantes, y lo mismo ocurre con los que se sitúan sobre la quilla, siendo los res- tantes algo más finos y uniformes. Abertura circular, con borde fino, engrosado en su parte externa por el final del cordón espiral sobresaliente. El animal es desconocido. Discusión: La especie ahora descrita es incluida en el género Anticlimax como una aproximación, ya que presenta algunas diferencias con la especie tipo. Estas diferencias son, principalmente, la carencia de escultura axial en la base y la ausencia de una prolongación de la abertura en la finalización del cordón espiral. Sin embargo, la subespecie A. tholus prodromus Pilsbry y Olsson, 1950, apenas presenta escultura axial en la base y también carece de la prolonga- ción de la abertura, habiendo sido no obstante incluida en este género. A. deco- rata spec. nov., además de parecerse a esta última especie, tiene muchas de sus características coincidentes con especies que han sido consideradas pertene- cientes al género Anticlimax, como la base aplanada o cóncava, el cordón peri- férico, la superficie convexa por encima del mismo, la existencia de cordones en ROLÁN E7 4£.: Una nueva especie de Anticlimax de Cuba Figuras 1, 2. Anticlimax decorata spec. nov. 1: paratipo, Rancho Luna, Cienfuegos, Cuba, American Museum of Natural History de Nueva York; 2: holotipo, Rancho Luna, Cienfuegos, Cuba, Museo Nacional de Ciencias Naturales de Madrid. Escala 0,5 mm. Figures 1, 2. Anticlimax decorata 2. sp. 1: paratype, Rancho Luna, Cienfuegos, Cuba, American Museum of Natural History New York; 2: holotype, Rancho Luna, Cienfuegos, Cuba, Museo Nacional de Ciencias Naturales, Madrid. Scale bar 0.5 mm. el dorso formados por perforaciones, y los cordones en zigzag de la base. Su parecido con A. athleenae (Pilsbry y McGinty, 1946) es bastante notable, aunque esta especie se diferencia de A. decorata porque presenta ondulaciones en la base, y carece, tanto en la base como en el cordón periférico, de cordon- cillos en zigzag. Tambien con A. tholus (Pilsbry y McGinty, 1946) tiene un cierto” parecido, pero el ombligo de esta última está ocluido por un fuerte callo. Las restantes especies incluidas en este género, o tienen escultura axial, o el 33 Iberus, 15 (1), 1997 ombligo está ocluido por un callo, o tienen una prolongación triangular en la abertura como continuación de la quilla periférica, por lo que son claramente diferentes de A. decorata. Según ESPINOSA, FERNÁNDEZ-GAR- CÉs Y ROLÁN (1995), para la fauna de Cuba, únicamente era conocida una especie en el género Anticlimax: A. pro- boscidea (Aguayo, 1949), la cual es más elevada y tiene una gran prolongación de la abertura. La principal diferencia entre los dos subgéneros, Anticlimax y Subclimax, en los que PILSBRY Y OLSSON (1950) dividie- ron las especies del género Anticlimax es que, en el primero, el ombligo es evi- dente mientras que, en el segundo, existe un callo columelar que cubre par- AGRADECIMIENTOS A Bernardo Fernández Souto del Servicio General de Apoyo a la Inves- tigación de la Universidad de A Co- BIBLIOGRAFÍA AGUAYO, C. G. Y BORRO, P., 1946. Nuevos mo- luscos del Terciario superior de Cuba. Revista de la Sociedad Malacológica “Carlos de la Torre”, 4 (1): 9-12, 1 lám. DaLL, W. H., 1903. Two new mollusks from the West Coast of America. The Nautilus, 17 (4): 37-38. ESPINOSA, J., FERNÁNDEZ GARCÉS, R. Y ROLÁN, E., 1995. Catálogo actualizado de los molus- cos marinos de Cuba. Reseñas Malacológicas, 9: 1-90. 34 cial o totalmente el ombligo. Estos autores mencionan que no conocen especies de estructura intermedia entre ambos subgéneros. Anticlimax decorata, por su ombligo abierto y carencia de callo columelar, no estaría incluida en el subgénero Subclimax; pero por la caren- cia de cualquier tipo de prolongación triangular en la abertura, tampoco pre- sentaría perfectamente las características del subgénero Anticlimax. Por tanto, Anticlimax decorata representa una forma intermedia entre ambos subgéneros. En cualquier caso, en ausencia de información sobre las partes blandas, es preferible no asumir su posición taxonó- mica genérica más que como probable y no hacer uso de asignación subgenérica alguna. ruña, por la realización de las fotogra- fías en el microscopio electrónico de barrido. PiLsBRY, H. A. Y MCGINTY, T. L., 1946a. “Cy- clostrematidae” and Vitrinellidae of Flo- rida and Bahamas, III. The Nautilus, 59: 77- 83, pl. 8. PiLSBRY, H. A. Y MCGINTY, T. L., 1946b. Vitri- nellidae of Florida, part. 4. The Nautilus, 60 (1) 12-18. PiLsBRY, H. A. Y OLSSON, A. A., 1950. Review of Anticlimax, with new tertiary species (Gas- tropoda, Vitrinellidae). Bulletin of American Pa- leontology, 33 (135): 105-116, lám. 17-20. O Sociedad Española de Malacología Iberus, 15 (1): 35-40, 1997 La familia Pyramidellidae Gray, 1840 (Mollusca, Gastro- poda) en África occidental. 1. El género Sayella Dall, 1885 The family Pyramidellidae Gray, 1840 (Mollusca, Castiopodaa in West Africa. 1. The genus Sayella Dall, 1885 Anselmo PEÑAS* y Emilio ROLÁN** Recibido el 8-VIIL-1996. Aceptado el 8-X-1996 RESUMEN Se describen dos especies nuevas del género Sayella Dall, 1885 de África Occidental, una de ellas encontrada en los Archipiélagos de Cabo Verde y de Sáo Tomé y Príncipe, y otra, sólo en el segundo de ellos. ABSTRACT Two new species of the genus Sayella Dall, 1885 from West Africa are described; one of them found in the archipelagos of Cape Verde, and Sáo Tomé and Principe, the other one only in the latter. PALABRAS CLAVE: Pyramidellidae, Sayella, África occidental, especies nuevas. KEY WORDS: Pyramidellidae, Sayella, West Africa, new species. INTRODUCCIÓN Después de realizada la revisión de los piramidélidos del Mediterráneo ibérico (PEÑAS, TEMPLADO Y MARTINEZ, 1996) y animados por los recientes tra- bajos de NOFRONI Y SCHANDER (1994) y SCHANDER (1994), en los que se descri- ben 31 especies nuevas de este grupo en África occidental, unido a la gran canti- dad de material que poseemos, nos decidió a abordar el estudio de esta familia en las costas atlánticas africanas. Comenzamos esta revisión con el género Sayella Dall, 1885, hasta ahora no men- cionado en el área geográfica de estudio, y continuaremos de forma inmediata con la de los géneros Turbonilla, Eulime- lla, Chrysallida y otros. El género Sayella fue descrito por DaLL (1885) para una especie fósil del Mioceno y bajo Plioceno de Louisiana. Inicialmente, la especie tipo, Leuconia hemphillii Dall, 1884, fue incluida por su autor en la familia Ellobiidae (Pulmo- nata). Sayella no aparece mencionado en las revisiones genéricas de THIELE (1931- 35) o de WENZ (1938). La posición de este género y sus relaciones con otros Pyramidellidae son discutidas por ODE (1994), que comenta sus diferencias con Odostomia y la dificultad para la separa- ción de especies, revisando también los taxones del Atlántico occidental. Las características del género son referidas por ABBOTT (1974) como “concha pu- * Carrer Olérdola, 39; 08800 Vilanova i la Geltrú. Barcelona. ** Cánovas del Castillo, 22; 36202 Vigo. 39 ber ISO poide-alargada, frágil, lisa, con vueltas convexas. Animal con tentáculos apla- nados, triangulares. Periostraco amari- llento”. Se ha señalado en aguas someras y fondos fangosos (ABBOTT, 1974). Recientemente, WISE (1996), basán- dose en datos anatómicos, crea la subfa- milia Sayellinae para incluir a las espe- cies de este género. Asimismo, describe para la especie S. crosseana (Dall, 1885) un nuevo género, Petitiella, por sus dife- rencias anatómicas, al que también incluye en esta subfamilia. En el presente trabajo abordamos la descripción de dos especies que consi- deramos nuevas y que asignamos provi- sionalmente a Sayella, pués carecemos de los datos anatómicos necesarios para confirmar su pertenencia a este género. Dado el escaso número de vueltas de su protoconcha, cabe pensar que estas especies carecen de un desarrollo larva- rio plantotrófico, y que, por tanto, es poco probable una relación estrecha con especies del Atlántico occidental. Dicho género no ha sido mencionado con RESULTADOS anterioridad en las costas occidentales de África. MATERIAL Y MÉTODOS El material estudiado en el presente trabajo procede de las expediciones efec- tuadas por el segundo de los autores al archipiélago de Cabo Verde entre los años 1978 y 1988, y a Sáo Tomé y Prín- cipe en los años 1989 y 1990. Dicho material fue separado de los detritos recogidos mediante buceo y dragado. Abreviaturas empleadas: AMNH: American Museum of Natural History, New York BMNH: The Natural History Museum, Londres CER: Colección de E. Rolán CPM: Colección de P. Micali MNCN: Museo Nacional de Ciencias Naturales, Madrid MNHN: Museum National d'Histoire Naturelle, Paris Familia PYRAMIDELLIDAE Gray, 1840 Subfamilia SAYELLINAE Wise, 1996 Género Sayella Dall, 1885 Las características del género Sayella son: concha pequeña, pupoide alargada, vueltas lisas, sutura poco profunda, bandas espirales de color, protoconcha corta con núcleo oculto, y periostraco acastañado. Sayella micalii spec. nov. (Figs 1-5) Material tipo: Holotipo (Fig. 1) de 2,86 x 1,35 mm depositado en MNCN (n* 15.05/23757) y 1 paratipo, ambos de la Bahía de Santo Antonio, Príncipe, República de Sáo Tomé y Príncipe. Otros paratipos en las siguientes colecciones: MNHN, 1 concha y dos juveniles de Bahía das Agulhas, Príncipe; BMNH, 1 concha, y AMNH, 1 concha, ambas a -10 m, de Regona, Isla de Sal, Archipiélago de Cabo Verde; CPM, 1 concha, -8 m, de Furna, Isla de Brava, Archipiélago de Cabo Verde; CER, 1 concha, -2 m, de la ciudad de Sáo Tomé, República de Sao Tomé y Príncipe y otra, -10 m, de Furna, Isla de Brava, Cabo Verde. Localidad tipo: Bahía de Santo Antonio, en la isla de Príncipe, Archipiélago de Sáo Tomé y Prín- cipe. Etimología: El nombre específico está dedicado al malacólogo Pasquale Micali, experto en Pyra- midellidae del Mediterráneo, por su habitual cooperación en el estudio de estos moluscos. 36 PEÑAS Y ROLÁN : El género Sayella (Pyramidellidae) en África occidental Figuras 1-4. Sayella micalii spec. nov. 1: holotipo (MNCN), Bahía de Santo Antonio, Príncipe, Archi- piélago de Sáo Tomé y Príncipe; 2, 3: protoconcha; 4: detalle de la columela y de la microescultura. Figures 1-4. Sayella micalii 2. sp. 1: holotype (MNCN), Santo Antonio Bay, Principe, Archipelago of Sáo Tomé and Principe; 2, 3: protoconch; 4: detail of the columela and microesculpture. Descripción: Concha (Figs. 1 y 5) oval-cónica, muy frágil. Protoconcha (Figs. 2 y 3) obtusa, muy grande, de tipo B (según Aartsen, 1981), con un diámetro de 490 yum. Eje con un ángulo de unos 100” en relación al de la teloconcha, y núcleo parcial- mente oculto. Coloración castaña clara. Teloconcha con vueltas convexas, algo escalonadas, que crecen deprisa; la última vuelta es muy grande, representando un 60% del total de su altura. Sutura pro- funda con una débil repisa subsutural. Vueltas de espira lisas, excepto débiles líneas de crecimiento ligeramente proso- clinas y estrías microscópicas espirales, más visibles en la zona umbilical (Fig. 4). Abertura grande, oval alargada, con un labio externo simple, cortante. Colu- mela ligeramente arqueada, delgada, pero muy replegada hacia la zona umbilical; tiene dos dientes, el mayor muy saliente 37 Iberus, 15 (1), 1997 y situado en la parte superior de la colu- mela; el segundo en la parte central, en forma de cordoncillo oblícuo. Fisura umbi- lical profunda. Color vítreo, semitranspa- rente, con una banda suprasutural castaña y otra en la base de la última vuelta, en la que también se aprecia otra banda subsu- tural (Fig. 5). Periostraco amarillento. Distribución: Conocida únicamente de los Archipiélagos de Cabo Verde y Sáo Tomé y Príncipe. Hábitat: En sedimentos de arena y fango, entre 4 y 10 m de profundidad. Discusión: No hay ninguna especie similar a Sayella micalíí spec. nov. en el Mediterráneo ni en las costas de África occidental. Todas las especies del Caribe asignadas a este género (ver ABBOTT, 1974) son diferentes porque tienen más vueltas de teleoconcha, con un incre- mento más lento de la espira y una forma más lanceolada que cónica. Sayella mercedordae spec. nov. (Figs. 6-8) Material tipo: Holotipo (Fig. 7) de 2,10 x 0,90 mm depositado en MNCN (n* 15.05/23758); 1 paratipo (Fig. 6) de Rife de Chaves, Boavista, en CER. Localidad tipo: Bahía de Mordeira, en la isla de Sal, Archipiélago de Cabo Verde. Etimología: El nombre específico está dedicado a Mercedes Dorda, esposa del malacólogo Esteban Calderón, por su colaboración a lo largo de toda su vida en la creación y mantenimiento de su colección. Descripción: Concha (Figs. 6 y 7) oval-cónica, pequeña, y frágil. Protoconcha (Figs. 8) pequeña, de tipo B, tendente a € (de acuerdo con AARTSEN, 1981). Su eje forma un ángulo de unos 100% en relación al de la telocon- : cha. Núcleo oculto. Teloconcha con vueltas convexas, que crecen deprisa en altura y anchura. La última vuelta es muy grande, repre- sentando un 70% del total de la altura de la concha. Sutura muy marcada, algo profunda. Vueltas de espira lisas, excep- tuando líneas de crecimiento ligera- mente prosoclinas y estrías microscópi- cas espirales en la base. Abertura oval-piriforme, estrechada hacia arriba. Columela opistoclina, algo arqueada, sin un diente claro, pero con un pliegue columelar en su parte cen- tral. No hay ombligo. Color blanquecino con dos estrechas bandas de color pardo por vuelta, y otra más, del mismo color, próxima a la zona columelar (Fig. 8). AGRADECIMIENTOS A José Bedoya por las fotografías al MEB realizadas en el MNCN de Madrid. 38 Distribución: Conocida únicamente del Archipiélago de Cabo Verde. Hábitat: Fondos arenosos entre 3 y 5 metros de profundidad, con zonas de fango próximas. Discusión: Sayella micalii spec. nov., es mucho grande, ancha, con forma más piramidal y con protoconcha mucho más gruesa. Las especies del Caribe son todas de mayor tamaño; Sayella fusca (C. B. Adams, 1839) tiene la sutura menos pro- funda y una banda clara subsutural; S. livida Rehder, 1935, tiene también una banda clara subsutural y un mayor número de vueltas de espira; S. crosseana (Dall, 1885) es mucho más alargada y proporcionalmente más estrecha; S. hemphilli (Dall, 1889) y S. chesapeakea Morrison, 1939, tienen la sutura apenas marcada y un número mayor de vueltas de espira. A Pasquale Micali por la lectura crítica del manuscrito. PEÑAS Y ROLÁN : El género Sayella (Pyramidellidae) en África occidental Figura 5. Sayella micalii spec. nov. Dibujo del paratipo (CER) mostrando la distribución de las ban- das de color. Figuras 6-8. S. mercedordae spec. nov. 6: dibujo de un paratipo (CER) mostrando la distribución de las bandas de color; 7: holotipo (MNCN), Bahía de Mordeira, Isla de Sal, Archipiélago de Cabo Verde; 8: protoconcha. Figure 5. Sayella micalii 2. sp. Drawing of paratype (CER) showing the distribution of the colour bands. Figures 6-8. S. mercedordae 7. sp. 6: drawing of a paratype (CER) showing the distribution of the colour bands; 7: holotype (MNCN), Mordeira Bay, Sal Island, Archipelago of Cape Verde; 8: protoconch. BIBLIOGRAFÍA AARTSEN, J. J. VAN, 1981. European Pyramide- llidae: HU. Turbonilla. Bollettino Malacologico, 17 (5-6): 61-88. ABBOTT, R. T., 1974. American seashells. Van Nos- trand Reinhold Co. New York. 663 pp., 24 láms. NOFRONI, I. Y SCHANDER, C., 1994. Description of three new species of Pyramidellidae (Gas- tropoda, Heterobranchia) from West Africa. Notiziario CISMA, 15: 1-10. ODE, H., 1994. Monograph. Distribution and Re- cords of the marine Mollusca in the Northwest Gulf of Mexico. Texas Conchologist, 31 (1): 9-32. 39 Iberus, 15 (1), 1997 Prxas, A., TEMPLADO, J. Y MARTINEZ, J. L., 1996. Contribución al conocimiento de los Pyra- midelloidea (Gastropoda: Heterostropha) del Mediterráneo español. Iberus, 14 (1): 1-82. SCHANDER, C., 1994. Twenty-eight new species of Pyramidellidae (Gastropoda, Hetero- branchia) from West Africa. Notiziario CISMA, 17: 11-78. THIELE, J., 1931-35. Handbook of systematic mala- cology. Gustav Fischer. 189 pp. 40 VAUGHT, K. C., 1989. A classification of the living Mollusca. American malacologists. Mel- bourne, Florida. 189 pp. WENZ, W., 1938. Handbuch der Paláozoologie. Von Gebrúder Borntraeger. Berlin. 1638 pp. WISE, J. B., 1996. Morphology and philogene- tic relationship of certain pyramidelid taxa (Heterobranchia). Malacologia, 37(2): 443-551. O Sociedad Española de Malacología —_—_—_—_—_—— Iberus, 15 (1): 41-93, 1997 Fauna malacológica del litoral del Garraf (NE de la Penín- sula Ibérica) Malacological marine fauna from Garraf coast (NE Iberian Penin- sula) Gonzalo GIRIBET* y Anselmo PEÑAS** Recibido el 22-VII-1996. Aceptado el 11-X-1996 RESUMEN Se presenta una lista de 622 especies de moluscos marinos (7 Poliplacóforos, 417 Gaste- rópodos, 190 Bivalvos y 8 Escafópodos) recolectados en el litoral del Garraf (Barcelona, NE de la Península Ibérica). De estas especies, 53 se citan por primera vez en el Medite- rráneo español, siendo dos de ellas primera cita para todo el Mediterráneo, Trophon bar- vicensis y Pleurotomella coeloraphe. De particular interés ha resultado el estudio de un nuevo yacimiento de sedimentos Wúrmienses, asociado a una biocenosis de corales blan- cos, entre 250 y 350 m de profundidad, y el análisis del contenido gástrico de unos tres mil ejemplares de estrellas de mar del género Astropecten, recolectadas entre 40 y 350 m de profundidad. Se incluyen, asimismo, comentarios sobre algunos de los taxones mencio- nados y se ilustran al MEB muchos de ellos, con especial atención a los de las familias Cerithiopsidae, Turridae de profundidad, Yoldiidae y Thyasiridae. ABSTRACT We report a checklist of 622 marine molluses (7 Poliplacophors, 417 Gastropods, 190 Bivalves and 8 Scaphopods) from “El Garraf” coast (Barcelona, NE Iberian Penninsula). From these species, 53 are new findings for the Spanish Mediterranean, and two of them, Trophon barvicensis and Pleurotomella coeloraphe, are reported for the first time for the whole Mediterranean. A new Wúrm bed associated with a white coral biocenosis has been found off Vallcarca at depths between 250 and 350 m, and is described here. Data about molluscs identified from the gut contents of about 3000 specimens of Astropecten sea stars found between 40 and 350 m depth are also reported. Also, we include com- ments about some of the listed taxa and a special SEM image collections, particularly of such groups as Cerithiopsidae, deep-sea Turridae, Yoldiidae and Thyasiridae. PALABRAS CLAVE: Moluscos marinos, Garraf, NE Península Ibérica, Mar Mediterráneo, tanatocenosis Wiiúrmiense, biocenosis de coral blanco, contenidos estomacales de Astropecten. KEY WORDS: Marine molluscs, Garraf, NE Iberian Penninsula, Mediterranean Sea, Wiirm tanatocenosis, white coral biocenosis, Astropecten gut contents. * Departament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Av. Diagonal 645, 08071 Barcelona. ** Carrer Olérdola 39, 08800 Vilanova i la Geltrú, Barcelona. 41 Iberus, 15 (1), 1997 INTRODUCCIÓN El presente trabajo trata sobre los moluscos marinos (exceptuando la Clase Cephalopoda) que se han encon- trado en el litoral de la comarca del Garraf (Barcelona) durante más de una década de recolección y estudio de material. Aunque, aparentemente, la región objeto de estudio no presente ninguna particularidad biogeográfica o física que la delimite desde un punto de vista biológico, la consideramos de especial interés, puesto que en un área relativamente pequeña se encuentran representados buena parte de los ecosis- temas marinos del Mediterráneo. Ésto se refleja en la gran diversidad de especies encontrada en esta zona. Otro factor importante que determina la riqueza faunística del Garraf es la existencia de diferentes tipos de fondos, con numero- sos cañones submarinos, por lo que a sólo 14 km del puerto principal (Vila- nova i la Geltrú) se alcanzan profundi- dades de unos 530 m, mientras que en otras zonas próximas la distancia se tri- plica para llegar a profundidades seme- jantes. Varios autores han estudiado la fauna malacológica marina de esta comarca (SAMA, 1916; HIDALGO, 1917; VILELLA, 1968; Ros, 1975; BALLESTEROS 1977, 1978 Y 1984; ASENSI, 1984), que ha ofrecido una gran riqueza en cuanto a número de especies, pero en ninguno de los casos anteriores se habían muestre- ado todos los hábitats encontrados en este litoral, o al menos no se había hecho de una forma tan extensiva. Ya HIDALGO (1917) citaba para la zona de estudio 307 especies de moluscos (191 Gasterópo- dos, 108 Bivalvos, 4 Escafópodos y 1 Cefalópodo), y SAMA (1916) citaba 319 especies y 81 variedades. ZONA DE ESTUDIO El Garraf es una pequeña comarca litoral situada al sur de Barcelona (Fig. 1), en cuyo interior se encuentra el Parque Natural del Garraf. Los aproxi- madamente 25 km de costa que presenta 42 esta comarca, están comprendidos entre la desembocadura del río Foix (41* 12' N, 1* 40” E) y punta Ginesta (41* 16' N, 1” 57' E). Tres son los municipios litora- les que se encuentran: Sitges, Vilanova i la Geltrú, y Cubelles, incluyendo el primero de ellos las pedanías de Garraf y Vallcarca. La existencia del importante puerto pesquero de Vilanova i la Geltrú, así como la colaboración de algunos de los pescadores de su cofradía, han sido factores decisivos para proporcionarnos gran parte del material en el que está basado este estudio. Las playas de arena fina con pen- dientes poco pronunciadas son domi- nantes en la zona, aunque antes de la construcción generalizada de espigones, había algunas calas de arenas gruesas con pendientes más pronunciadas, como Cala Morisca (en Vallcarca) o Aiguadolc (en Sitges). Los fondos de arena fina son el hábitat típico de algunos Nassariidae, Naticidae y varios bivalvos, sobre todo de las especies de aguas someras de las familias Tellinidae, Pharidae, Donacidae, Veneridae, Mactri- dae, Pandoridae y Thraciidae. También se encuentran gran cantidad de escolle- ras O espigones, que están proliferando por toda la zona, tanto para la creación de puertos deportivos, como para formar playas artificiales. Además son importantes las paredes rocosas de los acantilados calcáreos típicos del macizo del Garraf, que emergen casi verticalmente de fondos arenosos desde profundidades com- prendidas entre 0, 5 y 4 m. Estas rocas son poco ricas desde el punto de vista malacológico, aunque presentan grandes bancos de Mytilus galloprovin- cialis (Linnaeus, 1758), y pueden llegar a abundar especies como Thais haemastoma (Linnaeus, 1767) y, sobre todo por encima de la línea de marea, algunas especies de los géneros Patella, Gibbula o Littorina. La desembocadura de algunas rieras de cauce intermitente son muy intere- santes desde el punto de vista faunís- tico. Este es el caso de la riera de Vila- GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf 41%30' 410 030' 12 Figura 1. Mapa de la zona estudiada. Figure 1. Map of studied area. franca, que desemboca al sur de Sitges, y especialmente la desembocadura del río Foix en Cubelles. Esta última zona está formada por una base de arena fina, casi fangosa, rica en sedimentos orgáni- cos, cubierta casi completamente por cantos rodados y piedras aportadas esporádicamente por el río (suele estar seco por la presencia de una presa de contención situada unos kilómetros más arriba), que sólo lleva agua en forma de grandes avenidas. La profundidad máxima en esta zona de cantos rodados es de aproximadamente 1 m, y a partir de aquí ya se encuentra el típico fondo arenoso que caracteriza a las playas cir- cundantes. La base pedregosa presenta una gran riqueza malacológica, princi- palmente de Opistobranquios. Esta zona concreta de Cubelles ha sido previa- mente estudiada por Ros (1975) y, prin- cipalmente, por BALLESTEROS (1977, 1978 y 1984), y constituye la localidad tipo de Río Foix Barcelona ' SS 1%30' 2 Taringa faba (Ballesteros, Llera y Ortea, 1984). Una buena descripción del recu- brimiento algal así como de la fauna de Invertebrados acompañante se puede encontrar en BALLESTEROS (1984). Situada paralelamente a la costa y a una distancia media de ésta de aproxi- madamente 2, 5 km, se encuentra una pradera de Posidonia oceanica (L.) Dellile. Hace unos 25 años, esta pradera de fanerógamas era muy densa y extensa, encontrándose desde los 8 m de profun- didad (frente a Terramar, en Sitges) hasta 22 m en algunos puntos, y su lon- gitud era de unos 10 km de largo por casi 2 km de anchura. Esta pradera ha sufrido una regresión considerable durante las dos últimas décadas, y con ella muchas de las especies típicas de estos hábitats, como Pinna nobilis Linna- eus, 1758, muy común anteriormente, y que ya prácticamente no se encuentra viva en esta comarca litoral. 43 Iberus, 15 (1), 1997 Los fondos de aguas profundas se dividen en “La Mar de Terra” y “La Mar de Fora”, que están separadas por una serie de rocas dispuestas paralelamente a la costa. “La Mar de Terra” es una pla- nicie fangosa con una profundidad máxima que oscila entre los 74 y los 105 m, y que se halla a una distancia media de la costa de 9,5 km. Aquí destacan algunas zonas rocosas aisladas, varias zonas con gorgonias (Eunicella singularis (Esper) y Leptogorgia sarmentosa (Esper)), en las que abunda el bivalvo Pteria hirundo (Linnaeus, 1758). También es importante en esta zona una amplia extensión de concreciones calcáreas y coralinas situadas una entre Sitges y Vilanova, y otra frente a Vallcarca, for- madas por típicos fondos de maérl, en los que abundan las algas calcáreas Lit- hothamnion calcareum (Pallas) Areschoug y Lithothamniom corallioides Crouan, principalmente. “La Mar de Fora” es más variada y generalmente escarpada. Su profundi- dad va desde los 105 hasta los 1600 m en el canal de Foix. Destacan entre el S y SO de Vilanova una zona de barrancos sub- marinos que convergen hacia una pro- fundidad de unos 500 m, un caladero muy rico en el centro, y unas planicies en cuyos límites, frente a Vallcarca, se en- cuentran los fondos de “El Parrusset”. Éste es un cañón submarino profundo (los pescadores faenan a profundidades entre 200 y 450 m), de fondo rico en nó- dulos de ferromanganeso y que alberga una biocenosis de coral blanco (sensu PÉRES Y PICARD, 1964), asociada a una ta- natocenosis de fauna Wúrmiense de gran interés malacológico. MATERIAL Y MÉTODOS El presente trabajo-está basado en el material recolectado por los autores o proporcionado por pescadores durante más de una década. También se ha revi- sado material de colecciones malacoló- gicas privadas de la zona de estudio. El material ha sido recolectado en fondos arenosos, escolleras, rocas o acantilados mediante inmersión, y si- 44 multaneando con visitas a las diferentes playas de la zona, especialmente des- pués de los temporales. Hay que tener en cuenta que las especies de aguas pro- fundas que aparecen ocasionalmente en las playas o en aguas someras, suelen proceder de los restos arrojados al mar por los barcos pesqueros, o de la lim- pieza de las redes de pesca antes de en- trar en el puerto. La zona de la desembocadura del río Foix en Cubelles se ha muestreado quin- cenalmente durante más de un año (junio de 1992 a agosto de 1993) de una forma exhaustiva, volteando todas las piedras en dos transectos, uno de 15 x 1 m paralelo a la línea de costa y a 0,5 m de profundidad, y otro de 25 x 1 m, per- pendicular a la costa y a una profundi- dad comprendida entre 0 y 1 m. El material de aguas profundas ha sido proporcionado por pescadores de arrastre. Además, en más de 20 ocasio- nes durante el período de muestreo, se ha acompañado a los pescadores con objeto de separar el material por hábi- tats y profundidades in situ, para obtener una información más detallada sobre el mismo. Se ha analizado el contenido estoma- cal de unos 3000 ejemplares de los aste- roideos Astropecten aranciacus (L.) y As- tropecten irregularis (Linck), procedentes de más de 50 arrastres, principalmente de fondos fangosos entre 40 y 350 m de profundidad. Se han estudiado también seis muestras de detrito fangoso (aproxi- madamente 20 kg en total) procedente de “El Parrusset”, entre 200 y 350 m de profundidad, recolectadas entre 1994 y 1996. El detrito ha sido lavado y pasado por una serie de tamices, siendo el más fino de 0,4 mm de luz de malla. Además se han analizado unos 30 kg de sedi- mentos obtenidos en playas de la zona; unos 5 kg de sedimentos arenoso de 2 m de profundidad obtenidos en Sitges; y 1 kg de sedimento arenoso de 2 m de pro- fundidad obtenido en el interior del puerto de Vallcarca. Una gran cantidad de micromoluscos se ha obtenido del es- tudio de estos sedimentos, lo que per- mite obtener un elevado número de es- pecies, aunque de la mayor parte de GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf ellas se hallen sólo conchas (tanatoceno- sis), por lo que no es posible precisar sus hábitats. La mayor parte del trabajo de Opis- tobranquios se ha basado en las publica- ciones de BALLESTEROS (1977, 1978 y 1984), Ros (1975) y ASENSI (1984), y siempre que el material no haya sido recolectado por los autores, se indica la cita bibliográfica de la cual proviene. Además, se han revisado las colec- ciones de A. Tubau, M. Roca y P. Ortoll, malacólogos aficionados o pescadores de Vilanova i la Geltrú. No ha sido posible, sin embargo localizar la colec- ción de SAMA (1916), compuesta por 400 especies de moluscos procedentes del litoral entre Vilanova i la Geltrú (Barce- lona) y Calafell (Tarragona). El material fotografiado al microsco- pio electrónico de barrido (M.E.B.), ha sido previamente hervido en agua desti- lada y tratado con ultrasonidos, con el objeto de eliminar las impurezas deposi- tadas en las conchas, aunque en algunos casos, cuando las conchas eran dema- siado finas, no se ha realizado el trata- miento con ultrasonidos. Las muestras han sido fotografiadas en un MEB Hitachi S-2300 a 15KV. En algunos casos se han seleccionado para fotografiar ejemplares procedentes de otras zonas, con el objeto de ilustrar los ejemplares mejor conservados. El listado de especies ha sido confec- cionado siguiendo a SABELLI, GIANUZZI- SAVELLI Y BEDULLI (1990), excepto para algunos taxones, para los que se han empleado revisiones taxonómicas más recientes. Parte del material aquí tratado ha sido cedido al Museu del Mar de Vila- nova 1 la Geltrú. RESULTADOS El número total de especies de moluscos marinos recogidos en este trabajo es de 622 (7 Poliplacóforos, 417 Gasterópodos, 190 Bivalvos y 8 Escafó- podos), correspondiendo aproximada- mente un 2,7% a especies que se han encontrado exclusivamente en forma subfósil en el Garraf, algunas de las cuales no viven actualmente en el Medi- terráneo, O habitan en zonas más pro- fundas. Lista de especies (Tabla I): A la izquierda aparece el nombre de cada especie, que irá en negrita en el caso de que sea objeto de comentarios en la dis- cusión, irá precedida de un asterisco (*) cuando constituya primera cita en el Mediterráneo español, y de dos (**) cuando constituya primera cita en el Mediterráneo en general. A continua- ción se describe brevemente el tipo de hábitat donde se ha encontrado la especie (lo cual algunas veces no refleja su hábitat real) y el rango batimétrico: “s” (supralitoral), “m” (mesolitoral), “1” (infralitoral: de O a 30 m), “c” (circalito- ral: de 30 a 200 m), y “b” (batial: más de 200 m). En algunos casos no se disponía de estos datos, por lo que no se regis- tran. En el caso de las especies fósiles, tampoco se indica el hábitat. En la siguiente columna se señalan, con un número, las especies ilustradas, indi- cando dicho número el de la figura correspondiente. A continuación se señala la abundancia (+: 1-2 ejemplares, ++: 3-10, +++: 11-100, ++++: más de 100), y se identifican con una “p” a las especies procedentes del detrito de “El Parrusset” y con una “f” a aquellas que han sido halladas fósiles. En el caso de que las letras “p” y “f” aparezcan entre paréntesis, significa que la especie en cuestión no ha sido hallada exclusiva- mente en “El Parrusset” o no ha sido encontrada exclusivamente fósil, respec- tivamente. También se indica con una “y” si la especie ha sido hallada viva en el área de estudio, y con “Aa” o “Ai” se señalan las especies obtenidas en conte- nidos estomacales de Astropecten aran- ciacus O A. irregularis, respectivamente. Evidentemente no ha sido posible espe- cificar todos los ambientes donde se han recolectado las muestras, y es por esto que nos hemos limitado a mencionar la procedencia de aquellas muestras obte- nidas de estas formas particulares. 45 Iberus, 15 (1), 1997 Tabla I. Listado de especies encontradas en el área de estudio, hábitat donde se han encontrado, rango batimétrico, figuras en las que están representadas, abundancia y procedencia. Las especies en negrita están comentadas en el texto. No se incluyen los datos no disponibles de bari- metría, ni de hábitat en las especies fósiles. En las especies de opistobranquios que no han sido reco- lectadas por los autores se incluye la referencia bibliográfica de donde procede la cita. Códigos. *: primera cita en el Mediterráneo español; **: primera cita en el Mediterráneo; s: suprali- toral; m: mesolitoral; i infralitoral (0-30 m); c: circalitoral (30-200 m); b: batial (>200 m); +: 1-2 ejem- plares; ++: 3-10 ejemplares; +++: 11-100 ejemplares; ++++: más de 100 ejemplares; p: especie proce- dente del detrito de El Parrusset; f: especies halladas fósiles; (p): especie hallada no sólo en El Parrus- set; (£): especie hallada no exclusivamente fósil; v: especie encontrada viva en el área de estudio; Aa: especie obtenida en contenido estomacal de Astropecten aranciacus, Ai: idem de Astropecten irregularis. Table I. List of species found in the study area, habitat where they have been collected, bathymetric range, figures when included, abundance and other data related with their collection. Species in bold are discussed in the text. Data on bathymetric range are included only when known, habi- tat of fossil species always excluded. A bibliographic reference is given for the opisthobranch species not collected by the authors. Codes. *: first record in the Spanish Mediterranean; **: first record in the Mediterranean Sea; s: upper littoral; m: midlittoral; i lower littoral (0-30 m); c: circa littoral (30-200 m); b: bathyal (>200 m); +: 1-2 specimens; ++: 3-10 specimens; +++: 11-100 specimens; ++++: more than 100 specimens; p: spe- cies found in El Parrusset detritus; f. species found fossil; (p): species found not only in El Parrusset; (f): species found not only fossil; v: species found alive in the study area; Aa: species collected in Astropecten aranciacus gut contents; Ál: idem in Asttopecten irregularis gut contents. Close POLYPLACOPHORA Familia LEPTOCHITONIDAE Lepidopleurus cajetanus (Poli, 1791): piedras, ¡ + ov Familia ISCHNOCHITONIDAE Callochiton septemvalvis euplaeae (0. G. Costa, 1829): algas (Payssonnelia) y conchas muertas, ic Hov Lepidochitona cinerea (Linnaeus, 1767): piedras y espigones, mi H+ oV Lepidochitona corrugata (Reeve, 1848): piedras y espigones, mi H+ oV Familia CHITONIDAE Chiton olivaceus Spengler, 1797: piedras y espigones, m:i HE oV Familia ACANTHOCHITONIDAE Acanthochitona crinita (Pennant, 1777): piedras, ¡ ++ v A Acanthochitona fascicularis (Linnaeus, 1767): piedras y conchas muertas, ic Hov Clase GASTROPODA Familia PATELLIDAE Potella caerulea Linnaeus, 1758: rocas y espigones, mii HH v Patella rustica Linnaeus, 1758: rocas y espigones, s HH V Patella ulyssiponensis Gmelin, 1791: rocas y espigones, ¡ H+-ov Familia ACMAEIDAE Acmaea virginea (0. E. Múller, 1776): sedimentos en zonas rocosas, i ++ Familia LEPETIDAE lothia fulva (O. F. Múller, 1776): b + pÍ Familia COCCULINIDAE Coccopigya sp.: b + Pp Familia LEPETELLIDAE Lepetella cfr. espinosae Dantart y Luque, 1994: b H+ op Familia ADDISONIIDAE Addisonia excentrica Tiberi, 1857: en capsula ovígeras de Scyliorrhinus, ca Fig.3 ++ (p),v Familia NERITIDAE Smaragdia viridis (Linnaeus, 1758): fango, ic ++ Mo 46 GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Familia FISSURELLIDAE Fissurella nubecula (Linnaeus, 1758): en rocas y piedras, mi Diodora gibberula (Lamarck, 1822): en piedras, ¡ Diodora graeca (Linnaeus, 1758): en rocas y piedras, ¡ Emarginula fissura (Linnaeus, 1758): b Emarginula octaviana Coen, 1939: ic *Emarginula pustula Thiele in Kuester, 1913: b Emarginula rosea T. Bell, 1824: cb Familia SCISSURELLIDAE Anatoma aspera (Philippi, 1844): b Familia HALIOTIDAE Haliotis tuberculata lamellosa Lamarck, 1822: ¡ Familia TROCHIDAE Clanculus cruciatus (Linnaeus, 1758): piedras, ¡ Clanculus jussievi (Payraudeau, 1826): piedras, ¡ Jujubinus exasperatus (Pennant, 1777): ic Jujubinus montagui (W. Wood, 1828): fango, c Jujubinus striatus (Linnaeus, 1758): fango, c Gibbula albida (Gmelin, 1791) Gibbula magus (Linnaeus, 1758): cascajo y múerl, c Gibbula racketti (Payraudeau, 1826): piedras, ¡ Gibbula fanulum (Gmelin, 1791) Gibbula guttadauri (Philippi, 1836) Gibbula leucophaea (Philippi, 1836) Gibbula philberti (Récluz, 1843): piedras y espigones, m Gibbula richardi (Payraudeau, 1826): piedras, m Gibbula varia (Linnaeus, 1758): piedras, m Gibbula divaricata (Linnaeus, 1758): piedras, m Osilinus articulatus Lamarck, 1822: piedras, m Osilinus turbinatus (Bor, 1778): piedras, m Calliostoma conulus (Linnaeus, 1758): rocas y múers, ic Calliostoma dubium (Philippi, 1844) Calliostoma laugieri laugieri (Payraudeau, 1826): i Calliostoma zizyphinum (Linnaeus, 1758) Calliostoma granulatum (Born, 1778): fango, cb Danilia otaviana (Cantraine, 1835): b Familia SKENEIDAE Dikoleps pusilla (Jeffreys, 1847): b *Lissotesta gittenbergeri (van Aortsen y Bogi, 1988): b Familia TURBINIDAE Bolma rugosa (Linnaeus, 1767): rocas y fango, c-b Familia COLLONIIDAE Homalopoma sanguineum (Linnaeus, 1758) Familia TRICOLIIDAE Tricolia pullus (Linnaeus, 1758): ¡ Tricolia speciosa (von Múhlfeldt, 1824): ¡ Tricolia tenvis (Michaud, 1829): Familia CERITHIIDAE Cerithium alucaster (Brocchi, 1814): fango y múerl, i-c Cerithium lividulum Risso, 1826 Cerithium vulgatum Bruguiére, 1792: fango y múerl, i-c Bittium latreillei (Payraudeau, 1826): ix Bittium reticulatum (da Costa, 1778): b Bittium submamillatum (Rayneval y Ponzi, 1854): fango y fondos detríticos, c Fig. 4 + +++ Iberus, 15 (1), 1997 Familia FOSSARIDAE Fossarus ambiguus (Linnaeus, 1758): ¡ “Familia SILIQUARIIDAE Tenagodus obtusus (Schumacher, 1817) Familia TURRITELLIDAE Turritella communis Risso, 1826: fango, c Turritella monterosatoi Kobelt, 1888: fango, c Familia LITTORINIDAE Littorina neritoides (Linnaeus, 1758): rocas y espigones, s Littorina punctata (Gmelin, 1791): rocas y espigones, s Familia SKENEOPSIDAE Skeneopsis planorbis (Fabricius, 1780): arena, ¡ Familia RISSOIDAE Rissoa auriscalpium (Linnaeus, 1758): i Rissoa decorata Philippi, 1846: ¡ *Rissoa gemmula (Fischer in de Folin, 1871): i Figs. 2, 20-21 Rissoa querinii Récluz, 1843: ¡ Rissoa labiosa (Montagu, 1803): ¡ Rissoa lia (Monterosato, 1884 ex Benoit ms.): i Rissoa monodonta Philippi, 1836: ¡ Rissoa similis Scacchi, 1836: ¡ Rissoa ventricosa Desmarest, 1814: ¡ Rissoa violacea Desmarest, 1814: ¡ Alvania beani (Hanley in Thorpe, 1844): ¡ Alvania cancellata (da Costa, 1778): ¡ Alvania cimex (Linnaeus, 1758): i Alvania cimicoides (Forbes, 1844): fango y detrito coralígeno, c-b Alvania discors (Allan, 1818): ¡ Alvania geryonia (Nardo, 1847 ex Chiereghini ms.): ¡ Alvania lactea (Michaud, 1832): ¡ Alvania lineata Risso, 1826: ¡ Alvania punctura (Montagu, 1803): fango y detrito coralígeno, cb Alvania rudis (Philippi, 1844): i Alvania subcrenulata (B. D. D., 1884): ¡ * Alvania subsoluta (Aradas, 1847): b Figs. 8, 11, 12 Alvania testae (Aradas y Maggiore, 1844): fango y detrito coralígeno, c-b Figs. 7, 9, 10 *Alvania zylensis Gotas y Warén, 1982: b Figs. 13, 14 Alvania semistriata (Montagu, 1808): ib Alvania carinata (da Costa, 1778): i *Benthonella tenella (Jeffreys, 1869): fango, c Manzonia crassa (Kanmacher, 1798): i Manzonia zetlandica (Montagu, 1815): b Obtusella intersecta (S. W. Wood, 1857): b Obtusella macilenta (Monterosato, 1880): fango y detrito coralígeno, cb Fig. 6 Pusillina inconspicua (Alder, 1844): ib Figs. 17, 18, 19 Pusillina philippi (Aradas y Maggiore, 1844): +b Figs. 15, 16 Pusillina radiata (Philippi, 1836): ¡ Setia maculata (Monterosato, 1869): ¡ Rissoina bruguierei (Poyraudeau, 1826): ¡ Familia ADEORBIDAE Circulus striatus (Philippi, 1836): ¡ Familia ASSIMINEIDAE Paludinella sicana (Brugnone, 1876): ¡ 48 +++ ++ ++ + +++ ++ +++ +++ Ai (p), v, Ai GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Familia CAECIDAE Caecum auriculatum de Folin, 1868: ¡ Caecum clarkii Carpenter, 1858: ¡ Caecum trachea (Montagu, 1803): ¡ Familia HYDROBIIDAE Ventrosia ventrosa (Montagu, 1803) Familia IRAVADIIDAE Ceratia proxima (Forbes y Hanley, 1850 ex Alder ms.): fango y detrito coralígeno, c-b Hyala vitrea (Montagu, 1803): fango y detrito coralígeno, c-b Familia TORNIDAE Tornus subcarinatus (Montagu, 1803): i Familia TRUNCATELLIDAE Truncatella subcylindrica (Linnaeus, 1767): i Familia APORRHAIDAE Aporrhais pespelecani (Linnaeus, 1758): fango, c Aporrhais serresianus (Michaud, 1828): fango, cb Familia VANIKORIDAE * Tolassia dagueneti (de Folin, 1873): b Familia CALYPTRAEIDAE Colyptraea chinensis (Linnaeus, 1758): en conchas muertas, cb Crepidula fornicata (Linnaeus, 1758) Crepidula unguiformis Lamarck, 1822: en conchas muertas, cb Familia CAPULIDAE Capulus ungaricus (Linnaeus, 1758): sobre conchas, c-b Familia XENOPHORIDAE Xenophora crispa (Koenig, 1825): fango y fondos detríticos, b Familia VERMETIDAE Vermetus triquetrus Bivona, 1832: rocas, ¡ Serpulorbis arenaria (Linnaeus, 1767): rocas, ¡ Familia CYPRAEIDAE Erosaria spurca (Linnaeus, 1758) Luria lurida (Linnaeus, 1758) Zonaria pyrum (Gmelin, 1791): múer, c Familia OVULIDAE Aperiovula adriatica (G. B. Sowerby |, 1828): c Neosimnia spelta (Linnaeus, 1758): sobre Eunicella, c Pseudosimnia camea (Poiret, 1789): c Familia LAMELLARIIDAE Lamellaria latens (0. F. Múller, 1776): ¡ Familia TRIVIIDAE Trivia arctica (Pulteney, 1789): ¡ Trivia monacha (da Costa, 1778): ¡ Trivia multilirata (G. B. Sowerby 11, 1870) Erato voluta (Montagu, 1803) Familia NATICIDAE Naticarius cruentatus (Martyn, 1784): arena y fango, ic Naticarius dillwyni (Payraudeau, 1826): arena, i Naticarius punctatus (Chemnitz in Karsten, 1789) Naticarivs vittatus (Gmelin, 1791) Tectonatica filosa (Philippi, 1844): fango, c Lunatia fusca (Blainville, 1825): fango, c-b Lunatia guillemini (Poyraudeau, 1826): fango, ic Lunatia macilenta (Philippi, 1844): arena y fango, ic Lunatia nitida (Donovan, 1804): arena y fango, i-c Payraudeautia intricata (Donovan, 1804): fango y múerl, i-c (p), v, Ai (p), v Ai ==" MES 49 Iberus, 15 (1), 1997 Familia TONNIDAE Tonna galea (Linnaeus, 1758) + Familia CASSIDAE Galeodea echinophora (Linnaeus, 1758): fango, c HE 0V Galeodea rugosa (Linnaeus, 1771): fango, cb ++ (p),v Phalium granulatum (Born, 1778): fango, c H+ 0V Phalium saburon (Bruguiére, 1792): fango, c ++ Y Familia RANELLIDAE Ranella olearia (Linnaeus, 1758) + pf Cymatium corrugatum (Lamarck, 1816): fango, c HH V Cymatium parthenopeum parthenopeum (von Salis, 1793): rocas, ¡ + Y Cobestana cutacea cutacea (Linnaeus, 1767): fango y rocas, ic ++ v Charonia lampas lampas (Linnaeus, 1758) + Familia ATLANTIDAE Atlanta peronii Lesueur, 1817: fango y detrito coralígeno, b ++ (p), Ai Oxygyrus keraudrenii (Lesueur, 1817): b Hop Familia TRIPHORIDAE Marshallora adversa (Montagu, 1803): fondos detríticos, ¡ ++ vA Monophorus erythrosomus (Bouchet y Guillemot, 1978): ¡ ++ Monophorus perversus (Linnaeus, 1758): ¡ ++ * Obesula marinostri Bouchet, 1985: b + op Similiphora similior (Bouchet y Guillemot, 1978): i + Metaxia metaxae (delle Chiaje, 1828): ¡-b ++ (p), (f) Familia CERITHIOPSIDAE * Cerithiopsis diadema Monterosato, 1874 ex Watson ms.: b Fig. 22 + Pp Cerithiopsis jeffreysi Watson, 1885: b Fig.23 ++ (p), (1) Cerithiopsis minima (Brusina, 1865): i Fig. 24 +++ Cerithiopsis nana Jeffreys, 1867: ib Figs. 25,29 ++ (p), (1 Cerithiopsis scalaris (Monterosato, 1877): b Fig. 26 +++ (p), (f) * Cerithiopsis tiara Monterosato, 1874 ex Watson ms.: b Fig. 27 + (p), (1) Cerithiopsis tubercularis (Montagu, 1803): +b Figs. 28,30 +++ (p), (f) Familia ACLIDIDAE *Adlis attenuans Jeffreys, 1883: fango y detrito coralígeno, b ++ (p), v Ai Aclis gulsonae (W. Clark, 1850): fango y detrito coralígeno, b + pvA Cima minima (Jeffreys, 1858): b ++ (p) * Cioniscus gracilis Monterosato, 1874, ex Jeffreys ms.: b ++ (p) Graphis albida (Kanmacher, 1798): ib Figs. 32, 33,34 ++ (p) Familia EPITONIDAE * Epitonium aculeatum (Allan, 1818): fango y detrito coralígeno, b ++ (p),v Ai Epitonium olgerianum (Weinkauff, 1866): b ++ (p) *Epitonium celesti (Aradas, 1854): b + p Epitonium clathratulum (Kanmacher, 1798): b Fig.39 ++ (p) Epitonium commune (Lamarck, 1822): rocas y fango, i-c +++ v A *Epitonium dendrophylliae Bouchet y Warén, 1986: b 509 * Epitonium hispidulum (Monterosato, 1874): b ++ (p) * Epitonium linctum (de Boury y Monterosato, 1890): b Fig. 40 + p Epitonium pulchellum (Bivona, 1832): ¡ ++ Epitonium turtonis (Turton, 1819): fango, c +++ vw A Cirsotrema cochlea (6. B. Sowerby Il, 1844): ¡ + Gyroscala lamellosa (Lamarck, 1822): rocas y arena, i st *Opalia abboti Clench y Turner, 1952: b 5 Opalia crenata (Linnaeus, 1758): i ++ Opalia hellenica (Forbes, 1844): b 0d) 50 GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Familia EULIMIDAE Eulima bilineata Alder, 1848: fango y detrito coralígeno, b Eulima glabra (da Costa, 1778): ¡ *Crinophtheiros sp.: fango, b * Entoconcha mirabilis Múller, 1852: b Melanella alba (da Costa, 1778): i Melanella boscii (Payraudeau, 1827): ¡ Melanella praecurta (Pallary, 1904): fango, b Parvioris ibizenca (Nordsieck, 1968): ¡ Sticteulima jeffreysiana (Brusina, 1869): detrito coralígeno, b Vitreolina perminima (Jeffreys, 1883): detrito coralígeno, b Vitreolina sp.: detrito coralígeno, b Familia MURICIDAE Bolinus brandaris (Linnaeus, 1758): fango, arena y piedras, ¡ Hadriania craticuloides (Vokes, 1964): fango y detrito coralígeno, c-b Hexaplex trunculus (Linnaeus, 1758): fango, rocas y espigones, ¡ Murexul aradasii (Poirier, 1883 ex Monterosato ms.) Muricopsis cristatus (Brocchi, 1814): rocas y detritos, ic Ocenebra erinaceus (Linnaeus, 1758): rocas y detritos, ix Ocinebrina aciculata (Lamarck, 1822): fango y piedras, ix Ocinebrina edwardsi (Payraudeau, 1826): rocas y piedras, i *Trophon barvicensis (Johnston, 1825) Trophon echinatus (Kiener, 1840): b Trophon sp.: b Irophon muricatus (Montagu, 1803): fango y detrito coralígeno, b Familia BUCCINIDAE Buccinum humphreysianum Bennet, 1824: b Buccinum undatum Linnaeus, 1758: fango, c Buccinulum cormeum (Linnaeus, 1758): fango y rocas, ic Chauvetia brunnea (Donovan, 1804): ¡ Chauvetia turritellata (Deshayes, 1835): i Colus jeffreysianus (Fischer, 1868): i Neptunea contraria (Linnaeus, 1771) Pisania striata (Gmelin, 1791) Contharus dorbignyi (Payraudeau, 1826): i Familia CORALLIOPHILIDAE Coralliophila meyendorffi (Calcara, 1845): piedras y fango, i Coralliophila panormitana (Monterosato, 1869): b Coralliophila squamosa (Bivona, 1831): fango, cb Familia FASCIOLARIIDAE Fusinus pulchellus (Philippi, 1844): fango, ¡ Fusinus rostratus (Olivi, 1792): fango, c Fusinus rudis (Linnaeus, 1758): fango, i Familia NASSARIIDAE Nassarius corniculus (Olivi, 1792): piedras, ¡ Nassarius cuvierii (Payraudeau, 1826): arena, ¡ Nassarius incrassatus (Stróm, 1768): piedras, ¡ Nassarius mutabilis (Linnaeus, 1758): arena y fango, i Nassarius nitidus (Jeffreys, 1867): i Nassarius pygmaeus (Lamarck, 1822): fango y piedras, ix Nassarius reticulatus (Linnaeus, 1758): arena, i Nassarius unifasciatus (Kiener, 1835) Naytiopsis granum (Lamarck, 1822): arena, ¡ Cyclope neritea (Linnaeus, 1758) Figs. 37, 38 ++ Figs. 35,36 +++ + Fig. 45 + Figs. 41,42. ++ Figs. 43,44 ++ Figs. 46,47 +++ (p), y Ai 51 Iberus, 15 (1), 1997 Familia THAIDIDAE Orania fusulus (Brocchi, 1814) Stramonita haemastoma (Linnaeus, 1766): rocas, ¡ Familia COLUMBELLIDAE Columbella rustica (Linnaeus, 1758): rocos y algas, i Mitrella minor (Scacchi, 1836) Mitrella scripta (Linnaeus, 1758) Familia COSTELLARIIDAE Vexillum ebenus (Lomarck, 1811): ¡ Vexillum tricolor (Gmelin, 1790) Familia MARGINELLIDAE Gibberula caelata (Monterosato, 1877) bibberula miliaria (Linnaeus, 1758): arena, ¡ Gibberula philippii (Monterosato, 1877): ¡ Gibberulo turgidula (Locard y Caziot, 1900): fango, e Volvarina mitrella (Risso, 1826) Granulina clandestina (Brocchi, 1814): b Familia MITRIDAE Mitra zonata Marryot, 1818: fango y múerl, ic Familia CANCELLARIIDAE Concellaria cancellata (Linnaeus, 1767): arena y fango, ic Cancellaria similis Sowerby, 1833: fango, cb Familia CONIDAE Conus ventricosus (Gmelin, 1791): i Familia TURRIDAE Bela brachistoma (Philippi, 1844): fango y detrito coralígeno, c-b Belo laevigata (Philippi, 1836): arena, i Bela menkhorsti van Aartsen, 1988: b Bela nebula (Montagu, 1803): arena fangosa, i Bela ornata (Locard, 1897): fango, ic Bela zonata (Locard, 1892): fango, c Mongelia attenuata (Montagu, 1803): fango y detrito coralígeno, b Mangelia costata (Donovan, 1804): fango y detrito coralígeno, cb Mangelia cfr. goodalii Reeve, 1846: i Mangelia nuperrima (Tiberi, 1855): fango y detrito coralígeno, cb Mangelia paciniana (Calcara, 1839): arena, ¡ Mangelia serga (Dall, 1881): fango y detrito coralígeno, c-b Mangelia smithi (Forbes, 1844): arena, fango y detrito coralígeno, cb Mangelia stossiciana (Brusina, 1869): ¡ Mangelia unifasciata Deshayes, 1835: arena y fango, ¡ Mangelia vauquelini (Payraudeau, 1826): orena, ¡ Mangiliella bertrandii (Poyraudeau, 1826): ¡ Mangiliella taeniata (Deshayes, 1835): ¡ Taranis moerchi (Malm, 1861): detrito coralígeno, b Taranis sp.: detrito coralígeno, b Microdrilia loprestiana (Calcara, 1841): fongo y detrito coralígeno, cb Haedropleura septangularis (Montagu, 1803): en Posidonia, i **Pleurotomella coeloraphe (Dautzenberg y Fischer, 1896): b *Pleurotomella demosia (Duutzenberg y Fischer, 1896): b Crassopleura maravignae Bivona, 1838: ic Mitrolumna olivoidea (Cantraine, 1835): ¡ Raphitoma aequalis Jeffreys, 1867: fango, c- Raphitoma bicolor (Risso, 1826): arena, ¡ Raphitoma concinna (Scacchi, 1836): arena, ¡ 2 Figs. 50, 51 Figs. 52, 53 Figs. 54, 55 Figs. 57, 58, 59 Figs. 60, 61, 62 Fig. 56 Figs. 63, 64, 65 Figs. 66, 67, 68 ++ (p), Ai GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf *Raphitoma cordieri (Payraudeau, 1826): ¡ Raphitoma echinata (Brocchi, 1814): i Raphitoma horrida Monterosato, 1844: en Posidonia, ¡ Raphitoma leufroyi (Michaud, 1828): arena, ¡ Raphitoma linearis (Montagu, 1803): arena, ¡ Raphitoma cfr. nivea (Marshall in Sykes, 1906): i *Raphitoma pupoides (Monterosato, 1884): i Comarmondia gracilis (Montagu, 1803): fango y arena, i-c Teretia teres (Forbes, 1844): fango y detrito coralígeno, b Familia TJAERNOEIDAE Tjaernoeia exquisita (Jeffreys, 1883): b Familia ARCHITECTONICIDAE Basisulcata lepida (Bayer, 1942): mier, c Heliacus alleryi (Seguenza, 1876): b Heliacus architae (0. 6. Costa, 1867): b Familia MATHILDIDAE Mathilda cochlaeformis Brugnone, 1873: b Familia OMALOGYRIDAE Omalogyra atomus (Philippi, 1841): ¡ Ammonicera fischeriana (Monterosato, 1869): i Familia PYRAMIDELLIDAE Tiberia minuscula (Monterosato, 1880): fango y detrito coralígeno, c- Chrysallida brattstroemi Warén, 1991: b Chrysallida brusinaí (Cossmann, 1921): i Chrysallida dollfusí (Kobelt, 1903): b Chrysallida emaciata (Brusina, 1866): ¡ Chrysallida excavata (Philippi, 1836): ¡ Chrysallida fenestrata (Jrffreys, 1848): c Chrysallida flexuosa (Monterosato, 1874 ex Jeffreys): fango y detrito coralígeno, cb Chrysallida ghisottii van Aartsen, 1984: i Chrysallida indistincta (Montagu, 1808): ¡ Chrysallida intermixta (Monterosato, 1884) Chrysallida interstincta (J. Adams, 1797): Chrysallida juliae (de Folin, 1872): fango, c Chrysallida palazziiMicali, 1984: fango y detrito coralígeno, cb Chrysallida pellucida (Dillwyn, 1817) Chrysallida suturalis (Philippi, 1844): fango y detrito coralígeno, c-b Odostomella doliolum (Philippi, 1844): ib Euparthenia bulinea (Lowe, 1841): arena, ¡ Euparthenia humboldti (Risso, 1826) Eulimella acicula (Philippi, 1836) Eulimella ataktos Warén, 1991: fango y detrito coralígeno, cb Eulimella bogii van Aartsen, 1994: b Eulimella scillae (Scacchi, 1835): fango y detrito coralígeno, cb Eulimella unifasciata (Forbes, 1844): fango y detrito coralígeno, Eulimella ventricosa (Forbes, 1844): fango y detrito coralígeno, c-b Puposyrnola minuta (H. Adams, 1869): fango y detrito coralígeno, b Odostomia acuta Jeffreys, 1848: fango y arena, ic Odostomia afzelii (Warén, 1991): fango y detrito coralígeno, c-b Odostomia carrozzai van Aartsen, 1987: ¡ Odostomia clavulus (Lovén, 1846): fango y detrito coralígeno, cb Odostomia conoidea (Brocchi, 1814): fango y detrito coralígeno, ib Odostomia erjaveciana Brusina, 1869: arena, ¡ Odostomia eulimoides Hanley, 1844: ¡ Fig. 69 +++ p, Y, Ai (p), v Ai 53 Iberus, 15 (1), 1997 Odostomia hansgei (Warén, 1991): fango y detrito coralígeno, b Odostomia kromi van Aartsen, Menkhorst y Gittenberger, 1984: ¡ Odostomia lukisii Jeffreys, 1859: i Odostomia megerlei (Locard, 1886) Odostomia plicata (Montagu, 1803): ¡ Odostomia scalaris MacGillivray, 1843: ¡ Odostomia striolata Forbes y Hanley, 1850: ib Odostomia suboblonga Jeffreys, 1884: b Odostomia turriculata Monterosato, 1869: ¡ Odostomia turrita Honley, 1844: ib Odostomia umbilicaris Malm, 1863: fango y detrito coralígeno, c-b Odostomia unidentata (Montagu, 1803): fango y detrito coralígeno, ib Odostomia verduini van Aartsen, 1987: ¡ Noemiamea dolioliformis (Jeffreys, 1848): ¡ Ondina dilucida (Monterosato, 1844): fango, c Ondina obliqua (Alder, 1844): ¡ Turbonilla acuta (Donovan, 1804): ¡ Turbonilla acutissima Monterosato, 1884: b Turbonilla jeffreysii (Jeffreys, 1848): i Turbonilla pusilla (Philippi, 1844) Turbonilla rufa (Philippi, 1836): ¡ Turbonilla sinvosa (Jeffreys, 1884): ¡ Turbonilla striatula (Linnaeus, 1758) Ebala nitidissima (Montagu, 1803) Ebala pointeli (de Folin, 1868): i Ebala sp.: b Familia ACTEONIDAE Acteon tornatilis (Linnaeus, 1758): arena y fango, ic Crenilabrum exilis (Forbes in Jeffreys, 1870): b Familia DIAPHANIDAE Diaphana minuta Brown, 1827: b Familia RETUSIDAE Retusa semisulcata (Philippi, 1836): ¡ Retusa truncatula (Bruguiére, 1792): i Cylichnina umbilicata (Montagu, 1803): fango, ic Familia RINGICULIDAE Ringicula auriculata (Ménard de la Groye, 1811): fango, c *Ringicula cfr. leptocheila Brugnone, 1873: b Familia BULLIDAE Bulla striata Bruguiére, 1792: arena, ¡ Familia HAMINAEIDAE Haminaea hydatis (Linnaeus, 1758): i Haminaea orbignyana (Férussac, 1822): ¡ Weinkauffia turgidula (Forbes, 1844): b Familia PHILINIDAE Philine aperta (Linnaeus, 1767): ¡ Philine catena (Montagu, 1803): i Philine scabra (0. F. Miller, 1776): ib *Lgona pruinosa (Clark, 1827): fango y detrito coralígeno, cb Familia SCAPHANDRIDAE Cylichna cylindracea (Pennant, 1777): fango y detrito coralígeno, cb Roxania utriculus (Brocchi, 1814): fango, c Scaphander lignarivs (Linnaeus, 1758): fango, c Scaphander punctostriatus (Mighels y Adams, 1841): b 54 Figs. 80,81 ++ Figs. 78, 79 + Fig. 82. +++ Ai Ai A GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Familia CAVOLINIIDAE Cavolinia inflexa (Lesueur, 1813) Clio cuspidata (Bosc, 1802) Clio pyramidata Linnaeus, 1767 Creseis acicula Rang, 1828 Styliola subula (Quoy y Gaimard, 1827) Familia LIMACINIDAE * Limacina bulimoides (d'Orbigny, 1836): b Limacina inflata (d'Orbigny, 1836): b *Limacina retroversa (Fleming, 1822): b Familia PERACLIDAE Peracle reticulata (d'Orbigny, 1836): b Familia ELYSIIDAE Elysia viridis (Montagu, 1810) Familia HERMAEIDAE Stiliger sp. Familia UMBRACULIDAE Umbraculum mediterraneum (Lamarck, 1819): fango, c Familia APLYSIIDAE Aplysia depilans Gmelin, 1791 Apiysia fasciata Poiret, 1789: arena, i Aplysia punctata Cuvier, 1803: arena, ¡ Familia TRITONIIDAE Tritonia hombergi Cuvier, 1803 Familia DOTIDAE Doto koenneckeri Lemche, 1976 Familia TRIOPHIDAE Kaloplocamus ramosus (Cantraine, 1835) Familia POLYCERIDAE Polycera quadrilineata (0. E. Miller, 1776): piedras, i Familia DORIDIDAE Doris verrucosa Linnaeus, 1758: detritos y piedras, ¡ Familia ARCHIDORIDIDAE Archidoris tuberculata (Cuvier, 1804) Familia DISCODORIDIDAE Taringa faba (Ballesteros, Llera y Ortea, 1984): bajo piedras, zona detrítica, ¡ Familia CENTRODORIDIDAE Jorunna tomentosa (Cuvier, 1804) Familia DENDRODORIDIDAE Dendrodoris grandiflora (Rapp, 1827): bajo piedras, zona detrítica, ¡ Doriopsilla areolata Bergh, 1880 Familia ARMINIDAE Armina maculata Rafinesque, 1814 Familia FLABELLINIDAE Colmella cavolini (Verany, 1846) Coryphella pedata (Montagu, 1822) Familia TERGIPEDIDAE lergipes tergipes (Forskal, 1775) Familia EUBRANCHIDAE Eubranchus exiguus (Alder y Hancock, 1848) Eubranchus farrani (Alder y Hancock, 1844) Familia FACELINIDAE Cratena peregrina Gmelin, 1791 Facelina coronata (Forbes y Goodsir, 1839) Facelina drummondi (Thompson, 1844) Facelina sp. + + + DUDO OO ++ | + op Fig. 83 ++++ p Figs. 84, 85 + p Ballesteros (1984) Ballesteros (1984) + Y Ros (1975) ++ V ++ V Ros (1975) Ballesteros (1984) Ros (1975) H ov Ros (1975) ++ V Ballesteros (1984) ++ V Asensi (1984) Ballesteros (1981) Ballesteros (1978) Ballesteros (1984) Asensi (1984) Asensi (1984) Ballesteros (1984) Ballesteros (1978) Ballesteros (1984) Ballesteros (1984) Ballesteros (1984) DS) Iberus, 15 (1), 1997 Familia FAVORINIDAE Favorinus branchialis (Rathke, 1806): bajo piedras, zona detrítica, ¡ Favorinus vitreus (Ortea, 1982): bajo piedras, zona detrítica, ¡ + ov Familia AEOLIDIDAE Aeolidiella alderi (Cocks, 1852) Ros (1975) Berghia verrucicornis (0. G. Costa, 1864): bajo piedras, zona detrítica, ¡ H+ oV Spurilla neapolitana (delle Chiaje, 1841): bajo piedras, zona detrítica, ¡ HH oV Familia SIPHONARIIDAE Williamia gussonii (O. 6. Costa, 1829): zonas detríticas, ¡ ++ Familia TRIMUSCULIDAE Irimusculus mammilaris (Linnaeus, 1758): i ++ Familia ELLOBIIDAE Auriculinella erosa (Jeffreys, 1829) + Ovatella firminii (Payraudeau, 1826) ++ Ovatella myosotis (Draparnaud, 1801) ++ Close BIVALVIA Familia NUCULIDAE Nucula hanleyi Winckworth, 1930: fango, c +++ v A Nucula nitidosa Winckworth, 1930: fango y detrito coralígeno, c-b +++ (p), v, Ao Nucula cfr. nucleus (Linnaeus, 1758): fango, c +++ v A Nucula sulcata Bronn, 183: fango y detrito coralígeno, cb +++ (p), y Ao * Ennucula aegeensis (Forbes, 1844): fango y detrito coralígeno, c-b ++ (p),v Ai Familia NUCULANIDAE Nuculana commutata (Philippi, 1844): fango y detrito coralígeno, cb + lp), Ai Nuculana pella (Linnaeus, 1767): fango, c ++ Ao Familia YOLDIIDAE * Yoldiella lucida (Lovén, 1846): b Fig.86 ++ pA *Yoldiella nana (M. Sars, 1865): fango y detrito coralígeno, b Fig.87. ++ (p), Ai *Yoldiella philippiana (Nyst, 1845): fango y detrito coralígeno, b Figs. 8893 — +++ (p), v Ai Familia ARCIDAE Arca noae Linnaeus, 1758: rocas, ¡ H ov Arca tetragona Poli, 1795 + Barbatia barbata (Linnaeus, 1758): rocas, i + oV Barbatia clathrata (Defrance, 1816): b Hop Anadara diluvii (Lamarck, 1819): fango, c H+ oV Bathyarca pectunculoides (Scacchi, 1834): b Fig. 94 +++ (p),v Bathyarca philippiana (Nyst, 1848): b Fig. 95 +++ (p) Familia NOETIIDAE Striarca lactea (Linnaeus, 1758): piedras, i-c Sa 0 Familia GLYCYMERIDAE Glycymeris glycymeris (Linnaeus, 1758): fango, c: Hov Glycymeris insubrica (Brocchi, 1814): arena, i H+ V Familia MYTILIDAE Myrilus galloprovincialis Lamarck, 1819: rocas y espigones, i HH V Mytilaster minimus (Poli, 1795): rocas, m HH V *Crenella pellucida (Jeffreys, 1850): detrito coralígeno, b Fig.96 ++ p,v Gregariella subclavata (Libassi, 1859): rocas, ¡ TV Gregariella petagnae (Scacchi, 1832): rocas, i HE v Musculus costulatus (Risso, 1826) +. Musculus subpictus (Cantraine, 1835): fango, c +H ov Lithophaga lifhophaga (Linnaeus, 1758): en piedras, ic H ov Modiolus adriaticus Lamarck, 1819 + Modiolus barbatus (Linnaeus, 1758): rocas, i +++ 56 GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf *Idas cfr. ghisottii Warén y Carrozza, 1990: madera *Idas simpsoni (Marshall, 1900): esqueletos de peces y cetáceos, ix Modiolula phaseolina (Philippi, 1844): detrito coralígeno, b Familia PINNIDAE Atrina pectinata (Linnaeus, 1758): fango, c Pinna nobilis Linnaeus, 1758: praderas de Posidonia, ¡ Familia PTERIIDAE Pteria hirundo (Linnaeus, 1758): gorgonias y restos de redes, c Familia PECTINIDAE Palliolum incomparabile (Risso, 1826) Delectopecten vitreus (Gmelin, 1791): b Pseudamussium septemradiatum (0. F. Múller, 1776): fango, c Peplum clavatum (Poli, 1795): fango, c Karnekampia bruei (Payraudeau, 1826): b Manupecten pesfelis (Linnaeus, 1758): cb Chlamys islandica (0. E. Múller, 1776) Chlamys multistriata (Poli, 1795): fango, ic Chlamys varia (Linnaeus, 1758): roca y fango, i Lissopecten hyalinum (Poli, 1795) Flexopecten flexuosus (Poli, 1795): arena y fango, ic Flexopecten glaber (Linnaeus, 1758) Aequipecten opercularis (Linnaeus, 1758): fango, c Perapecten commutatus (Monterosato, 1875): fango, c Pecten jacobaeus (Linnaeus, 1758): arena y fango, i-c Similipecten similis (Loskey, 1811): fango y detrito coralígeno, c-b *Propeamussium lucidum (Jeffreys in Thompson, 1873): detrito coralígeno, b Propeamussium fenestratum (Forbes, 1844): detrito coralígeno, b *(yclopecten hoskynsi (Forbes, 1844): detrito coralígeno, b Familia SPONDYLIDAE Spondylus gaederopus Linnaeus, 1758 Familia ANOMIIDAE Anomia ephippium Linnaeus, 1758: conchas de moluscos *Heteranomia squamula (Linnaeus, 1758): b Monia patelliformis (Linnaeus, 1761) Familia LIMIDAE Limaria hians (Gmelin, 1791): rocas y espigones, ¡ Limaria inflata Link, 1807: ¡ Notolimea crassa (Forbes, 1844): detrito coralígeno, b Limatula subauriculata (Montagu, 1808): detrito coralígeno, b *Limatula cfr. gwyni (Sykes, 1903): detrito coralígeno, b Familia OSTREIDAE Ostrea edulis Linnaeus, 1758: rocas, ¡ Fomilia GRYPHAEIDAE Neopycnodonte cochlear (Poli, 1795): cb Familia LUCINIDAE Ctena decussata (0. 6. Costa, 1829): arena, i Loripes lacteus (Linnaeus, 1758): arena, ¡ Lucinella divaricata (Linnaeus, 1758): arena, ¡ Lucinoma borealis (Linnaeus, 1767): fango y detrito coralígeno. cb Myrtea spinifera (Montagu, 1803): ¡ Familia THYASIRIDAE Thyasira (Thyasira) biplicata (Philippi, 1836): b *Thyasira (Thyasira) obsoleta (Verrill y Bush, 1898): b *Thyasira (Parathyasira) granulosa (Mont., 1874 ex Jeffreys): b Fig. 97 Fig. 98 Fig. 99 Figs. 101, 102 + +H ++ ++ p, (1), v a. Iberus, 15 (1), 1997 *Thyasira (Parathyasira) subovata (Jeffreys, 1881): detrito coralígeno, b *Thyasira (Leptaxinus) incrassata (Jeffreys, 1876): b *Thyasira (Axinulus) croulinensis (Jeffreys, 1847): b *Thyasira (Axinulus) eumyaria (M. Sars, 1870): b Thyasira (Mendicula) ferruginea (Locard, 1886): fango y detrito coralígeno, b Familia CHAMIDAE Chama gryphoides Linnaeus, 1758: en piedras o Microccosmus, ¡ Pseudochama gryphina (Lamarck, 1819): rocas, i Familia ERYCINIDAE Scacchia ovata Philippi, 1844: ¡ Familia KELLIIDAE Bornia sebetia (0. 6. Costa, 1829): ¡ Kellia suborbicularis (Montagu, 1803): b Familia LASAEIDAE Hemilepton nitidum (Turton, 1822): b Familia MONTACUTIDAE * Mancikellia pumila (Sowerby, 1846): b * Montacuta phascolionis Dautzenberg y Fischer, 1925: b Montacuta substriata (Montagu, 1808): b Mysella bidentata (Montagu, 1803): i Mysella obliguata (Chaster, 1897) Tellimya ferruginosa (Montagu, 1808): b Epilepton clarkiae (Clark, 1852): b Epilepton sp.: detrito coralígeno, b Familia NEOLEPTONIDAE *Arculus sp.: detrito coralígeno, b Familia CARDITIDAE Venericardia antiquata (Linnaeus, 1758): fango, c Glans aculeata (Poli, 1795): fango y detrito coralígeno, cb Glans trapezia (Linnaeus, 1758): fango y piedras, c Familia ASTARTIDAE Astarte fusca (Poli, 1795): fango, c Astarte sulcata (da Costa, 1778): fango, c Goodallia triamgularis (Montagu, 1803): i. b Goodallia sp.: detrito coralígeno, b Familia CARDIIDAE Acanthocardia aculeata (Linnaeus, 1758): arena, Acanthocardia echinata (Linnaeus, 1758): fango, c Acanthocardia paucicostata (Sowerby, 1834): fango, ic Acanthocardia tuberculata (Linnaeus, 1758): arena, ¡ Parvicardium exiguum (Gmelin, 1791): ¡ Parvicardium minimum (Philippi, 1836). fango y detrito coralígeno, cb Parvicardium ovale (6. B. Sowerby, 1840): i Parvicardium roseum (Lamarck, 1819): fango, c Plagiocardium papillosum (Poli, 1795): arena, i Laevicardium crassum (Gmelin, 1791): fango, c Loevicardium oblongum (Chemnitz, 1782): fango, cb Cerastoderma glaucum (Poiret, 1789): arena fangosa, ¡ Familia MACTRIDAE Mactra glauca (Bor, 1778): arena, ¡ Mactra stultorum (Linnaeus, 1758): arena, ¡ Spisula subtruncata (da Costa, 1778): arena, ¡ Lutraria angustior Philippi, 1844 Lutraria lutraria (Linnaeus, 1758): ic Lutraria magna (da Costa, 1778): Posidonia y fango, ix 58 Fig. 100 Fig. 103 Fig. 104 Fig. 105 Fig. 108 GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Familia MESODESMATIDAE Donacilla corea (Poli, 1795) Ervilia castanea (Montagu, 1803): fango, c Familia SOLENIDAE Solen marginatus Pulteney, 1799: fango, i Familia PHARIDAE Ensis ensis (Linnaeus, 1758): arena, ¡ Ensis minor (Chenu, 1843): arena, ¡ Pharus legumen (Linnaeus, 1758): arena, ¡ Familia TELLINIDAE Arcopagia balaustina (Linnaeus, 1758): fango, cb Arcopagia crassa (Pennant, 1777): fango y rocas, ic Gastrana fragilis (Linnaeus, 1758) Macoma cumana (0. 6. Costa, 1829): arena, ¡ Tellina donacina Linnaeus, 1758: arena y fango, ic Tellina incarnata Linnaeus, 1758: areno, ¡ Tellina nitida Poli, 1791: arena, ¡ Tellina planata Linnaeus, 1758: arena, i Tellina pulchella Lamarck, 1818: arena y fango, ic Tellina serrata Brocchi, 1814: fango, c-b Tellina tenvis da Costa, 1778: arena, ¡ Familia DONACIDAE Donax semistriatus Poli, 1795: arena, i Donax trunculus Linnaeus, 1758: arena, ¡ Familia PSAMMOBIIDAE Gari fervensis (Gmelin, 1791) Familia SCROBICULARIIDAE Serobicularia coftardi (Payraudeau, 1826) Familia SEMELIDAE Abra longicallus (Scacchi, 1834): detrito coralígeno, b Familia SOLECURTIDAE Solecurtus scopula (Turton, 1822): fango, c Solecurtus strigilatus (Linnaeus, 1758): fango, c Azorinus chamasolen (da Costa, 1778): fango, c Familia ARCTICIDAE Arctica islandica (Linnaeus, 1767) Familia KELLIELLIDAE Kelliella abyssicola (Forbes, 1844): fango, cb Familia TRAPEZIIDAE Coralliophaga lithophagella (Lamarck, 1819): rocas, c Familia GLOSSIDAE Glossus humanus (Linnaeus, 1758): fango, c Familia VENERIDAE Callista chione (Linnaeus, 1758): arena, i Chamelea gallina (Linnaeus, 1758): arena, ¡ Clausinella fasciata (da Costa, 1778): fango, Dosinia exoleta (Linnaeus, 1758): fango, c Dosinia lupinus (Linnaeus, 1758): arena y fango, ic Globivenus effosa (Bivona, 1836) Gouldia minima (Montagu, 1803): b lrus irus (Linnaeus, 1758): interior de piedras calcáreas, mi Paphia aurea (Gmelin, 1791): i Paphia rhomboides (Pennant, 1777): fango, ic Pitar mediterranea Tiberi, 1855: b ++ v Al 59 Iberus, 15 (1), 1997 Pitar rudis (Poli, 1795): fango, cb Tapes decussatus (Linnaeus, 1758): arena fangosa, ¡ Timoclea ovata (Pennant, 1777): fango, cb Venerupis corrugata (Gmelin, 1791): arena, i Venus casina Linnaeus, 1758: fango, c Venus nux Gmelin, 1791 Venus verrucosa Linnaeus, 1758: fango, c Familia PETRICOLIDAE Petricola lifhophaga (Retzivs, 1786): interior de piedras calcáreas, mii Mysia undata (Pennant, 1777): fango, ic Familia CORBULIDAE Corbula gibba (Olivi, 1792): fango y detrito coralígeno, c-b Lentidium mediterraneum (0. 6. Costa, 1829): arena, i Familia GASTROCHAENIDAE Gastrochaena dubia (Pennant, 1777): interior de piedras calcáreas, ¡ Familia HIATELLIDAE Hiatella arctica (Linnaeus, 1767): restos de conchas y piedras, ic Hiatella rugosa (Linnaeus, 1767): restos de conchas, piedras y detrito coralígeno, ib Panopea norvegica (Spengler, 1793) Familia PHOLADIDAE Barnea candida (Linnaeus, 1758): arena con bloques de fango, i Pholas dactylus Linnaeus, 1758: arena y fango, ¡ Familia TEREDINIDAE *Bankia carinata (Gray, 1827): madera *lyrodus pedicellatus (Quatrefages, 1849): madera Nototeredo norvegica (Spengler, 1792): madera Familia XYLOPHAGIDAE Xylophaga dorsalis (Turton, 1819): madera y detrito coralígeno Familia THRACIDAE Thracia convexa (Wood, 1815): fango, c Thracia corbuloides Deshayes, 1830: fango, c Thracia papyracea (Poli, 1795): arena, i Thracia pubescens (Pulteney, 1799): fango, c Familia PANDORIDAE Pandora inaequivalvis (Linnaeus, 1758): arena, ¡ Pandora pinna (Montagu, 1803): fango y detrito coralígeno, b Familia POROMYIDAE Poromya granulata (Nyst y Westendorp, 1839): fango y detrito coralígeno, b Familia CUSPIDARIIDAE * Cardiomya striolata (Locard, 1898): fango y detrito coralígeno, b * Cuspidaria abbreviata (Forbes, 1843): b Cuspidaria cuspidata (Olivi, 1792): b Cuspidaria rostrata (Spengler, 1793): b Clase SCAPHOPODA Familia DENTALIIDAE Dentalium agile Sars, 1872 Dentalium inaequicostatum Dautzenberg, 1891: fango y detrito coralígeno, cb Dentalium panormum Chenu, 1842: b Dentalium vulgare da Costa, 1778: arena y detrito rocoso, ¡ Fustiaria rubescens (Deshayes, 1825): ¡ Familia SIPHONODENTALIDAE Pulsellum lofotense (Sars, 1865): fango y detrito coralígeno, b Codulus jeffreysi (Monterosato, 1875): b Entalina teftragona (Brocchi, 1814): b 60 Fig. 109 ++ Fig. 110 +++ Fig. 111 +++ Fig. 108 +++ GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figura 2. Concha de Rissoa gemmula (Sitges, 1,7 mm). Figure 2. Shell of Rissoa gemmula (Sitges, 1.7 mm). DISCUSIÓN COMENTARIOS SOBRE ALGUNOS TÁ- XONES: De la mayoría de especies citadas para el Garraf existen fotografías y des- cripciones actualizadas en la literatura, aunque algunas un poco dispersas. En este apartado nos hemos limitado a comentar algunos de los taxones que nos han parecido de mayor interés, ya sea por su rareza, por su importancia pesquera- comercial en el Garraf, por la escasa documentación bibliográfica existente, o bien por su importancia biológica en la zona. De muchas de estas especies repor- tamos fotografías, la mayoría al M.E.B., de la concha o de la protoconcha, según convenga para su identificación. Clase GASTROPODA Familia LEPETIDAE lothia fulva (O. E. Muller, 1776) Esta especie atlántica ha sido citada para el Mediterráneo por TAvIANI (1974), más concretamente para el Adriático, en fondos de fango entre 180 y 320 m de profundidad, aunque como comenta el propio autor, seguramente se trataba de una concha semifósil del Wuúrmiense. CECALUPO Y GIUustTI (1986) citan otro ejemplar en buenas condiciones de la Isla de Capraia, entre 400 y 440 m de profundidad. Nuestro único ejemplar también parece ser un fósil Wúrmiense, puesto que se trata de una concha mal conservada, procedente del detrito de “El Parrusset” entre 250 y 350 m de pro- fundidad. 61 Uber Familia LEPETELLIDAE Lepetella cfr. espinosae Dantart y Luque, 1994 No se han encontrado ejemplares vivos de esta especie, y la diagnosis sólo es posible estudiando la morfolo- gía del animal. De todas formas las otras dos especies con las que podría confundirse, Lepetella sierrai Dantart y Luque, 1994 y L. barrajoni Dantart y Luque, 1994, no se han encontrado en el Mediterráneo (ver DANTART Y LUQUE, 1994). Familia ADDISONIDAE Addisoniía excentrica Tiberi, 1857 (Fig. 3) MCLEAN (1985) señala que la princi- pal diferencia entre A. paradoxa Dall, 1882 del Atlántico occidental, y A. excen- trica (Tiberi, 1857) es el tamaño del adulto, dando 20, 3 mm de talla máxima para la primera y 10, 5 mm para la segu- nada. DANTART Y LUQUE (1994), tras una detallada discusión, consideran a A. paradoxa sinónimo posterior de A. excen- trica, y reportan ejemplares de esta última de hasta 12 mm. Nosotros hemos encontrado un ejemplar de A. excentrica de 20 mm, por lo que ratificamos esta sinonimia. Familia FISSURELLIDAE Emarginula pustula Thiele in Kuester, 1913 (Fig. 4) Esta especie ha sido considerada por PIANI (1984) como un “endemismo del archipiélago Toscano y de la costa Sarda oriental”, pero el hallazgo de un ejemplar en el detrito de “El Parrusset”, amplía su distribución al Mediterráneo occidental, hecho éste que era de espe- rar. Familia SCISSURELLIDAE Anatoma aspera (Philippi, 1844) Esta especie ha sido considerada si- nónina de A. crispata Fleming, 1828, o como una subespecie de ésta (SCHIRO, 1986), pero presenta una espira más alta, y parece ser que A. crispata no vive al sur de Escocia (Gofas, com. pers.), y ade- más ambas especies presentan diferen- cias en la rádula (DANTART, com. pers.). Familia RISSOIDAE Alvania cimicoides (Forbes, 1844) y Alvania testae (Aradas y Maggiore, 1843) (ies 790) Los miles de ejemplares de A. testae hallados en los contenidos estomacales de Astropecten irregularis, contrastan con los tan sólo cuatro de A. címicoides (hallados 62 todos ellos en la misma estrella), a pesar de que en sedimentos como el de “El Pa- rrusset” A. cimicoides es aproximadamente tres veces más frecuente que A. testae. GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figura 3. Addisonia excentrica (El Parrusset), 20 mm. Figura 4. Emarginula pustula (El Parrusset), 1,22 mm. Figura 5. Protoconcha de Danilia otaviana (El Parrusset). Figura 6. Obtusella macilenta (Vilanova). Figura 7. Alvania testae (Vilanova), 2,16 mm. Figura 8. Alvania subsoluta (isla de Capraia, Italia), 2,1 mm. Figure 3. Addisonia excentrica (El Parrusset), 20 mm. Figure 4. Emarginula pustula (El Parrusset), 1.22 mm. Figure 5. Protoconch ofDanilia otaviana (El Parrusset). Figure 6. Obtusella macilenta. (Vilanova). Fi- gure 7. Alvania testae (Vilanova), 2.16 mm. Figura 8. Alvania subsoluta (Capraia Island, Italy), 2.1 mm. 63 Iberus, 15 (1), 1997 Alvania zylensis Gotas y Warén, 1982 (Figs. 13, 14) Esta especie fue descrita por GOFAS Y WARÉN (1982) para las costas atlánticas de Marruecos. AARTSEN, MENKHORST Y GITTENBERGER (1984) la citan en la Bahía de Algeciras, y posteriormente, BOGI, COPPINI Y MARGELLI (1989) la mencionan por primera vez para el Mediterráneo, en el Tirreno. En el detrito de “El Parrus- set” hemos encontrado algunas conchas que asociamos a esta especie, concreta- mente a la forma de profundidad des- crita por BOGI ET AL. (1989), que presenta una teleoconcha con una escultura débil. El diámetro máximo de la protoconcha es de 530 mm. Aportamos, además, la fo- tografía de un ejemplar de la Isla de Al- borán (Fig. 14), que aunque presenta una teleoconcha idéntica a la de los ejempla- res de Vallcarca, tiene una protoconcha más pequeña, de 450 mm de diámetro. Obtusella macilenta (Monterosato, 1880) (Fig. 6) Es una especie abundante en todos los fondos fangosos del Garraf, mientras que sólo hemos hallado unos pocos ejempla- res de O. intersecta (Wood, 1857). Sinembar- go, lo normal en fondos similares de otras regiones es que la proporción sea inversa. Rissoa gemmula Fischer in de Folin, 1871 (Figs. 2, 20, 21) Se han encontrado dos conchas en el litoral de Sitges, más una en Es Caló (For- mentera, Islas Baleares). Aunque no se ha estudiado el material tipo, los tres ejem- plares se corresponden con la descripción aparecida en FOLIN (1871), que reprodu- cimos a continuación, y con la figura representada en NORDSIECK (1972): “... Long 1*/3 millim. Coquille conique- allongée, blanche, subdiaphane, ornée de có- tes longitudinales obsoletes, a peine indi- quées, et de stries spirales, visibles a la partie inférieure des tours. Sept tours de spire ven- trus: les trois premiers translucides, brillants, globuleux, papilliformes; le quatrieme dilaté, proportionnellement trés large; les dernieurs peu dilatés; suture bordée, ornée en dessous d'une petite zone transverse, brune, inte- rrompue de blanc; dernier tour orné, a sa partie moyenne, d'une zonule de meme co- loration; overture petite, ovale; periostoma simple. Observation. - On ne pourrait rap- procher ce Rissoa que du R. dolium (Nyst), (Nassa Philippi); mais notre espece est plus élancée, plus petite, á cótes obsoletes, et sa coloration est spéciale, comme la présence de la zone suturale et de la zone médiane du dernier tour”. En cuanto a la protoconcha, es lisa de 2!/4 vueltas de espira, con un diáme- tro máximo de 390 mm. Hemos incluído en el trabajo un dibujo detallado de la concha (Fig. 2), aparte de las fotografías realizadas al M.E.B. (Figs. 20, 21). Además, hemos fotografiado las espe- cies de Rissoidae a nuestro juicio más cer- canas, Pusillina inconspicua (Alder, 1844) (Figs. 17-19) y P. philippi (Aradas y Mag- giore, 1844) (Figs. 15, 16), de las que se diferencia por la coloración y forma. Familia CALYPTRAEIDAE Crepidula fornicata (Linnaeus, 1758) Aunque se ha hallado una única concha, su presencia en la zona puede explicarse por la introducción artificial adherida, al casco de un barco. Ade- 64 más, en el puerto de Barcelona (a tan sólo 45 km de distancia) han aparecido numerosos ejemplares vivos de esta especie. GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figuras 9-14. Género Alvania, protoconchas. 9, 10: A. testae (Vilanova); 11, 12: A. subsoluta (isla de Capraia, Italia); 13: A. zylensis (El Parrusset); 14: A. zylensis (isla de Alborán). Escalas, 9, 11-14: 200 um; 10: 100 ym. Figures 9-14. Genus Alvania, protoconchs. 9, 10. A. testae (Vilanova); 11, 12: A. subsoluta (Capraia Island, Italy); 13: A. zylensis (El Parrusset); 14: A. zylensis (Alborán Island). Scale bars, 9, 11-14: 200 ym; 10: 100 um. 65 Iberus, 15 (1), 1997 Figuras 15, 16. Pusillina philippi (cala Montjoy, Roses, Girona). 15: ejemplar de 2,1 mm; 16: pro- toconcha. Figuras 17-19. Pusillina inconspicua (lossa de Mar, Girona). 17: ejemplar de 1,7 mm; 18, 19: protoconcha. Figuras 20, 21. Ríssoa gemmula (Sitges). 20: ejemplar de 1,7 mm; 21: protocon- cha. Escalas, 16, 19, 21: 100 um; 18: 200 pm. Figures 15, 16. Pusillina philippi (cala Montjoy, Roses, Girona). 15: shell of 2.1 mm; 16: protoconch. Figures 17-19. Pusillina inconspicua (Tossa de Mar, Girona). 17: shell of 1.7 mm; 18, 19: protoconch. Figures 20, 21. Rissoa gemmula (Sitges). 20: shell of 1.7 mm; 21: protoconch. Scale bars: 16, 19, 21: 100 ym, 18: 200 um. 66 GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figuras 22-28. Protoconchas de Cerithiopsis. 22: C. diadema (Isla de Alborán); 23: C. jeffreysi (bahía de Almería); 24: C. minima (Sitges); 25: C. nana (Sitges); 26: C. scalaris (bahía de Almería); 27: C. tiara (isla de Alborán); 28: C. tubercularis (La Herradura, Granada). Figura 29: C. nana (Sitges), concha de 1,8 mm. Figura 30: C. tubercularis (La Herradura, Granada), concha de 2,4 mm. Escalas, 22-28: 200 um; 29, 30: 1 mm. Figures 22-28. Cerithiopsis protoconchs. 22: C. diadema (Alborán Island); 23: C. jeftreysi (Almería bay); 24: C. minima (Sitges); 25: C. nana (Sitges); 26: C. scalaris (Almería bay); 27: C. tiara (Alborán Island); 28: C. tubercularis (La Herradura, Granada). Figure 29: C. nana (Sitges), shell of 1.8 mm. Figure 30: C. tubercularis (La Herradura, Granada), shell of 2.4 mm. Scale bars, 22-28: 200 um; 29, 30: 1 mm. 67 Iberus, 15 (1), 1997 Familia EULIMIDAE La familia Eulimidae es una de las más ricas en aguas profundas, quizás más que la familia Turridae (BOUCHET Y WARÉN, 1986). A diferencia de otras regiones, la zona de estudio es pobre en especies del género Vitreolina, que prin- cipalmente viven en aguas infralitorales. Los pocos ejemplares de este género encontrados proceden de aguas profun- das. Crinophthetros sp. (Figs. 37, 38) Se han encontrado tres ejemplares frescos del género Crinophtheiros en con- tenidos estomacales de Astropecten irre- gularis, a profundidades superiores a los 200 m. La especie C. comatulicola (Graf, 1875) es frecuente en fondos infralitorales, siempre asociada a Ante- don mediterranea (Lamarck) (TEMPLADO, com. pers.), pero a profundidades supe- riores a 200 m, el crinoideo presente es Leptometra phalangium (Muller), por lo que podría tratarse de otra especie dife- rente perteneciente al género Crinoph- theiros. Parvioris ibizenca (Nordsieck, 1968) La especie Parvioris ibizenca ha sido referida normalmente en la bibliografía como P. microstoma (Brusina, 1864), pero según Gofas (com. pers.) el nombre correcto sería el primero de éstos, porque P. microstoma ya está preocu- pado, con lo que se considera sinoni- mía. Vitreolina sp. La especie tipo del género Vitreolina es Eulima incurva Bucquoy, Dautzen- berg y Dollfus, 1883, especie poco clara (BOUCHET Y WARÉN, 1986). Además, hay una gran confusión con las especies adscritas al género Vitreolina (eulímidos de pequeño tamaño,con forma curva- da), por lo que hemos preferido men- cionar estos ejemplares como Vitreolina sp. Familia MURICIDAE Trophon echinatus (Kiener, 1840) y Trophon sp. (Figs. 41-44) T. echinatus presenta una considerable variación de formas de la teleoconcha, espe- cialmente con relación a la profundidad. Sin embargo, no se ha descrito variabili- dad en la protoconcha. En el detrito de “El Parrusset” hemos encontrado dos tipos de protoconcha que presentan tamaños muy diferentes; una con un diámetro máximo de unos 670 mm, que asignamos a T. echi- natus, y otra con un diámetro máximo regis- trado entre 770 y 830 mm, que denomi- namos provisionalmente Trophon sp. Trophon barvicensis (Johnston, 1825) (Fig. 45) Según BOUCHET Y WARÉN (1985), no han visto ningún especimen mediterrá- neo que pueda asignarse inequívoca- 68 mente a este taxón. También comentan que algunas citas de Trophonopsis richardi (Dautzenberg y Fischer, 1896) para el GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Eigura 31. Talassia daguenetí (El Parrusset), concha juvenil de 0,88 mm. Figura 32. Graphis albida (El Parrusset), 1,83 mm. Figuras 33, 34. Graphis albida (La Herradura, Granada). 33: concha de 2,4 mm, 34: protoconcha. Figuras 35, 36. Vitreolina perminima (El Parrusset). 35: concha de 2,2 mm; 36: protoconcha. Figuras 37, 38. Crinophtheiros sp. (Vilanova). 37: concha de 4,6 mm; 38: proto- concha. Figuras 39, 40. Protoconchas de Epitonium. 39: E. clathratulum (Mijas Costa, Málaga); 40: E. linctum (El Parrusset). Escalas, 34: 100 um; 36: 300 um; 38: 300 um; 39, 40: 200 pm. Figure 31. Talassia dagueneti (El Parrusset), juvenile shell 00.88 mm. Figure 32. Graphis albida (El Parrusset), 1.83 mm. Figures 33, 34. Graphis albida (La Herradura, Granada). 33: shell 0f2.4 mm; 34: protoconch. Figures 35, 36. Vitreolina perminima (El Parrusset). 35: shell of 2.2 mm; 36: protoconch. Figures 37, 38. Crinophtheiros sp. (Vilanova). 37: shell of 4.6 mm; 38: protoconch. Figures 39, 40. Pro- toconchs ofEpitonium. 39: E. clathratulum (Mijas Costa, Málaga); 40: E. linctum (El Parrusset). Scale bars, 34: 100 um; 36: 300 ym; 38: 300 um; 39, 40: 200 um. 69 Iberus, 15 (1), 1997 Figuras 41, 42. Trophon echinatus (El Parrusset). 41: concha juvenil de 4,75 mm; 42: protoconcha. Figuras 43, 44. Trophon sp. (El Parrusset). 43: concha juvenil de 2,7 mm; 44: protoconcha. Figura 45. Trophon barviciensis (El Parrusset), 19 mm. Figuras 46, 47. Trophon muricatus. 46: concha juve- nil de 5,9 mm; 47: protoconcha. Escalas 500 um. Figures 41, 42. Trophon echinatus (El Parrusset). 41: juvenile shell of 4.75 mm; 42: protoconch. Figures 43, 44. Trophon sp. (El Parrusset). 43: juvenil shell of 2.7 mm; 44: protoconch. Figure 45. Trophon barviciensis (El Parrusset), 19 mm. Figures 46, 47. Trophon muricatus. 46: juvenil shell of 5.9 mm; 47: protoconch. Scale bars 500 um. 70 GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figuras 48, 49. Bela brachystoma (Vilanova). 48: ejemplar de 3,6 mm; 49: protoconcha. Figuras 50, 51. Mangelia attenuata (Vilanova). 50: ejemplar de 2,5 mm; 51: protoconcha. Figuras 52, 53. Mangelia nuperrima (El Parrusset). 52: ejemplar de 7 mm; 53: protoconcha. Figuras 54, 55. Mangelia serga (El Parrusset). 54: ejemplar de 2,75 mm; 55: protoconcha. Escalas 500 um. Figures 48, 49. Bela brachystoma (Vilanova). 48: shell of 3.6 mm; 49: protoconch. Figures 50, 51. Mangelia attenuata (Vilanova). 50: shell 0f2.5 mm; 51: protoconch. Figures 52, 53. Mangelia nupe- rrima (El Parrusset). 52: shell of 7 mm; 53: protoconch. Figures 54, 55. Mangelia serga (El Parrusset). 54: shell 0f 2.75 mm; 55: protoconch. Scale bars 500 yum. 71 Iberus, 15 (1), 1997 Mediterráneo (ver DI GERONIMO Y PaA- NETTA, 1973 y FRANCHINI Y FRILLI, 1970), podrían corresponder a T. barvicensis. Nuestro único ejemplar proveniente de “El Parrusset” parece un ejemplar nor- mal de T. barvicensis (Warén, com. pers.). Familia TURRIDAE Mangelia costata (Donovan, 1804) AARTSEN ET AL. (1984) diferencian M. coarctata (Forbes, 1840) de M. costata, siendo la primera más grande y alar- gada, con una o dos costillas más y pre- sentando una coloración uniforme. De todas maneras, dichos autores no des- cartan la posibilidad de que M. costata sea la forma litoral y M. coarctata la de aguas profundas de una misma especie. El análisis de más de 300 ejemplares del Mediterráneo español de la colección de uno de los autores, desde aguas someras hasta profundidades de 350 m, y cu- briendo una área geográfica desde el Mar de Alborán hasta Cataluña, nos sugiere que, efectivamente, ambas son formas de una misma especie, con una ligera tendencia a aumentar el tamaño y a atenuar la coloración a medida que aumenta la profundidad. Además, los escasos ejemplares de coloración uni- forme encontrados en aguas profundas son ejemplares subfósiles. Por lo tanto, y atendiendo sólo a las características de la concha, consideramos a M. coarctata como sinónimo de M. costata. Mangelia attenuata (Montagu, 1803) (Figs. 50, 51) Del mismo modo que en el caso anterior, creemos que M. tenuicostata (Brugnone, 1868) es sinónimo de M. atte- nuata. Esta especie, que se encuentra desde el litoral hasta los 250 m, dismi- nuye de tamaño al aumentar la profun- didad, y atenúa la coloración. Además, las vueltas de espira se van haciendo más escalonadas y las costillas más mar- cadas, pero la protoconcha no sufre variación alguna. Las Figuras 50 y 51 ilustran la forma de aguas profundas. Mangelia nuperrima (Tiberi, 1855) (Figs. 52, 53) Esta especie se diferencia de Man- gelia serga (Dall, 1881) por poseer vueltas de espira más redondeadas y una boca más ancha. Se ha encontrado un ejemplar fresco en el contenido es- tomacal de Astropecten irregularis, y cuatro conchas en el detrito de “El Pa- rrusset”. Mangelia serga (Dall, 1881) (Figs. 54, 55) Esta especie se ha citado pocas veces en el Mediterráneo; una vez para Cerdeña (CECALUPO, 1984) y otra vez para el Tirreno Central (SMRIGLIO, MARIOTTINI Y GRAVINA, 1987b). Se ha encontrado un ejemplar con restos de partes blandas en contenidos estomacales de Astropecten irregularis y tres conchas en el detrito de “El Parrusset”. Taranis moerchi (Malm, 1861) (Figs. 57, 58, 59) Esta especie, de concha extremada- mente variable, no es rara en el Medite- 72 rráneo. Las medidas de la protoconcha ilustrada son las siguientes: 500 mm de GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figura 56. Microdrillia loprestiana (EL Parrusset), 3,1 mm. Figuras 57-59 Taranis moerchi (El Parrusset). 57: concha de 4,4 mm, 58, 59: protoconcha. Figuras 60-62. Taranis sp. (El Parrusset). 60: concha de 2,8 mm; 61, 62: protoconcha. Escalas, 58, 61, 62: 500 um; 59: 200 um. Figure 56. Microdrillia loprestiana (EL Parrusset), 3.1 mm. Figures 57-59. “Taranis moerchi (El Parrusset). 57: shell of 4.4 mm; 58, 59: protoconch. Figures 60-62. Taranis sp. (El Parrusset). 60: shell of 2.8 mm; 61, 62: protoconch. Scale bars, 58, 61, 62: 500 ym; 59: 200 ym. Iberus, 15 (1), 1997 altura y 550 mm de diámetro máximo. Se han encontrado 18 ejemplares (inclu- yendo juveniles) en el detrito de “El Parrusset”. Taranis sp. (Figs. 60, 61, 62) Junto a los ejemplares de T. moerchi, hemos encontrado una concha de aspecto similar a ésta, pero que difiere en la protoconcha, que es mucho más grande (650 mm de altura y 730 mm de diámetro máximo), y además presenta una escultura puntiforme alineada en cordones desde el mismo ápice de la protoconcha, mientras que en T. moerchi los cordones de puntos se desordenan en el ápice. Este ejemplar mide 2, 8 mm de longitud. Microdrillia loprestiana (Calcara, 1841) (Fig. 56) Esta especie es común en contenidos estomacales de estrellas a partir de los 60-80 metros, y en el detrito de “El Parrusset”. Pleurotomella demosia (Dautzenber y Fischer, 1896) (Figs. 66, 67, 68) Nuestros ejemplares se ajustan a la descripción y figuras aportadas por BOu- CHET Y WARÉN (1980), aunque el diá- metro de la protoconcha es menor. Las medidas que presenta la protoconcha fo- tografiada (Figs. 67, 68) son las siguien- tes: 325 mm de diámetro de la P1 y 580 mm de diámetro de la P2. Esta especie fue citada por primera vez para el Mediterráneo por BoGI (1985), y posteriormente por CECALUPO (1988) para Cerdeña y por BOGI ET AL. (1989) para el Tirreno. Hemos encontrado dos ejemplares frescos en contenidos estoma- cales de Astropecten irregularis y dos conchas en el detrito de “El Parrusset”. Pleurotomella coeloraphe (Dautzenberg y Fischer, 1896) (Figs. 63, 64, 65) De esta especie, de aspecto más glo- boso que P. eurybrocha y que P. demosia, sólamente se conoce el material de la zona batial de Azores, recolectado en va- rias estaciones de la expedición MO- NACO y en una estación de la expedi- ción PORCUPINE (BOUCHET Y WARÉN, 1980). Presenta una protoconcha similar a la de P. eurybrocha en cuanto a forma, pero se diferencia en que la protoconcha embrionaria es reticulada (como en P. de- mosia) y no granulada como en P. eury- brocha. Las medidas que presenta la pro- toconcha fotografiada son las siguientes: 210 mm de diámetro de la P1 y 460 mm de diámetro de la P2. El diámetro de la protoconcha a 100 mm del ápice es de 200 mn, al igual que en P. eurybrocha. Hemos encontrado 4 ejemplares juveniles en el detrito de “El Parrusset”. Se trata por tanto de la primera cita de esta especie para el Mediterráneo. Familia TJAERNOEIDAE Familia monogenérica de Hetero- branchia descrita por WARÉN (1991) en base a la anatomía externa del animal. Previamente, el género Tjaernoeia, des- 74 crito por WARÉN Y BOUCHET (1988), ha- bía sido situado provisionalmente en la familia Pyramidellidae por los mismos autores. GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figuras 63-65. Pleurotomella coeloraphe (El Parrusset). 63: juvenil de 1,6 mm; 64, 65: protoconcha. Figuras 66-68. Pleurotomella demosía (El Parrusset). 66: juvenil de 2,1 mm; 67, 68: protoconcha. Figura 69. Teretía teres (El Parrusset), 4,4 mm. Escalas, 64, 67: 500 pm; 65, 68: 200 ym. Figures 63-65. Pleurotomella coeloraphe (El Parrusset). 63: juvenile shell of 1.6 mm; 64, 65: proto- conch. Figures 66-68. Pleurotomella demosia (El Parrusset). 66: juvenile shell of 2.1 mm; 67, 68: pro- toconch. Figure 69. Teretia teres (El Parrusset), 4.4 mm. Scale bars, 64, 67: 500 pm; 65, 68: 200 ym. 73 Iberus, 15 (1), 1997 Figuras 70-77. Familia Pyramidellidae. 70: Chrysallida brattstroemi (El Parrusset), 1,1 mm. 71: Chry- sallida dollfusi (LEscala, Girona), 2,8 mm. 72: Eulimella ataktos (Vilanova), 2,7 mm. 73: Eulimella ventricosa (Isla de Alborán), 2,36 mm. 74: Enlimella unifasciata (Blanes, Girona), 5,5 mm. 75: Odos- tomia afzelii (Vilanova), 1,5 mm. 76: Odostomia hansgei (Vilanova), 1,6 mm. 77: Turbonilla acutis- sima (Mijas, Málaga), 4,4 mm. Escalas, 70: 200 pm; 71, 72, 73, 75, 76: 500 um; 74, 77: 1 mm. Figures 70-77. Family Pyramidellidae. 70: Chrysallida brattstroemi (El Parrusset), 1.1 mm. 71: Chry- sallida dollfusi (L'Escala, Girona), 2.8 mm. 72: Eulimella ataktos (Vilanova), 2.7 mm. 73: Eulimella ventricosa (Alborán Island), 2.36 mm. 74: Eulimella unifasciata (Blanes, Girona), 5.5 mm. 75: Odos- tomia afzelii (Vilanova), 1.5 mm. 76: Odostomia hansgei (Vilanova), 1.6 mm. 77: Turbonilla acutis- sima (Mijas, Málaga), 4.4 mm. Scale bars, 70: 200 um; 71, 72, 73, 75, 76: 500 um; 74, 77: 1 mm. 76 GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figuras 78, 79. Ringicula cfr. leptocheila (El Parrusset). 78: concha de 2,37 mm; 79: protoconcha. Figuras 80, 81. Ringicula auriculata (Vilanova). 80: concha de 3,75 mm; 81: protoconcha. Figura 82. Cylichnina umbilicata (Vilanova), 1,3 mm. Figura 83: Limacina retroversa (El Parrusset), 2,7 mm. Figuras 84, 85. Peracle reticulata (El Parrusset). 84: concha de 2 mm; 85: protoconcha. Escalas, 79, 85: 200 um; 81: 500 um. Figures 78, 79. Ringicula cfr. leptocheila (El Parrusset). 78: shell 02.37 mm; 79: protoconch. Figures 80, 81. Ringicula auriculata (Vilanova). 80: shell of 3.75 mm, 81: protoconch. Figure 82. Cylichnina umbili- cata (Vilanova), 1.3 mm. Figure 83: Limacina retroversa (El Parrusset), 2.7 mm. Figures 84, 85. Peracle reticulata (El Parrusset). 84: shell of 2 mm; 85: protoconch. Scale bars, 79, 85: 200 ym; 81: 500 ym. 7 Iberus, 15 (1), 1997 Tjaernoeia exquisita (Jeffreys, 1883) Especie poco frecuente en el Medite- rráneo, en donde WARÉN (1991) la señala entre 25 y 200 m. Nosotros hemos encontrado ejemplares a más profundidad en el detrito de “El Parrus- set”. Familia ARCHITECTONICIDAE Basisulcata lepida (Bayer, 1942) Sólo se han encontrado dos ejempla- res vivos en “La Mar de Nit”, a unos 40 m de profundidad, en una zona detrítica con grandes colonias de Bolinus brandaris (L., 1758), cuyas conchas estaban recubiertas por la anémona Calliactis parasitica (Couch). Familia PYRAMIDELLIDAE (Figs. 70-77) En el trabajo sobre Pyramidellidae del Mediterráneo español de PEÑAS, TEMPLADO Y MARTÍNEZ (1996) se aporta una gran cantidad de datos sobre la pre- sencia de esta familia en fondos del Garraf. Asimismo, se citaba por primera vez para el Mediterráneo las siguientes especies: Chrysallida brattstroemi Warén, 1991, Eulimella ataktos Warén, 1991, Odostomia afzelii (Warén, 1991), y Odosto- mia hansgei (Warén, 1991), las cuatro comunes en esta zona, y ninguna hallada hasta ahora en ninguna otra localidad del Mediterráneo español. Además, en el mismo trabajo, numero- sas especies son citadas por primera vez para el Mediterráneo español, también procedentes del Garraf. Las especies que no habían sido cita- das para el Garraf en el trabajo anterior de PEÑAS ET AL. (1996), y que hemos hallado posteriormente, son: Chrysallida dollfusi (Kobelt, 1903), Eulimella unifasciata (Forbes, 1844), Eulimella ventricosa (Forbes, 1844), y Turbonilla acutissima Monterosato, 1884. El hallazgo en “El Parrusset” de una concha subfósil de Chrysallida pellucida (Dillwyn, 1817), no contradice la hipóte- sis de PEÑAS ET AL., (1996), que comen- tan que, en la actualidad, esta especie atlántica no penetra en el Mediterráneo más allá del mar de Alborán. Familia SCAPHANDRIDAE Scaphander punctostriatus (Mighels y Adams, 1841) Esta especie ha sido citada raras veces en el Mediterráneo (MONTERO- SATO, 1880; Ros, 1975; BOUCHET Y TAvIANI, 1989), y ha sido tradicional- mente considerada una especie de aguas profundas del Atlántico. Se han encon- trado 6 conchas en el detrito de “El Parrusset”. Familia DISCODORIDIDAE Taringa faba (Ballesteros, Llera y Ortea, 1984) Esta especie se encuentra en la playa del búnker de Cubelles, bajo piedras a menos de 50 centímetros de profundidad. Ésta es la localidad tipo 78 de la especie, descrita por BALLESTE- ROS, LLERA Y ORTEA (1984), donde se puede encontrar con relativa frecuen- cia. GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figura 86. Yoldiella lucida, 4,25 mm. Figura 87. Yoldiella nana, 2,9 mm. Figuras 88-93. Yoldiella philippiana. 88, 89: juveniles de 2,9 y 2,3 mm; 90: adulto de 4,3 mm; 91: juvenil, detalle de los dientes de la charnela; 92: charnela; 93: protoconcha. Todos las especies de El Parrusset, contenido estomacal de Astropeceten, 250-350 m. Escalas; 92: 200 pm; 93: 100 pm. Figure 86. Yoldiella lucida, 4.25 mm. Figure 87. Yoldiella nana, 2.9 mm. Figures 88-93. Yoldiella phi- lippiana. 88, 89: juvenile shells of 2.9 and 2.3 mm; 90: adult shell of 4. 3 mm; 91: juvenil shell, hinge lateral view; 92: hinge; 93: protoconch. All the especies from El Parrusset, gut content of Astropecten, 250-350 m. Scale bars, 92: 200 ym; 93: 100 ym. Y Iberus, 15 (1), 1997 Familia FAVORINIDAE Favorinus vitreus (Ortea, 1982) Esta especie fue descrita por ORTEA (1982) en Tenerife (Islas Canarias), a par- tir de dos ejemplares, y únicamente se ha citado 1 ejemplar para el Mediterrá- neo, en al Cabo de Palos (Murcia) en ri- zomas de Posidonia a 5 m de profundi- dad (TEMPLADO, 1982). Se ha recolec- tado un ejemplar de esta especie en Ju- nio de 1993 en Cubelles, debajo de una piedra a unos 20-30 cm de profundidad. Clase BIVALVIA Familia NUCULIDAE Nucula cfr. nucleus (Linnaeus, 1758) Gofas (com. pers.) opina que su pre- sencia en el Mediterráneo es dudosa, siendo N. hanleyi Winckworth, 1930, la especie más normal, que presenta un periostraco más brillante, con estrías radiales oscuras, y más alargada ante- riormente. SALAS (1996) ilustra un ejem- plar de N. nucleus del Mediterráneo. Hemos encontrado ejemplares juveniles con una protoconcha que coincide con la descripción de GOFAS Y SALAS (1996), y ejemplares adultos que, aunque han perdido parte del periostraco, asigna- mos a N. nucleus. Familia YOLDIIDAE La familia Yoldiidae es la principal familia de Nuculanoidea en el Medite- rráneo, y en el Garraf sólo hemos ha- llado el género Yoldiella. Las otras fami- lias de Nuculanoidea no han sido reco- lectadas en este estudio por presentar una distribución batimétrica de mayor profundidad. Aunque tenemos constan- cia de la existencia de especies de otras familias en zonas más profundas frente a la costa del Garraf (Dantart, com. pers.). Hemos tratado de recopilar aquí las citas modernas de las especies de la sub- familia Yoldiellinae en el Mediterráneo debido a que la informacion sobre el género es muy dispersa. Sin embargo, no se han tenido en cuenta descripcio- nes originales, citas antiguas ni sinoni- mias, que se pueden encontrar en los artículos mencionados en esta sección. De todas las citas recopiladas para el Mediterráneo, reconocemos sólo 5 espe- cies. Se han dado entre corchetes los nombres utilizados incorrectamente, o los que han sido considerados como sinonimias. Yoldiella lucida (Lovén, 1846) (Fig. 86) Di Geronimo y Panetta (1973): Golfo de Taranto, 940-1000 m Warén (1989): distribucion Mediterránea entre 100 y 1000 m de profundidad Es la especie de Yoldiella menos fre- cuente de las halladas en Garraf, habién- dose encontrado sólo tres conchas, dos en contenidos estomacales de estrellas y 80 una en el sedimento, todas del detrito de “El Parrusset”, entre 250 y 350 m de profundidad, por lo que se deduce que vive en la zona. GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figura 94. Bathyarca pectunculoides (El Parrusset), 1,3 mm. Figura 95. Bathyarca philippiana (El Parrusset), 1,3 y 1,6 mm. Figura 96. Crenella pellucida (El Parrusset), 710 um. Figura 97. Modiolula phaseolina (El Parrusset), 1,96 mm. Figure 94. Bathyarca pectunculoides (El Parrusset), 1.3 mm. Figure 95. Bathyarca philippiana (El Parrusset), 1.3 and 1.6 mm. Figure 96. Crenella pellucida (El Parrusset), 710 pm. Figure 97. Modiolula phaseolina (El Parrusset), 1.96 mm. 81 Iberus, 15 (1), 1997 Yoldiella nana (M. Sars, 1865) (Fig. 87) Cecalupo y Giusti (1986): Isla de Capraia, 400-440 m [Portlandia frigida (Torell, 1859)] Bogi et al. (1989): Capo Corso, 150 m [Portlandia frigida (Torell, 1859)] Warén (1989): Banyuls, 650-770 m Se han encontrado varias conchas en contenidos estomacales de estrellas entre 100 y 350 m de profundidad, y otras en el detrito de “El Parrusset”, por lo que se deduce que también vive en la zona. Yoldiella philippiana (Nyst, 1845) (Figs. 88-93) Di Geronimo y Panetta (1973): Golfo de Taranto, 350-1000 m [Yoldiella tenuis (Philippi)] Di Geronimo (1974): Jónico [Yoldiella tenuis (Philippi)] Cecalupo y Giusti (1986): varios ejemplares vivos, Isla de Capraia, 400-440 m Warén (1989): Mediterráneo, 100-300 m Bonfitto y Sabelli (1995): Cerdeña, 245-1707 m Es la especie más abundante, apare- ciendo tanto en sedimentos como en contenidos intestinales de Astropecten, desde los 60 hasta los 350 m de profun- didad. Se han encontrado numerosos juveniles vivos. Las otras dos especies mediterráneas de este género, Yoldiella messanensis (Seguenza, MS, Jeffreys, 1870) y Yoldiella seguenzae Bonfitto y Sabelli, 1995, no se han encontrado en la zona de estudio. La primera ha sido citada en el Medite- rráneo en general (150-3000 m) y en el mar de Alborán (60-1235 m) por TERRENI (1980) CECALUPO Y GIUST1 (1986), WARÉN (1978, 1989), ALLEN Y HANNAH (1989) y SALas (1996). La se- gunda sólo se conoce de Cerdeña -(245- 1707 m) y mar de Alborán, entre 480 y 1005 m (BONFITTO Y SABELLL 1995; SALAS 1996). Familia MYTILIDAE El género Idas en el Mediterráneo está representado por tres especies (Wa- RÉN, 1991), asociadas siempre a restos de esqueletos de cetáceos. Además de este hábitat, 1. argenteus Jeffreys, 1876 e 1. ghisotti Warén y Carozza, 1990 también pueden encontrarse en restos de troncos (WARÉN, 1991, 1993). La utilización del género Idas Jeffreys, 1878, en vez de Ida- sola Iredale, 1915, contrariamente a lo propuesto por DELL (1987), se basa en lo expuesto por WARÉN (1991). Idas cfr ghisottii Warén y Carrozza, 1990 En un tronco de madera arrojado a la playa tras una tormenta, se ha encontrado un ejemplar que asignamos a esta especie, junto con varios ejem- plares de Teredinidae. Esta especie sólo se ha encontrado asociada a restos de madera en el Mediterráneo por CARROZZA (1984), que la identificó erróneamente como Myrina modiolaefor- 82 mis Sturany, 1896. Fue corregido poste- riormente por WARÉN Y CARROZZA (1990), que la describieron como nueva especie. De todas formas, y debido a la pérdida del único ejemplar hallado, la asignación de este especimen podría ser errónea, aunque estamos seguros de que no se trata de /. simpsoni (Mars- hall, 1900). GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figura 98. Thyasira (1) biplicata (El Parrusset), 4,9 mm. Figura 99. Thyasira (1) obsoleta (El Parrusset), 1,46 y 1,87 mm. Figura 100. Thyasira (P) subovata (El Parrusset), 1 y 1,16 mm. Figuras 101, 102. Thyasira (Parathyasira) granulosa. 101: ejemplar de 5.4 mm. 102: detalle de la granulación de la concha. Escala 400 um. Figure 98. Thyasira (T.) biplicata (El Parrusset), 4.9 mm. Figure 99. Thyasira (T.) obsoleta (El Parrusset), 1.46 and 1.87 mm. Figure 100. Thyasira (P.) subovata (El Parrusset), 1 and 1.16 mm. Figures 101, 102. Thyasira (Parathyasira) granulosa. 101: shell of5.4 mm. 102: shell granulation detail. Scale bar 400 ym. 83 Iberus, 15 (1), 1997 Idas simpsoni (Marshall, 1900) Se trata de otro Mytilidae interesante, que vive en restos orgánicos de esquele- tos de cetáceos o peces, aunque mucho más común que el anterior y de distribu- ción más amplia. Se han encontrado va- rios ejemplares asociados a esqueletos. Modiolula phaseolina (Philippi, 1844) (Fig. 97) Se han encontrado unos pocos ejem- plares vivos en el detrito de “El Parrus- set”, aunque esta especie destaca por la gran cantidad de conchas Wúrmienses de gran tamano (más de 20 mm) encon- tradas en esta localidad. Familia PECTINIDAE Hemos incluido en esta familia las especies clasificadas por algunos auto- res como Propeamussiidae (o Amussii- dae). De todas formas, esta clasificación es provisional hasta que se realice un es- tudio filogenético que resuelva la cues- tión. Hemos seguido la sistemática utili- zada por WAGNER (1991), excepto en la separación de Pectinidae y Amussiidae. Para algunas especies de “Amussiidae”, hemos seguido a SMRIGLIO Y MARIOT- TINI (1990). Pseudamussium septemradiatum (O. E. Muller, 1776) Se han encontrado ejemplares vivos de esta especie en fondos de fango entre 150 y 200 m de profundidad, mientras que VinYas (1981) la consideraba como una especie de aguas frías, que supues- tamente se había extinguido del Medite- rráneo, aunque abundante en los sedi- mentos Wúrmienses (MArs, 1958; MAR- TINELL Y JULIA-BRUGUES, 1973; VINYAS, 1981; DOMENECH Y MARTINELL, 1982). SALAS (1996) la cita viva para el mar de Alborán. Flexopecten glaber (Linnaeus, 1758) Gofas (com. pers.) ha comentado que se trata de una especie más bien lagunar, rara en el Mediterráneo occidental, y que algunos juveniles grandes de E. flexuosus, que no presentan aún la flexuosidad típica de la especie, pueden ser confundidos con F. glaber. Hemos encontrado una única valva de esta especie, de 33 mm de altura con 10 costillas principales, claramente diferente de FE. flexuosus, cuyos mayores ejemplares encontrados en la zona miden 28 mm y presenta 5 costillas principales. Familia LIMIDAE Limatula cfr. gwyni (Sykes, 1903) Se ha encontrado una única valva fresca procedente del detrito de “El Parrusset”. El ejemplar se destruyó durante el proceso de montaje para el 84 M.E.B. El color era blanco y presentaba una forma y tamaño similar a L. subovata, aunque totalmente lisa y sin restos de costillas radiales. GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figura 103. Thyasira (Leptaxinus) incrassata (El Parrusset), 1,66 mm. Figura 104. Thyasira (Axinulus) croulinensis (El Parrusset), 1,26 mm. Figura 105. Thyasira (A.) eumyaria (El Parrusset), 2,9 mm. Figura 106. 7hyasira (Mendicula) ferruginea (El Parrusset), 1,46 mm. Figura 107. Arculus sp. (El Parrusset), 1,02 mm y 1,16 mm. Figure 103. Thyasira (Leptaxinus) incrassata (El Parrusset) 1.66 mm. Figure 104. Thyasira (Axinulus) croulinensis (El Parrusset), 1.26 mm. Figure 105. Thyasira (A.) eumyaria (El Parrusset), 2.9 mm. Figure 106. Thyasira (Mendicula) ferruginea (El Parrusset), 1.46 mm. Figure 107. Arculus sp. (El Parrusset), 1.02 mm and 1.16 mm. 95 Iberus, 15 (1), 1997 Familia LUCINIDAE Lucinoma borealis (Linnaeus, 1767) Se han encontrado ejemplares vivos de gran tamaño (hasta 47,9 mm, un ejemplar de la coleccion de J. L. Ferrer), en fondos de fango entre 60 y 80 m de profundidad. Familia THYASIRIDAE (Figs. 98-106) Los tiasíridos son la principal familia de bivalvos que habita en el talud. Muchas especies viven desde el circalito- ral hasta profundidades abisales. Las especies atlánticas han sido reciente- mente revisadas a nivel morfológico por PAYNE Y ALLEN (1991), pero son pocos los trabajos sobre este grupo en el Mediterrá- neo. La descripción de una nueva especie mediterránea por CARROZZA (1981), así como las redescripciones de algunas especies de Monterosato publicadas por GAGLINI (1991) son las obras modernas más importantes para este mar. Además, existen algunas citas dispersas (p. e. DI GERONIMO Y PANETTA, 1973; TERRENI, 1980; CARROZZA, 1984; CIANFANELLI Y TALENTI, 1987; CECALUPO Y GIusTI, 1989), pero no hay ningún trabajo taxonómico de revisión importante. Tampoco preten- demos aquí realizar una revisión taxonó- mica del grupo, pero hemos querido foto- grafiar las 8 especies aparecidas en el detrito de “El Parrusset” (Figs. 98-106), constituyendo 6 de ellas la primera cita para el Mediterráneo español. Thyasira biplicata (Philippi, 1836) (Fig. 98) Esta especie, fue descrita por PHI- LIPPI (1836) para zonas abisales del Mar Mediterráneo, pero ha sido referida en la literatura como T. flexuosa (Montagu, 1803), que es una especie litoral atlántica con la ondulación posterior más suave (Gofas, com. pers.), de presencia dudosa en el Mediterráneo. Familia MONTACUTIDAE Epilepton clarkiae (Clark, 1852) Todos los ejemplares se han encon- trado en una comunidad dominada por Turritella communis Risso, 1826, a unos 60 m de profundidad. Epilepton sp. Este pequeño Epilepton, aparecido en el detrito coralígeno de “El Parrusset” está en este momento en proceso de des- cripción. Familia NEOLEPTONIDAE Arculus sp. (Fig. 107) Hemos identificado provisionalmen- te estos ejemplares, hallados también en 86 “El Parrusset”, dentro del género Arcu- lus. GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Figura 108. Xylophaga dorsalis (Vilanova): exterior valva derecha, 3,4 mm; interior valva izquierda, 1,6 mm. Figura 109. paleta de Bankia carinata (Vilanova), 4,7 mm. Figura 110. Lyrodus pedicellatus (Vilanova): exterior valva izquierda, 1,9 mm; interior valva derecha, 2,16 mm; paleta de 2,5 mm. Fi- gura 111. Nototeredo norvegica (Vilanova): exterior valva izquierda, 1, 38 mm; interior valva derecha, 1,5 mm; paleta de 1,7 mm. Figures 108. Xylophaga dorsalis (Vilanova): outside of right valve, 3.4 mm; inside of lefi valve, 1.6 mm. Figure 109. pallet ofBankia carinata (Vilanova), 4.7 mm. Figures 110. Lyrodus pedicellatus (Vilanova): outside of lefi valve, 1.9 mm; inside of right valve, 2.16 mm; pallet of 2.5 mm. Figure 111. Nototeredo norvegica (Vilanova): outside of lefi valve, 1.38 mm; inside of right valve, 1.5 mm), pallet of 1.7 mm. 87 Mars SANS Familia ASTARTIDAE Goodallia sp. En el detrito de “El Parrusset” han aparecido ejemplares pertenecientes al género Goodallia, similares al taxon Los MOLUSCOS DE “EL PARRUSSET”: Como se comentaba anteriormente, “El Parrusset” es una biocenosis de coral blanco (PÉRES Y PICARD, 1964) con fan- go, cuyas especies predominantes son los madreporarios Dendrophyllia corni- gera (Lamarck) y Caryophyllia sp., aso- ciada a una tanatocenosis del Wur- miense, con la presencia de algunas es- pecies subfósiles que actualmente están extintas en el Mediterráneo. En esta lo- calidad se han obtenido numerosas es- pecies de moluscos de profundidad, siendo Trophon barvicensis y Pleurotomella coeloraphe la primera vez que se citan para el Mediterráneo, y otras la primera vez que se citan para el litoral Medite- rráneo español. Una comunidad muy similar a la de “El Parrusset” ha sido descrita por SMRI- GLIO, MARIOTTINI Y GRAVINA (1987a; 1987b) para el mar Tirreno Central, donde estudiaron una biocenosis de co- ral blanco con detrito fangoso situada entre 400 y 600 m de profundidad, en la que la especie de madreporario predo- minante es Dendrophyllia cornigera (La- marck). Aunque se trate de dos comuni- dades muy parecidas desde un punto de vista biológico y paleontológico, las es- pecies de moluscos que se encuentran en la localidad del Tirreno Central (Sm- RIGLIO ET AL., 1987a; 1987b, 1988a; 1988b; 1989; 1993) son características de aguas más profundas. Sin embargo, algunas de las especies son coincidentes en am- bas biocenosis, como por ejemplo Micro- drillia lopestriana (Calcara, 1841), Mange- lia serga (Dall, 1881) y Teretia teres (For- bes, 1844), entre los túrridos. En cuanto a los bivalvos actuales en- contrados en esta biocenosis, los grupos mayoritarios son Nuculidae, Yoldielli- dae, Bathyarca, Astarte, Veneridae, Thya- siridae, Pectinidae y Xylophaga, siendo 88. G. macandrewi Smith, 1881, que posi- blemente se trate de una nueva espe- cie. Kelliella abyssicola (Forbes, 1844) la espe- cie viva más abundante, tanto en detri- tos como en contenidos estomacales de estrellas. ALLEN (1979) en un trabajo so- bre bivalvos abisales Atlánticos, co- menta que el 95% de especies de bival- vos de sustratos blandos son Protobran- chia, Septibranchia y Thyasiridae, mientras que para sustratos duros, pre- dominan las especies con biso, como Bathyarca, Limopsis y Dacrydium. Ade- más, encuentra especies pequeñas, ge- neralmente menores de 5 mm. En este trabajo se han encontrado prácticamente los mismos grupos, aunque representa- dos por especies o géneros de menor profundidad y por norma general, de ta- llas mayores. El primer yacimiento Wúrmiense conocido para el litoral catalán fue el Cap de Creus (Girona), descrito por PRUVOT Y ROBERT (1897), posteriormente estudiado por Mars (1958). Otro yaci- miento situado en el Cap de Begur (Girona) ha sido descrito por MARTINELL Y JULIA-BRUGUES (1973). Trabajos poste- riores sobre estos yacimientos Wur- mienses de la costa de Girona son, entre otros, los de VINYAS (1981) y DOMENECH Y MARTINELL (1982). En cuanto a los fondos blandos del litoral catalán, la primera fauna malacológica Wúrmiense fue descrita por MARTINELL, DOMENECH Y DE VILLALTA (1986) para el delta del Ebro (Tarragona). La tanatocenosis Wurmiense que aquí se describe es el origen de algunas de las especies subfósiles que se encuen- tran en el área. Éste es el caso de los Gas- terópodos: lothia fulva (O. E. Múller, 1776), Calliostoma zizyphinum (Linnaeus, 1758), Danilia otaviana (Cantraine, 1835), Capulus ungaricus (Linnaeus, 1758), Trivia multili- rata (Sowerby, 1870), Erato voluta (Mon- tagu, 1803), Ranella olearia (Linnaeus, GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf 1758), Buccinum undatum Linnaeus, 1758, Neptunea contraria (Linnaeus, 1771), y Murexul aradasi (Poirier, 1883 ex Montero- sato ms.), que se encuentran sólo como especies subfósiles en el área de estudio. Entre los Bivalvos, destacan los grandes ejemplares de Modiolula phaseolina (Phi- lippi, 1844), Chlamys islandica (O. F. Mú- ller, 1776), Pseudamussium septemradiatum (O. F. Múller, 1776), Arctica islandica (Lin- naeus, 1767), Glossus humanus (Linnaeus, 1758), Globivenus effosa (Bivona, 1836), Gouldia minima (Montagu, 1803), Pitar me- diterranea Tiberi, 1855, Venus casina Linna- eus, 1758 y Panopea norvegica (Spengler, 1793). Al contrario que los Gasterópodos, muchas de estas especies también se en- cuentran vivas a menos profundidad, o incluso en el mismo sedimento, como es el caso de M. phaseolina. Las especies de Bivalvos que se encuentran en forma subfósil exclusivamente son C. islandica, A. islandica, y P. norvegica, especies de aguas frías que se han extinguido del Me- diterráneo (VINYAS, 1981), y G. effosa, que vive en zonas profundas del Mediterrá- neo (Mar de Alborán). La malacofauna del sedimento Wir- miense de fondos blandos que aquí se- ñalamos, tiene una composición similar a la de los yacimientos de la costa de Gi- rona descritos por los autores menciona- dos más arriba. De todas formas, la ma- yoría de los trabajos anteriores están únicamente basados en especies grandes debido a la metodología de muestreo empleada (Martinell, com. pers.). En los moluscos de “El Parrusset” destaca la ausencia de Buccinum humphreysianum Bennet, 1824 (aunque se han encontrado conchas en otras zonas del Garraf), y de Modiolus modiolus (Linnaeus, 1758), pre- sentes en casi todos los otros yacimien- tos de la costa catalana, así como la au- sencia de Colus islandicus (Gmelin, 1791), abundante en los yacimientos de la costa de Girona. CONTENIDOS ESTOMACALES DE AS- TROPECTEN: Los contenidos estomacales de Equinodermos y otros depredadores han sido investigados por diversos mala- cólogos, como método sencillo para reco- lectar numerosas especies, principalmente micromoluscos de profundidad. Trabajos científicos de este tipo han sido realizados en el Mediterráneo español, concretamente en las islas Baleares (GASULL Y CUERDA, 1974) y en la bahía de Almería (SIERRA, GARCÍA Y LLORIS, 1978), así como en la costa atlántica española, en la Ría de Ares (Galicia) (CRISTOBO-RODRÍGUEZ, TRON- COSO, URGORRI-CARRASCO Y RíOs-LÓPEZ, 1988), entre otros. En este trabajo, el número de espe- cies de Moluscos encontradas en los contenidos estomacales de Astropecten spp. es de 118: 1 Poliplacóforo, 90 Gaste- rópodos, 25 Bivalvos y 2 Escafópodos). Si bien no se ha recopilado información cuantitativa en los muestreos, sí se tie- nen datos cualitativos, de los que se ha podido extraer la siguiente información: - En el Garraf A. aranciacus es más común entre los 40 y los 80 m de pro- fundidad en fondos fangosos, mientras que A. irregularis es abundante en un rango batimétrico mucho más amplio y en todo tipo de fondos. Casi todas las especies halladas en A. aranciacus se encuentran en A. irregularis, aunque en esta última se trata de individuos más pequeños o juveniles. - El Gasterópodo predominante en todas las profundidades estudiadas fue Alvania testae (Aradas y Maggiore, 1843), representando aproximadamente el 50% de individuos encontrados. - El Bivalvo predominante hasta los 80 m fue Timoclea ovata (Pennant, 1777), y a partir de esta profundidad predo- minó Kelliella abyssicola (Forbes, 1844). - En “El Parrusset”, existe una rela- ción directa entre el número de ejempla- res vivos de Nuculidae, Nuculanidae y Yoldiidae encontrados en los contenidos estomacales y los hallados en el sedi- mento, pero ésto no ocurre en el resto de grupos estudiados. - En general, se observó una reduc- ción del tamaño de muchas especies (so- bre todo apreciable en grupos abundan- tes, como Pyramidellidae) a medida que aumentó la profundidad. Este hecho ha sido comentado por ALLEN (1979), que encuentra una disminución de las tasas de crecimiento relacionada con el incre- mento de la profundidad. 89 Iberus, 15 (1), 1997 NUEVAS CITAS PARA EL MEDITERRÁ- NEO ESPAÑOL: Como se señaló anterior- mente, en la lista de especies se identifi- can con un asterisco aquellas que, de acuerdo con la bibliografía, se citan por primera vez en el litoral Mediterráneo español, y con dos asteriscos las que se citan por primera vez para el Meditérra- neo en general. En este apartado no se han tenido en cuenta las citas del estre- cho de Gibraltar como pertenecientes al Mediterráneo. La publicación del catálogo provisio- nal no crítico de los Bivalvos del Medi- terráneo español de BONNIN Y RODRÍ- GUEZ-BABÍO (1990), basado en la biblio- grafía, nos ha facilitado el trabajo de recopilación de citas. En cuanto a los Gasterópodos, no existe ningún catá- logo actualizado, exceptuando algunas monografías de varios grupos, como Cocculiniformia (DANTART Y LUQUE, 1994); Pyramidellidae (PEÑAS ET AL,, 1996); Opistobranchia (CERVERA, TEM- PLADO, GARCÍA-GÓMEZ, BALLESTEROS, ORTEA, GARCÍA, ROs, Y LUQUE, 1988); o el género Mitrella (LUQUE, 1986). Por ello, la constatación de nuevas citas se hace dificultosa y requiere la consulta de numerosos trabajos muy dispersos. En total se reportan 53 nuevas citas para el Mediterráneo español, siendo la mención de Trophon barvicensis y Pleuro- tomella coeloraphe las primeras citas para el Mediterráneo en general. CONCLUSIONES Este estudio confirma la presencia en el Garraf de gran parte de la malaco- fauna típica del Mediterráneo español (exceptuando la zona de Alborán). Sin embargo, se ha constatado la ausencia de algunas especies consideradas comu- nes en áreas cercanas como Scissurella costata d'Orbigny, 1824, Sinezona cingu- lata (O. G. Costa, 1861), Gibbula ardens (von Salis, 1793), G. adansonil (Payrau- deau, 1826), G. rarilineata (Michaud, 1829), G. umbilicaris (Linnaeus, 1758), Ju- jubinus gravinae (Dautzenberg, 1881), Rissoa variabilis (von Muhlfeldt, 1824), Alvania mamillata Risso, 1826, o Mitrella 90 gervillei (Payraudeau, 1826), o por ejem- plo, la presencia de sólo 2 skenéidos pertenecientes a dos especies diferentes, y que no se haya encontrado ningún ejemplar de esta familia en fondos de maérl, en los que son comunes en otras localidades. Finalmente, podemos concluir que la comarca del Garraf es una zona rica en moluscos marinos debido a las particu- laridades de sus fondos. Prueba de ello son las 53 nuevas citas para el Medite- rráneo español. Muchas de estas espe- cies viven probablemente en otras zonas de nuestras costas, pero, debido a su pequeño tamaño, podrían haber pasado desapercibidas. Los autores consideran que la prospección con medios adecua- dos de las zonas más profundas de esta comarca podría contribuir a ampliar el conocimiento de la malacofauna del litoral español, y que sería interesante recolectar vivas algunas de las especies que aquí se han encontrado, de las que no se conoce el animal. AGRADECIMIENTOS Los autores quieren mostrar su agra- decimiento a los hermanos J. y F. Ayza, pescadores ya jubilados, que nos han proporcionado numerosas muestras, así como comentarios sobre los tipos de fondos, etc., y a M. Roca y a sus hijos Je- sús y Pavel que nos facilitaron desinte- resadamente numerosos ejemplares de asteroideos y los sedimentos de “El Pa- rrusset”. Sin ellos, este trabajo no hu- biera podido llevarse a cabo. También queremos agradecer a A. Tubau el ha- bernos cedido información de su colec- ción. Especialmente agradecemos a C. Palacín y M. Ballesteros (Dpt. de Biolo- gia Animal, Universitat de Barcelona) y a J. Templado (Museo Nacional de Cien- cias Naturales, Madrid) sus comentarios sobre el manuscrito original. Las correc- ciones y comentarios constructivos de S. Gofas (Muséum National d'Histoire Na- turelle, Paris) y de otro revisor anónimo han contribuido notablemente en la me- jora de este trabajo. También agradecer a L. Dantart (Dpto. de Biologia Animal, GIRIBET Y PEÑAS: Fauna malacológica del litoral del Garraf Universitat de Barcelona) y C. Salas (Dpto. de Biología Animal, Universidad de Málaga) por la determinación de al- gunos ejemplares. A. Warén (Swedish Museum of Natural History, Stockholm) y S. Gofas realizaron comentarios sobre algunas especies, y Y. -R. Kim (Ameri- can Museum of Natural History, New BIBLIOGRAFÍA AARTSEN, J. J. VAN, MENKHORST, H. P. M. G. Y GITTENBERGER, E., 1984. The marine Mo- llusca of the Bay of Algeciras, Spain, with general notes on Mitrella, Marginellidae and Turridae. Basteria, suppl. 2: 1-135. ALLEN, J. A., 1979. The adaptations and radia- tion of deep-sea bivalves. Sarsia, 64: 19-27. ALLEN, J. A. Y HANNAH, F. J., 1989. Studies on the deep-sea Protobranchia: the subfamily Ledellinae (Nuculanidae). Bulletin of the British Museum of natural History (Zoology Series), 55 (2): 123-171 ASENSI, J. M., 1984. Invertebrats del Mar de Sit- ges. Diputació de Barcelona, Servei del Medi Ambient, Barcelona, 54 pp. BALLESTEROS, M., 1977. Sobre Spurilla neapoli- tana Delle Chiaje (1824) y Berghia verruci- cornis A. Costa (1864), dos Aeolidacea (Gastropoda: Opistobranchia) recolectados en Cubellas (Barcelona). Publicaciones del Departamento de Zoología (Barcelona), 2: 7-12. BALLESTEROS, M., 1978. Contribución al cono- cimiento de la fauna bentónica de Cube- llas. Publicaciones del Departamento de Zoolo- gía (Barcelona), 3: 11-23. BALLESTEROS, M., 1984. Adición a la fauna de Opistobranquios de Cubelles (Tarragona). Miscelánea Zoológica, 8: 41-49. BALLESTEROS, M., LLERA, E. M. Y ORTEA, J. A., 1984. 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ANA Í hoi 8 ik ay 19 O Sociedad Española de Malacología Iberus, 15 (1): 95-111, 1997 Scaphopoda from the Spanish coasts Escafópodos de las costas españolas Gerhard STEINER* Recibido el 20-IX-1996. Aceptado el 18-X1-1996 ABSTRACT The scaphopod molluscs collected at 29 stations of the Fauna Iberica projects | - 1Il along the entire Spanish mainland coast and the Balearic Islands belong to 12 species from 7 genera and 5 families. These species are listed together with their synonyms, original des- criptions, and geographic and bathymetric ranges in this material and from the literature. None of the species are found to extend either range in these samples. On the other hand, some otherwise common scaphopods are not represented. Some of the species have extensive fossil records, and in Entalina tetragona and Antalis inaequicostata, problems of synonymy and historical biogeography are discussed. RESUMEN En el presente trabajo se relacionan aquellas especies de escafópodos encontradas en 29 estaciones de las costas españolas de la península Ibérica y de las islas Baleares durante las campañas FAUNA |-1II. En total se han hallado 12 especies pertenecientes a 7 géneros y 5 familias. Para cada especie se proporciona una lista de sinónimos, la descripción ori- ginal y los rangos de distribución geográfica y batimétrica, obtenidos a partir de este material y de la bibliografía. Los datos que proporcionan estas muestras no permiten extender rangos de distribución de ninguna de las especies. Por otra parte, algunas de las especies comúnes de escafópodos en el área no están aquí representados. Algunas de las especies tienen registros fósiles extensos, y en Entalina tetragona y Antalis inaequi- costata se discuten problemas relacionados con su sinonimia y biogeografía histórica. KEY WORDS: Scaphopoda, systematic biogeography, Iberian Peninsula and Balearic Islands PALABRAS CLAVE: Escafópodos, sistemática biogeografía, península Ibérica e islas Baleares. INTRODUCTION The main purpose of this paper is the systematic treatment of Scaphopoda collected by recent sampling cruises of the Fauna Ibérica programme along the Spanish coasts. The cruises Fauna I (July, 1989), Fauna II (June - July, 1991) and Fauna lll (June - July, 1994) covered the Sea of Alborán, the Gulf of Cádiz, the Bay of Biscay, the Atlantic coast off South Galicia, and the waters around the Balearic Islands. Scaphopod speci- mens are represented in 29 stations of these cruises. A list of stations of the “Fauna I” cruise is already published (TEMPLADO, GUERRA, BEDOYA, MORENO, REMÓN, MALDONADO AND RAMOS, * Institute of Zoology, University of Vienna. Althanstr. 14, A-1090 Vienna, Austria. 95 Iberus, 15 (1), 1997 1993); publications with the stations of the remaining cruises are in preparation (Iemplado, pers. comm.). Sources of primary faunistic information on Scap- hopoda are reports of the marine expe- ditions “Travailleur” and “Talisman” (LocarD, 1898), “Porcupine”, “Valo- rous” and two smaller cruises (JEFFREYS 1870, 1877, 1882), and ALZURIA (1986, 1987). Faunistic and systematic accounts of the Eastern Pyrenean Seas were made by BucQuoY, DAUTZENBERG AND DOLLFUS (1882) and Mars (1965). The only synopsis of Iberian and Balearic SYSTEMATIC ACCOUNT The samples revealed 12 species from 7 genera and 5 families. Both orders of Scaphopoda, Dentaliida and Gadilida are represented, with 8 and 4 species respec- tively. Each species is listed with sy- nonyms, original description, type locality, molluscs to date (HIDALGO, 1917) also includes Scaphopoda. The fundamental monography of PILSBRY AND SHARP (1897), the comprehensive studies of CAPROTTI (1965, 1968, 1979), and the classification of higher taxa Of STEINER (1992, 1996) and SCARABINO (1995) form the basis of the systematic treatment. Complementary information on syno- nyms is drawn from MONTEROSATO (1875) and STORK (1934). The material studied in this paper is deposited in the Museo Nacional de Ciencias Naturales, Madrid. distribution in the present material (with distribution maps) and as reported in the literature, and with the earliest fossil oc- currence. The numbers of empty shells (e) and of shells with soft body (b) are given for each station. Order DENTALIDA Da Costa, 1776 Family DENTALIDAE Gray, 1847 Genus Antalis H. and A. Adams, 1854 Antalis agilis (M. Sars, 1872) (Figs. 1, 4A) Synonyms Dentalium incertum Philippi 1844, Enum. Moll. Sicil. 1: 207, non Deshayes 1825. Dentalium abyssorum var. agilis Jeffreys 1870, Ann. Mag. N. Hist. Ser. 4 VI: 74. Dentalium agile M. Sars 1872, Some remarkable forms etc., Christiania 1872: 34. Dentalium fusticulus Brugnone 1876, Misc. Malac. 11: 21. Dentalium vagina Jeffreys 1877, Ann. Mag. N. Hist. Ser. 4, XIX: 155. Antalis agilis (M. Sars): G. O. Sars 1878, Moll. Reg. Arct. Norv., Christiania 1878: 102. Dentalium (Antalis) calabrum Crema 1910, p. 68. Original description: Shell slender, very narrow, slightly curved, almost straight, gradually attenuated towards the apex. White, faint luster, posterior part fre- quently darker. Apex very narrow, obli- quely truncated, with a tolerably deep inci- sion and a short, hardly protruding sup- plementary tube. Shell surface with cir- cular growth lines, rarely longitudinally striated in the posterior part, the striae being little distinct and never prominent ribs. [... ] Largest shell 58 mm long and 4 mm in diameter, 1 mm at the apex. 96 Type locality: Lofoten Islands, North Atlantic, at 360-540m. Present material: 5 stations; Gulf of Cádiz, 500-546 m (76A: 1b, 20e; 77A: 3b); Cape Finisterre, 129-133 m (91A: 1b); Biscay, 540-1025 m (124A: 2b, 9e; 1594: 2b, 14e). Reported distribution: North Atlan- tic: Portugal to Lofoten, Halifax to Cuba, Gulf of Mexico, Azores; Mediterranean; ?Red Sea; 60-5000 m. STEINER: Scaphopoda from the Spanish coasts Earliest fossil: Pliocene. Remarks: The largest specimen from station 159A is 65 mm long, which exce- eds indications of SARS (1872), PILSBRY AND SHARP (1897) and CAPROTTI (1965), but matches those of LOCARD (1898). There are no shells with apical slits in the present material, but most of the speci- mens are empty shells and fragments. The shell surface in the apical region may be eroded by boring organisms even in live animals. The shell then has a chalky and often crackled aspect, as was also remarked by Sars (1872). Young in- dividuals are often more or less dis- tinctly ribbed near the apex. The ribs intercalate to about 20 in number and then gradually become obsolete. The younger parts of the shell (towards the anterior opening) with an intact surface are glossy and bear closely spaced growth lines. LOCARD (1898) distinguis- hed a number of varieties according to size, curvature and sculpture. Antalis pa- norma (Chenu, 1842-47), similar in size and shape to A. agilis and reported from the Mediterranean and the Bay of Biscay, differs in being more curved and having a more solid shell with 12 narrow but pronounced primary ribs. Some of the shells from station 76A have bore holes of naticid gastropods being, apart from different fishes, important predators of scaphopods in the Mediterranean. Antalis entalis (Linné, 1758) (Figs. 1, 4B) Synonyms Dentalium entalis Linné 1758, Syst. Nat. (10): 785. Dentalium entalum L.: Blainville 1819, Dict. Sc. Nat. XUl: 70. Dentalium labiatum Brown 1827, 111. Conch. Gr. Brit. and Irel.: pl. 1, fig. 4. Dentalium striolatum Stimpson 1851, Proc. Bost. Soc. Nat. Hist. IV: 114. non Jeffreys, Watson, Sars, Risso. Entalis striolata (Stimpson 1851): Gould-Binney 1870, Invert. of Mass.: 266. Original description: Shell smooth, moderately curved, continuous, not fractured. Type locality: Atlantic Ocean. Present material: 2 stations; La Coruña, 151-152 m (101A: le); Biscay, 119-122 m (112DH: 2e). Reported distribution: North Atlan- tic from Spain north to Spitzbergen and Maine, Massachusetts to Bay of Fundy; 6-3500 m. Earliest fossil: Pliocene. Remarks: The shell is up to 42 mm long, solid, white, sometimes glossy, mode- rately curved but in the apical region, the wider, anterior part of the shell being only slightly curved. The shell surface is smooth, very fine longitudinal striae may be present in the apical region only. Towards the anterior opening growth lines become more distinct. There may be a shallow apical notch on the convex side. This species is infrequently cited from the Mediterranean as well, due to confusion with the apically striated An- talis vulgaris. However, HIDALGO (1917) retains Gibraltar and Mataró / Catalunya as localities for A. entalis. Antalis dentalis (Linné 1766) (Figs. 1, 4C) Synonyms Dentalium dentalis Linné 1766, Syst. Nat. XII: 1263. Dentalium dentale L.: Locard 1886, Ann. Soc. Agricult., Lyon Ser. 5, IX: 145. Dentalium linnaeum Locard 1886, Ann, Soc. Agricult., Lyon Ser. 5, IX: 145. Dentalium mutabile Dóderlin in Hórnes 1856, Abhandl. K. -K. Geol. Reichsanst. Il: 654. 2 Iberus, 15 (1), 1997 Original description: Shell striated, moderately curved, fractured. Type locality: Mediterranean Sea. Present material: 1 station; Gulf of Cádiz, 13-15 m (714: 1e). Reported distribution: Mediterra- nean Sea, East Atlantic from Galicia to Cape of Good Hope (?), Azores, Canary Islands; 0-300 m. Earliest fossil: Miocene. Remarks: This rather small species is up to 24 mm long (13 mm from station 71A), mostly white, only the apex sometimes with a rose tinge. There are about 10 sharp and narrow primary ribs, becoming doubled by intercalation towards the anterior opening. The secondary ribs are of about the same height as the primary ribs. The intercos- tal spaces are much wider than the ribs, and smooth except for widely spaced growth lines. Antalis dentalis is often confused with A. inaequicostata (see below). Antalis inaequicostata (Dautzenberg, 1891) (Figs. 1, 4D) Synonyms Dentalium dentalis Lamarck 1818, Anim. sans vert. V: 344; Deshayes 1825, Anat. et Monogr. du genre Dentale: 33; Risso 1826, Hist. Nat. Europ. Merid. IV: 398; Philippi 1836, Enum. Moll. Sicil. 1: 243; Jeffreys 1870, Ann. Mag. Nat. Hist. VI: 10; Monterosato 1872, Not. Int. alle conch. Medit.: 28; non Linné 1766. Dentalium fasciatum Lamarck 1818, Anim. sans vert. V: 343; non Gmelin 1790 Dentalium pseudo-antalis Scacchi 1836, Catal. Conch. Regni Neap.: 17; non Lamarck 1818 Dentalium novem-costatum Réquien 1848, Coq. de Corse: 90; non Lamarck 1818. Dentalium novemcostatum var. tenuis Monterosato 1878, Enum. et Sinon.: 16; non Lamarck 1818. Dentalium novemcostatum Réquien: Monterosato 1884, Nom. Gen. e Spec.: 31; non Lamarck 1818. Dentalium alternans Bucquoy, Dautzenberg and Dollfus 1891, Moll. Mar. Roussillon 1: 561; non Chenu 1842. Dentalium inaequicostatum Dautzenberg 1891, Mem. Zool. Soc. France 1891: 53. Antale novemcostatum (Réquien): Sacco 1897, Moll. terr. terz. Piemonte e delle Liguria XXI: 104. Dentalium (Antalis) inaequicostatum B. D. D.: Pilsbry and Sharp 1897-98, Man. Conch. XVII: 52. Caprotti 1965, Atti. Soc. Ital. Sci. Nat. Milano 105: 343. Dentalium novemcostatum var. inaequicostata Fantinet 1959, Serv. Carte Geol. Algérie 1: 46. Dentalium (Antalis) novemcostatum Réquien: Caprotti 1961, Atti. Soc. Ital. Sci. Nat. Milano 100: 353. Dentalium (Antalis) mutabile inaequicostatum Dautzenberg: Caprotti 1979, Boll. Malacol. Milano 15: 233. Original description: Shell solid, opaque, slightly to moderately curved, straightening towards the anterior aper- ture. Sculpture of 9 or 10 primary ribs alternating with same number of wider and less protruding secondary ribs. All ribs become obsolete towards the ante- rior. Numerous transverse growth lines, sometimes with irregular fractures or interruptions. Anterior shell aperture slightly polygonal. Posterior aperture truncated, polygonal, with an oval, short central pipe. No slit or notch. Colour light rose, more intense at the posterior end, transversal bands of lighter and darker colour. Shell 35 mm long, 5 mm at anterior aperture. 98 Type locality: Mediterranean Sea. Present material: 4 stations; Gulf of Cádiz, 13-28 m (44A: 2e; 66A: 9b, 3e; 69A: le; 71A: about 10b, many empty shells). Reported distribution: Mediterra- nean from Greece to Algeria; 5-120m. Earliest fossil: Miocene (BUCQUOY ET AL., 1886) or Pliocene (CAPROTTI, 1979). Remarks: This species is extremely variable in its longitudinal sculpture causing considerable confusion in studies of both fossil and recent scapho- STEINER: Scaphopoda from the Spanish coasts MW Antalis agilis O Antalis dentalis Ba Antalis novemcostata A Antalis entalis O Antalis inaequicostata e Antalis vulgaris Figure 1. Localities where Antalis agilis, A. entalis, A. dentalis, A. inaequicostata, A. novemcostata, and A. vulgaris were collected. Figura 1. Localidades donde se encontraron las especies Antalis agilis, A. entalis, A. dentalis, A. inae- quicostata, Á. novemcostata y A. vulgaris. pods. Small individuals of this species closely resemble Antalis dentalis. The latter differs, however, in having conspi- cuous transversal striae between the ribs, the ribs themselves are less acute, and the intercostal space is not as wide. If PILSBRY AND SHARP (1897-98, p. 52) describe “... 9-12 strong primary ribs towards the apex, narrower than their intervals... “ for A. inaequicostata, they obviously did not consider direct com- parison with A. dentalis. The atlantic A. novemcostata is stouter and has more conspicuous transverse striae. Many of the empty shells shows cha- racteristic signs of sipunculid occupa- tion. Phascolion strombus (Montagu, 1804) is known to close the anterior ope- nings of scaphopod and other mollusc shells by agglutinations of sediment, leaving open only a small tube for their introvert (TÉTRY, 1959). 99 Iberus, 15 (1), 1997 Antalis novemcostata (Lamarck, 1818) (Figs. 1, 4E) Synonyms Dentalium novemcostatum Lamarck 1818, Anim. sans vert., V: 344. Dentalium dentalis Risso 1826, Hist. Nat. Europ. Mérid., IV: 398. Dentalium dentale Risso: Weinkauff 1862 (partim), J. Conch., X: 364. Antale novemcostatum (Lamarck): Sacco 1896, Boll. Mus. Zool. Anat. Comp. Univ. Torino, XI: 97. Dentalium (Antalis) novemcostatum (Lamarck): Pilsbry and Sharp 1897-98, Man. Conch., 17: 51 Original description: Shell small, greenish-white, with nine ribs, subde- cussate transverse striae. Type locality: Atlantic coast near La Rochelle. Present material: 1 station, Gulf of Cádiz, 110-112 m (694: 1e). Reported distribution: East Atlantic from La Rochelle to South Spain; 20-300 m. Earliest fossil: Pliocene. Remarks: The rather stout shell is up to 32 mm long and has 8 to 10 rounded ribs decreasing in heigth towards the ante- rior opening. The intercostal spaces are more concave than in the other Antalis species of the region and show faint lon- gitudinal striae. Transverse striae are not always developed, although BUCQUOY ET AL. (1886) and CAPROTTI (1965) take the strong transverse sculpture as an impor- tant character to distinguish A. novem- costata from the mediterranean A. inae- quicostata. The apex often has a plug with a central pipe in larger specimens. Antalis novemcostata seems to be living on the european Atlantic coast only, although HIDALGO (1917) lists several mediterranean locations for this species. This seems to be due to misiden- tifications of A. inaequicostata (CAPROTTI, 1961, 1965; Mars, 1965). According to CAPROTTI (1965), the extremely rare A. novem-costatum from the Italian Pliocene could be intermediate between A. inae- quicostata and A. novemcostata. Antalis vulgaris (Da Costa, 1778) (Figs. 1, 4F) Synonyms Dentale vulgare Da Costa 1778, Brit. Conch.: 24. Dentalium fasciatum Gmelin 1791, Syst. Nat., 13: 3737. Dentalium striatum Montagu 1803, Test. Brit., II: 492, non Born 1780, Test. Mus. Caes. Vindob.: 431. Dentalium tarentinum Lamarck 1818, Anim. sans vert., V: 345. Forbes and Henley 1853, Hist. Brit. Moll., II: 451. Sowerby 1860, Thes. Conch., 11: 100. Jeffreys 1882, Brit. Conch., II: 195. Clessin 1896, Conchyl. Cab.: 3. Dentalium politum Blainville 1819, Dict. Sci. Nat., XI: 70. Turton 1819 (partim), Conch. Dict. Brit. Sh.: 38. Dentalium labiatum Turton 1819 (partim), Conch. Dict. Brit. Sh.: 38. Brown 1827, Illustr. Conch. Gr. Brit.: 117. Dentalium striolatum Risso 1826, Hist. Nat. Europ. Mérid., IV: 398. Dentalium multistriatum Risso 1826, Hist. Nat. Europ. Mérid., IV: 398, non Deshayes 1825, Anat. et Monogr. du genre Dentale. Dentalium affine Biondi 1859, Atti Accad. Gioenia Sci. Nat. (2), XIV: 120. Original description: Dentalium with a slender, smooth, glossy, subar- cuated shell, tapering to a small point, pervious: sometimes marked with a few circular wrinkles or annulations: colour white or yellowish. Length an inch and 100 a half [38 mm]; diameter at the larger end two-tenths of an inch [5 mm]; and one fourth as much [1.25 mm] at the smaller end. [... ] A variety is marked with dusky bands; and sometimes a little striated towards the point. STEINER: Scaphopoda from the Spanish coasts E Fissidentalium capillosum O Entalina tetragona Xx Dischides politus A Episiphon filum O Pulsellum lofotense e Cadulus jeffreysi Figure 2. Localities where Fissidentalium capillosum, Episiphon filum, Entalina tetragona, Pulsellum lofotense, Dischides politus, and Cadulus jeffreysi were collected. Figura 2. Localidades donde se encontraron las especies Fissidentalium capillosum, Episiphon filum, Entalina tetragona, Pulsellum lofotense, Dischides politus y Cadulus jeffreysi. Type locality: British shores, espe- cially Scilly Islands, Cornwall, Devons- hire, Hampshire. Present material: 5 stations; Biscay, 119-122 m (112DH: 1e); Balearic Islands, 5-59 m (190B: 2e; 192A: 1b; 203B: 1e; 258B: 3b). Reported distribution: Mediterranean Sea and East Atlantic Ocean; 5-1100 m. Earliest fossil: Miocene. Remarks: The shell is up to 60 mm long, white with a rose-coloured apex, rather broad, and moderately curved in the posterior half. There are about 30 longitudinal striae near the apex, oblite- rating gradually towards the anterior opening. The apical opening is entire and may have a plug with a short central pipe. 101 Iberus, 15 (1), 1997 Genus Fissidentalium Fischer, 1885 Fissidentalium capillosum (Jeffreys, 1876) (Figs. 2, 4G) Synonyms Dentalium capillosum Jeffreys 1876, Proc. Roy. Soc., 25: 185. (nomen nudum) Dentalium capillosum Jeffreys: Jeffreys 1877, Ann. Mag. Nat. Hist. Ser. 4, 19: 153. Dentalium (Fissidentalium) capillosum Jeffreys: Pilsbry and Sharp 1897-98, Man. Conch., 17: 77. Original description: Shell tapering to a fine point, slightly curved, rather solid, opaque, and mostly lusterless; sculpture: numerous and sharp (not rounded) lon- gitudinal striae, some of which are inter- mediate and smaller than the rest; they disappear towards the posterior or narrow end, which is quite smooth and glossy for a quarter of an inch [6.4 mm]; colour whitish; margin at the posterior end having a short and narrow notch. L [length]: 1.4 [35.6 mm]. B [maximum diameter]: 0.15 [3.8 mm]. (...) This appears to attain a size considerably exceeding that given in the above description, as fragments measure nearly 0. 4 inch [10 mm] in breadth. Type locality: North Atlantic, Valo- rous st. 12, 13, 16; 1242-3213 m. Present material: 1 station; Biscay, 925-1025 m (159A: 5e). Reported distribution: North Atlan- tic, Caribbean Sea to Portugal, Azores to Hebrides; 400-3500 m. Earliest fossil: No fossil record. Remarks: This species can be 81 mm long. The shell is white or grey and may be somewhat eroded. There are about 65 fine ribs throughout most of the length. Pilsbry and SHARP (1897-98) supplement JEFFREYS' (1877) description saying that the ribs are sharply cut but rounded on the top. This is particularly obvious in the anterior part of the shell where the ribs become wider. Family GADILINIDAE Chistikov, 1975 Subfamiliy EPISIPHONINAE Chistikov, 1975 Genus Episiphon Pilsbry and Sharp, 1897-98 Episiphon filum (Sowerby, 1860) (Figs. 2, 4H) Synonyms Dentalium filum Sowerby 1860, Thes. Conch., III: 89. Dentalium gracile Jeffreys 1870, Ann. Mag. Nat. Hist. (4), VI: 74. Fischer 1873, Journ. Conchyl.: 140. Pseudantalis filum (Sowerby): Monterosato 1884, Nom. Gen. Spec. Conch. Medit.: 33. Dentalium rufescens Weinkauff 1868 (partim), Conch. Mittelm., Il: 420. Original description: Shell slender, very narrow, thin, finely pointed, mantle reddish brown, apex entire. Type locality: Gibraltar. Present material: 2 stations; Biscay, 104-132 m (152A: 2b, le; 153A: 2b). Reported distribution: North Atlan- tic from Florida to Cape Hatteras, Algeria to Biscay; Mediterranean Sea from Aegean to Gibraltar; 20-4784 m. 102 Earliest fossil: Miocene. Remarks: This is a very characte- ristic species, being very narrow, hardly tapering and almost straight. The length is up to 13 mm. The shell is white and glossy, semitransparent and extremely fragile. The sculpture consists of growth lines only. Typical for the genus is a long pipe at the apex continuous with the shell. It may be wanting because of its fragi- lity. STEINER: Scaphopoda from the Spanish coasts Figure 3. Bathymetric ranges of species reported in literature (shaded) and in the present material (black). Figura 3. Rangos batimétricos de las especies: datos bibliográficos (gris), datos del presente material (negro). Order GADILIDA Starobogatoy, 1982 Suborder ENTALIMORPHA Steiner, 1992 Family ENTALINIDAE Chistikov, 1979 Subfamily ENTALININAE Chistikov, 1979 Genus Entalina Monterosato, 1872 Entalina tetragona (Brocchi, 1814) (Figs. 2, 5A) Synonyms Dentalium tetragonum Brocchi 1814, Conch. foss. subapen., Milano: 627. Dentalium quinquangulare Forbes 1844, Rep. Brit. Ass. Adv. Sci. for 1844: 188. Siphonodentalium pentagonum M. Sars 1865, Forh. Videsk. Selsk. Christiania 1864: 307. Dentalium quinquangulatum Reeve 1872, Conch. Icon.: pl. 5, fig 45. Siphonodentalium quinquangulare (Forbes): Jeffreys 1867, Ann. Mag. Nat. Hist. Ser. 3, XX: 251. Weinkauff 1868, Conch. Mittelm., 1: 421. Locard 1886, Ann, Soc. Agricult., Lyon Ser. 5, IX: 149. Dautzenberg 1891, Mem. Soc. Zool. France, IV: 609. Friele and Grieg 1901, Norv. N. Atlant. Exp. etc., Christiania 1901, Mollusca III: 50. Siphonentalis tetragona (Brocchi): G. O. Sars 1878, Moll. Reg. Arct. Norv., Christiania 1878: 105. 103 Iberus, 15 (1), 1997 Siphodentalium tetragonum (Brocchi): Norman 1879, J. Conch., London, 2: 49. Entalina tetragona (Brocchi): Monterosato 1880, Bull. Soc. Malac. Ital., VI: 64. Caprotti 1961, Atti Soc. Ital. Sci. Nat. Mus. Civ. Stor. Nat. Milano, Vol. C, IV: 356. Siphodentalium quinquangulare (Forbes): Jeffreys 1882, Proc. Zool. Soc. London, 1882: 662. Siphonentalis quinquangularis (Forbes): Carus 1889, Prodr. faunae Medit., 11: 176. Pulsellum quinquangulare (Forbes): Norman 1893, Ann. Mag. Nat. Hist. Ser. 6, XII: 344, 362. Entalina quinguangularis (Forbes): Hidalgo 1917, Trab. Mus. Nac. Cienc. Nat. Ser Zool. 30: 306. Chistikov and Sagaidachniy 1982, Zool. Zhurn., 60: 38. Original description: Shell four- angled, finely longitudinally striated, sides weakly carinated. Type locality: Pliocene of Italy (BROCcHI, 1814), Aegean Sea (FORBES, 1844). Present material: 6 stations; Medite- rranean, 1001-1005 m (247A: le); Sea of Alborán, 276-306 m (15A: 2e); Gulf of Cádiz, 500-546 m (76A: 3e; 77A: 2e); Galicia, 81-84 m (86DL: 1b), Biscay, 540- 543 m (124A: 5b, 15€e). Reported distribution: Mediterra- nean, East Atlantic from Biscay to Nort- hern Norway; 10-2664 m. Earliest fossil: Miocene. Remarks: The shell is strongly curved, at least in the apical half. It has five primary ribs, four of them forming almost right angles, the fifth rib on the midline of the concave side forms an obtuse angle. In specimens larger than about 10 mm, 3 - 25 secondary ribs gra- dually appear, the pentagonal form of the cross-section smoothing out to become subcircular. The anterior opening is oblique, the apex simple and entire. The shells may be up to 93 mm long but are usually around 15 mm. al There is some confusion in the Euro- pean species and the use of the names te- tragona Brocchi, pentagona Sars and quin- quangularis Forbes. BROCCHI (1814) des- cribed tetragona as a Pliocene fossil from Piemont and the Vienna Basin. FORBES (1844) described the extant species from the Aegean Sea as quinquangularis. MiI- CHAEL SARS (1865) described pentagona from the Norwegian coasts, without refe- rring to either Brocchi or Forbes. His son, G. O. SArs (1878) considers pentagona and quinquangularis as janior synonyms of te- tragona. MONTEROSATO (1880) comes to the same conclusion claiming the fossil and recent mediterranean species being iden- tical. JEFFREYS (1870, 1882), however, is of different opinion and gives priority to quinquangularis for the recent Atlantic and Mediterranean form. PILSBRY AND SHARP (1897-98), LOCARD (1898) and FRIELE AND GRIEG (1901) agree with this view. CAPROTTI (1968), finally, comparing re- cent specimens with the type material, confirmes the synonymity of quinquangu- laris and tetragona, stressing the priority of Brocchi's name. Later, CHISTIKOV AND SAGAIDACHNIY (1982), however, not only separate tetragona and quinquangularis, but also split the latter into the Mediterranean quinguangularis and the Atlantic pentagona on grounds of shell and radula characters. However, they do not include fossil spe- cimens in their study. Suborder GADILIMORPHA Steiner, 1992 Family PULSELLIDAE Scarabino in Boss, 1982 Genus Pulsellum Stoliczka, 1868 Pulsellum lofotense (M. Sars, 1865) (Figs. 2, 5B) Synonyms Siphonodentalium lofotense M. Sars 1865, Forh. Videsk. Selsk. Christiania 1864: 297. Siphonentalis lofotensis (M. Sars): G. O. Sars 1878, Moll. Reg. Arct. Norv., Christiania 1878: 104. Mon- terosato 1884, Nom. Gen. Spec. Conch. Medit.: 33. 104 STEINER: Scaphopoda from the Spanish coasts Figure 4. A: Antalis agilis, B: Antalis entalis, C: Antalis dentalis, D: Antalis inaequicostata; E: Antalis novemcostata; E: Antalis vulgaris, G: Fissidentalium capillosum; H: Episiphon filum. Scale bars, A, G: 10 mm; B-F: 2 mm; H: 1 mm. Figura 4. A: Antalis agilis; B: Antalis entalis; C: Antalis dentalis; D: Antalis inaequicostata; E: Antalis novemcostata; F: Antalis vulgaris; G: Fissidentalium capillosum; A: Episiphon filum. Escalas, A, G: 10 mm, B-E: 2 mm; H: 1 mm. Iberus, 15 (1), 1997 Siphodentalium lofotense M. Sars: Jeffreys 1882, Ann. Mag. Nat. Hist., Ser. 5, XI: 395. Siphonodentalium (Pulsellum) lofotense M. Sars: Pilsbry and Sharp 1897-98, Man. Conch., 17: 138. Siphonodentalium lofotensis M. Sars: Stork 1934, Thalassia, 1: 10. Pulsellum lofotensis (M. Sars): Emerson 1962, Journ. Paleontol. 36: 475. Original description: Shell smooth, moderately curved, anteriorly wide and tapering to the posterior, white, walls transparent or semitransparent, thin, shiny, very fine and dense obliquely transverse growth lines well visible, posterior shell margin entire [plain]. Length 5-6 mm, basal width 0.66 mm, apical about 0.33 mm. [Description of soft body omitted.] Type locality: Lofoten Isl., Norway; 90-216 m. Present material: 4 stations; Gulf of Cádiz, 535-546 m (76A: 3e); Galicia, 80- 120 m (1684: 3e; 171A: 3e); Biscay, 129- 132 m (153A: 5b, about 20e). Reported distribution: Mediterra- nean Sea; North Atlantic from Spain to Finmark, Ireland, New England; 26- 3500 m. Earliest fossil: Pliocene. Remarks: The small shell is rather fragile in the anterior third and easily breaks into cylindric fragments upon handling. Breakage occurs along the oblique growth lines. The anterior ope- ning is circular as the apical opening but slightly oblique. Most live animals have perfectly transparent shells, empty shells are opaque. The apical rim is al- ways entire without notches or lobes, although JEFFREYS (1882) mentions spe- cimens with regularly jagged tips. This may have been a siphonodentaliid spe- cies, also because he did not see a bul- bous larval shell, which is well develo- ped in Pulsellum lofotense (Steiner, 1995). Family GADILIDAE Stoliczka, 1868 Subfamily SIPHONODENTALIINAE Simroth, 1894 Genus Dischides Jeffreys, 1867 Dischides politus (Wood, 1842) (Figs. 2, 5C) Synonyms Ditrupa polita Wood 1842, Ann. Mag. Nat. Hist., 9: 459. Dentalium coarctatum Philippi 1844, Enum. Moll. Sicil., UI: 208, non Lamarck 1818, Anim. sans Vert., 5: 346. Dentalium laevigatum de Rayneval, Hecke and Ponzi 1854, Cat. Foss. Mont Mario, Versailles, non Schlotheim 1830. Dentalium bifissum Jeffreys 1867, Ann. Mag. Nat. Hist., Ser. 3, XX: 251. Weinkauff 1868, Conch. Mittelm., 1: 421. Monterosato 1884, Nom. Gen. Spec. Conch. Medit.: 34. Dischides olivi Jeffreys 1870, Ann. Mag. Nat. Hist., Ser. 4, VI: 73. Dischides bifissus (Jeffreys): Jeffreys 1882, Proc. Zool. Soc. London, 1882: 663. Cadulus politus (Wood): Pilsbry and Sharp 1897-98, Man. Conch., 17: 144. Stork 1934, Thalassia, 1: 9. Original description: Shell slightly arcuated, thin, smooth, subcylindrical; anterior opening plain, posterior cleft, bilateral, with unequal terminations. The body of the [... ] shell is not inflated or enlarged like that of Dentalium gadus, but has the posterior opening laterally cleft, somewhat resembling that of Den- talium coarctatum Deshayes [... ] but the dorsal part of the posterior end of this 106 fossil is produced beyond the edge beneath and rounded, the ventral edge is shorter and truncated, an enamel-like polish covers the exterior, and was pro- bably when inhabited subhyaline, but is now opaque. Length half an inch [12.7 mm] nearly. Type locality: Coralline England (Pliocene). Crag, STEINER: Scaphopoda from the Spanish coasts Figure 5. A: Entalina tetragona; B: Pulsellum lofotense, C: Dischides politus, D: Cadulus jeffreysi. Scale bars 1 mm. Figura 5. A: Entalina tetragona; B: Pulsellum lofotense; C: Dischides politus; D: Cadulus jeffreysi. Escalas 1 mm. Present material: 1 station; Gulf of Cádiz, Trafalgar, 34 m (58A: 1e). Reported distribution: Northeast Atlantic from Marocco to Biscay, Medi- terranean; 9-324 m. Earliest fossil: Pliocene. Remarks: The recent members of this species attain maximum lengths of 7 mm. The shell is white, glossy, mode- rately curved and only slightly tapering. The sculpture consists of growth lines only. The greatest diameter of the shell lies just behind the anterior opening. The posterior opening has two lateral notches producing a dorsal and a ventral lobe. Tubes of the serpulid poly- chaete Ditrupa sp. must not be confused with Dischides politus. The polychaete tubes are often reddish brown in colour, the outer layer has a semitransparent aspect. They are sharply constricted at the anterior opening and lack growth lines. Dischides has repeatedly been chan- ging between subgenus and genus status. It was recently confirmed in its generic rank and transferred to the sub- family Siphonodentaliinae by SCARA- BINO (1995). 1107 Iberus, 15 (1), 1997 Subfamily GADILINAE Stoliczka, 1868 Genus Cadulus Philippi, 1844 Cadulus jeffreysi (Monterosato, 1875) (Figs. 2, 5D) Synonyms Helonyx jeffreysi Monterosato 1875 Cadulus jeffreysi (Monterosato): Jeffreys 1882, Verrill 1882, Pilsbry and Sharp 1897-98, Muus 1959 Cadulus propinquus Verrill 1885, non G. O. Sars Cadulus subfusiformis Stork 1934, non M. Sars Original description: Anterior aper- ture obliquely truncated, base or [?and] posterior aperture compressed, slightly deformed at each side. Type locality: Aegean Sea, 234-450 m. Present material: 1 station; Gulf of Cádiz, 535-546 m (76A: 18€). Depth range: 535-546 m. Reported distribution: Mediterra- nean; North Atlantic from Canary Ís., Biscay to Ireland and Norway, West Atlantic from Martha's Vineyard to Bar- bados, South Atlantic at St. Helena; 90- 2200 m. Earliest fossil: Pliocene DISCUSSION Of the 19 valid species listed for the Iberian coasts by HIDALGO (1917), 12 are represented in the present material. Ta- king into account the recent finding of the Eastern Mediterranean species Anta- lis rossati near Barcelona (Alzuria, 1986) and the deep-water Biscayan material of the “Talisman”, “Travailleur” (LOCARD, 1898) and “Valorous” (JEFFREYS, 1877) expeditions, the species list of Iberian scaphopods grows to 31. Species treated in these reports but not present in this material are listed in Table I. The present findings provide no bio- geographic novelties. Species living in both the Atlantic and Mediterranen Sea are Antalis agilis, A. dentalis, A. vulgaris, Episiphon filum, Entalina tetragona, Pulse- llum lofotense, Dischides politus and Cadu- 108 Remarks: Shell small, smooth and shiny, moderately curved, with a conspi- cuous swelling just anterior to the middle; ventral side regularly curved, dorsal side with distinct convex area due to the swe- lling; anterior shell aperture obliquely trun- cated facing downwards, slightly laterally compressed; posterior aperture without lobes, dorsoventrally depressed. This species may be confused with Cadulus subfusiformis (M. Sars 1865). It differs in being larger, conspicuously swollen near the middle of the shell and having a distinct convex area in the dorsal line. The anterior aperture is slightly laterally compressed, the poste- rior aperture dorsoventrally depressed, while C. subfusiformis has a faintly dorso- ventrally depressed anterior end and a round posterior one. lus jeffreysi. The only typical Mediterra- nean forms in the material is A. inaequi- costata. This latter shallow-water form extends, however, beyond the Strait of Gibraltar into the Gulf of Cádiz. On the other hand, A. entalis, A. novemcostata and Fissidentalium capillosum are known from Atlantic waters only. Comparing the bathymetric data of this material with reported ranges (Fig. 3), only Antalis agilis, A. inaequicostata and Entalina tetragona cover the greater part of their ranges. Most of the other species are found a one or two stations only, which may explain their relatively restricted bathymetric occurrence. CAPROTTI (1968) considers Cadulus oli- vii, C. strangulatus and C. tumidosus doubt- ful species or mere variations of C. ovulus. STEINER: Scaphopoda from the Spanish coasts Table I. Scaphopoda reported from the Iberian coasts but not in present material. Asterisks mark doubtful species (see Discussion). Tabla 1. Escafópodos citados en las costas ibéricas pero no hallados en el presente material. Los asteríscos indican especies dudosas (ver Discusión) Species Area Order DENTALIIDA Family DENTALIIDAE Antalis panorma (Chenu, 1842-47) Antalis rossati (Caprotti, 1966) Fissidentalium candidum (Jeffreys, 1877) Biscay FUSTIARIIDAE Fustiaria rubescens (Deshayes, 1825) Mediterranean 1917, Alzuria, 1987 Order GADILIDA Suborder ENTALIMORPHA Family ENTALINIDAE Subfamily BATHOXIPHINAE Bathoxiphus ensiculus (Jeffreys, 1877) Subfamily HETEROSCHISMOINAE Heteroschismoides subterfissum Suborder GADILIMORPHA Family GADILIDAE Subfamily SIPHONODENTALIINAE Siphonodentalium lobatum (Sowerby, 1860) Atlantic, Portugal Subfamily GADILINAE Cadulus artatus Jeftreys, 1880 Cadulus subfusiformis (M. Sors, 1865) Atlantic, Biscoy Cadulus gracilis Jeffreys, 1877 Biscoy Cadulus propinquus 6. O. Sars, 1878 Biscay Cadulus cylindratus Jeffreys, 1877 Biscay Cadulus monterosatoi Locard, 1897 Codulus gibbus Jeffreys, 1882 Cadulus ovulus (Philippi, 1844) Cadulus amphorus Jeffreys, 1882 *Codulus olivii (Scacchi, 1835) *Cadulus tumidosus Jeffreys, 1877 *Cadulus strangulatus Locard, 1897 Biscoy Biscay The occurrence of C. subfusiformis, a wide- spread Atlantic species, in the Mediterra- nean remains controversary. Although MONTEROSATO (1880) identifies a varia- tion (var. abyssicola) from Palermo, CA- PROTTI (1968) does not list this species as constant inhabitant of the Mediterranean. Recently, GAGLINI (1985) reconfirmed C. subfusiformis, and also MIFSUD's (1996) pho- Mediterranean, Northeast Spain, Balearic ls. Mediterranean, Northeast Spain Atlantic, Portugal, Biscay Atlantic, Portugal, Atlantic, Biscay, Mediterranean Atlantic, Galicia, Portugal Biscoy, Mediterranean -Aílantic, South Portugal Atlantic, Biscay, Galicia, Southern Portugal Biscay, Mediterranean off Morseilles Reference Locard, 1898; Hidalgo, 1917 Alzuria, 1986 Jeffreys, 1877 Locard, 1898; Hidalgo, Jeffreys, 1877 Locard, 1898 Jeffreys, 1882 Locard, 1898 Locard, 1898; Monterosato, 1875 Locard, 1898 Locard, 1898 Locord, 1898 Locard, 1898 Locard, 1898 Locard, 1898 Locard, 1898 Jeffreys, 1882 Locord, 1898 Locard, 1898 tograph of C. jeffreysi from Malta looks more like subfusiformis. However, until further reports come in, it remains possi- ble that in the Mediterranean C. subfusi- formis occurs in episodic pseudopopula- tions only (BOUCHET AND TAVIANI, 1992). Long lists of synonyms as for some Mediterranean scaphopod species may have several reasons. One of them, espe- 109 Iberus, 15 (1), 1997 cially when generic designations vary, is progress in supraspecific systematics. Another reason may be closely related but morphologically variable species and / or similar but not identical fossil forms. The species complex of Antalis inaequicostata, novemcostata and the Miocene mutabile (Do- derlein in HÓRNES, 1856) is a good exam- ple for a combination of these causes. An- talis mutabile and A. inaequicostata are very similar and both highly variable in their shell features. BUCQUOY ET AL. (1886) re- cognize the close relationship of mutabile and inaequicostata (=alternans). PILSBRY AND SHARP (1897-98) go further and list muta- bile as synonym of both dentalis (p. 53) and novemcostata (p. 211). RUGGIERI (1948) sug- gests mutabile being ancestral to novem- costata. CAPROTTI (1979) takes the alterna- tive view and presents the recent inaequi- costata as subspecies of the fossil mutabile. On the other hand, he assigns species sta- tus to novemcostata, although considering it the Atlantic descendant of mutabile. Fi- nally, PavIA's (1991) questionable assign- ment of mutabile to the genus Fissidenta- lium leaves little doubt about its species status. In this case, there are two argu- ments for treating the presumed ancestor and the descendants as separate species. First, according to CAPROTTI (1979: 232), there is a “typical mutabile” to be distin- guished from inaequicostata, the latter being extremely variable. Second, there is no fos- sil record of either species from the Ta- bianian (Lower Pliocene). Thus, there is a gap between the latest mutabile fossils from BIBLIOGRAPHY ALZURIA, M., 1986. Antalis rossati (Caprotti, 1966), nuevo escafópodo para la fauna es- pañola. Publicaciones del Departamento de Zo- ología (Barcelona), 12: 37-39. ALZURIA, M., 1987. El género Fustiaria (Mo- llusca; Scaphopoda) en el delta de Ebro. Pu- blicaciones del Departamento de Zoologia (Bar- celona), 13: 63-69. BOUCHET, P. AND TAVIANI, M., 1992. The Me- diterranean deep-sea fauna - pseudopopu- lations of Atlantic species. Deep-Sea Research, 39: 169-184. BroccHI, G., 1814. Conchiologia fossile subapen- nina. Vol. IL. Milano. 110 the Miocene and the earliest inaequicostata remains from the Middle Pliocene. This suggests that mutabile became extinct du- ring the desiccation of the Mediterranean in the Upper Miocene (RÓGL AND STEI- NINGER, 1983, 1984), and inaequicostata ra- diated back into the Mediterranean Basin from the Atlantic, perhaps splitting from novemcostata, the possible sister species from the European southwest coasts. A study of the relationships of the recent Eu- ropean shallow water Antalis species with molecular techniques is needed to bring more clarity on this question. That the re-colonisation of the Medi- terranean by scaphopods around the Miocene-Pliocene transition is not necessarily connected with speciation is demonstrated by the cases of A. dentalis and Entalina tetragona. Both species are represented in the fossil record almost unchanged (CaAPRrOTTI, 1965; 1968; 1979) from the Miocene on. ACKNOWLEDGEMENTS I am indepted to José Templado, Museo Nacional de Ciencias Naturales, for making the material available and for the information on station data. I also thank Luitfried Salvini-Plawen for his comments on the manuscript. An anony- mous reviewer provided the abstract in Spanish. This paper is a contribution to the Fauna Ibérica Programmes (DGCYT PB87-0397, PB89-0081, PB92-0121). BucQuoY, E., DAUTZENBERG, P. AND DOLLFUS, G., 1882. Mollusques marins du Roussillon. Bailliere and Fils, Paris, Tome l, Gasteropodes: 570 pp. CAPROTTI, E., 1961. Scaphopodi piaceziani di Castell Arquato (Piacenza). Atti della Societá Italiana di Science Naturali e del Museo Cívico di Storia Naturale di Milano, 100 (4): 345-359. CAPROTTI, E., 1965. Note on the Mediterranean Dentaliidae. Atti della Societá Italiana di Science Naturali e del Museo Civico di Storia Naturale di Milano, 104 (3): 339-354. CAPROTTI, E., 1968. Nota sui generi Cadulus ed Entalina nel Mediterraneo. Archiv fúr Mo- lluskenkunde, 98: 77-84. STEINER: Scaphopoda from the Spanish coasts CAPROTTI, E., 1979. Scaphopodi neogenici e re- centi del bacino mediterraneo. Iconograp- hica ed epitome. Bollettino Malacologico, 15: 213-288. CHISTIKOV, S. D. AND SAGAIDACHNYI, A. Y., 1981. Comparative morphology of shells in two species of the family Entalina. Zoologi- cheskii Zhurnal, 60: 36-41. (in Russian). FORBES, E., 1844. Report on the mollusca and ra- diata of the Aegean Sea and on their distri- bution, considering as bearing on geology. Re- ports of the British Association for the Advance- ment of Science 13 (for 1843): 130-193. FRIELE, H. AND GRIEG, J. A., 1901. The Norwe- gian North Atlantic Expedition 1876-1878. Zoology. Mollusca III. Scaphopoda. Chris- tiania, 49-51. GAGLINI, A., 1985. Riconferma di Cadulus sub- fusiformis (M. Sars). Notiziario CISMA., 6: 1-8. HIDALGO. J. G., 1917. Fauna malacológica de España, Portugal y las Baleares. Moluscos testáceos marinos. Trabajos del Museo Nácio- nal de Ciencias Naturales, Serie Zoologia, no. 30: 752 pp- HÓRNES, 1856. Die fossile Mollusken des Tertiar- Beckens von Wien. Jahrbúcher der Geologis- chen Reichsanstalt, Wien, 654 pp. JEFFREYS, J. G., 1870. Mediterranean Mollusca. Annals and Magazines of Natural History, (4) 6: 56-86. JEFFREYS, J. G., 1877. New and peculiar Mo- llusca of the order Solenoconchia procured by the “Valorous” expedition. Annals and Magazines of Natural History, (4) 19: 153-158. JEFFREYS, J. G., 1882. On the Mollusca procured during the Lightning and Porcupine expe- ditions 1868-70. Proceedings of the Royal Society of London, 1882: 656-687. LOCARD, A., 1898. Mollusques Testaces. Expe- ditions Scientifigues Travailleur et Talisman 1880-1883, Masson, ed., 2: 1031 pp. MArs, P., 1965. Faune marine des Pyrénées- Orientales. 5. Mollusques Aplacophores, Polyplacophores, Scaphopodes et Bivalves. Vie et Milieu, Supplément a Tome XV, fasc. 4: 156 pp. MIFSsUD, C., 1996. Living Mollusca from circa- littoral coastal muds, off Western Malta. La Conchiglia, 1996: 23-41. MONTEROSATO, T. A. DI, 1875. Nuova Revista delle Conchiglie Mediterranee. Atti dell“Ac- cademia Palerm. Sciencias, Lettri e Arti, Vol. V, Ser. 2: 1-50. MONTEROSATO, T. A. DI, 1880. Conchiglie della zona edgli abissi. Bullettino della Societa Ma- lacologica Italiana, VI: 50-82. Pavia, G., 1991. I molluschi del Messiniano di Borrelli (Torino). 2. Scaphopoda. Museo Re- gionale di Scienze Naturali Bollettino (Turin), 9: 105-172. PiLsBRY, H. A. AND SHARP, B., 1897. Scaphopoda. In Tyron, G. W. (Ed.): Manual of Conchylology, 17: 1-280. RÓGL, F. AND STEININGER, F. F., 1983. Vom Zer- fall der Tethys zu Mediterran und Paratethys. Die neogene Paláogeographie und Pa- linspastik des zirkum-mediterranen Raumes. Annalen des Naturhistorischen Museums Wien, 85/A: 135-163. RÓGL, F. AND STEININGER, F. F., 1984. Neogene Paratethys, Mediterranean and Indo-Pacific Seaways. Implications for the paleobiogeo- graphy of marine and terrestrial biotas. In Brenchley, P. (Ed.): Fossils and Climate, Wiley and Sons, Ltd., 171-200. RUGGIERI, G., 1948. La malacofauna del Cala- briano Romagnolo. Giornale di Geologia. An- nali del Museo Geologico di Bologna, 20: 63-110. SARS, G. O., 1872. Some remarkable forms of ani- mal life from the great deeps off the Nor- wegian coast. I. Partly from posthumous ma- nuscripts of the late Professor Dr. Michael Sars. Christiania University Program, 1% half- year 1869: 1-VIII, 1-85, 6 pl. SARS, G. O., 1878. Bidrag til Kundskaben om Norges arktiske Fauna. I. Mollusca Regio- nis Arcticae Norvegiae. Oversigt over de i Norges arctiske region forkommende bloddyr. Universitetsprogram Christiana, forste halvaar 1878: 466 pp., 18pl. SARS, M., 1865. Malakozoologiske Jagtagelser. II. Nye Arter af Slaegten Siphonodentalium. Forhandlinger Videnskabs-Selskabet i Christia- nia, 1864: 296-315. SCARABINO, V., 1995. Scaphopoda of the tropi- cal Pacific and Indian Oceans, with descrip- tions of 3 new genera and 42 new species. Mé- moirs du Museum national d'Histoire naturelle. Résultats des Campagnes Musorstom, vol. 14, P. Bouchet (ed.), 167: 189-379. STEINER, G., 1992. Phylogeny and classification of Scaphopoda. Journal of Molluscan Studies, 58: 385-400. STEINER, G., 1995. Larval and juvenile shells of four North Atlantic scaphopod species. Ame- ican Malacological Bulletin, 11: 87-98. STEINER, G., 1996. Suprageneric phylogeny in Scaphopoda. In Taylor, J. (Ed.): Origin and Evolutionary Radiation of the Mollusca, Oxford University Press, 329-335 pp. STORK, H. A., 1934. Scaphopoden der Adria. Thalassia, 1(9): 3-18. TEMPLADO, J., GUERRA, A., BEDOYA, J., MORENO, D., REMÓN, J. M., MALDONADO, M. AND RA- MOS, M. A., 1993. Fauna marina circalitoral del sur de la península Ibérica. Resuldados de la Campana Oceanografica “Fauna 1”. Museo Na- cional de Ciencias Naturales Consejo Supe- rior de Investigaciones Científicas, 183 pp. TÉrry, A., 1959. Classe des Sipunculiens. In: Gras- sé, P. P. (Ed.): Traitéde Zoologie. Anatomie, Sys- tematique, Biologie. Tome V /1: 785-854. = 111 ad mA d A ds m NE: A” 7 rr EM ct eno A ooo ME EN 7d did ' (3 lo ps ab mt ' MS, ep das Hd, erat RA ¿gee Y MAA Mes a AA E LEE Qe e de 3 PA Y amo St Las ias un mo A Y JN O Sociedad Española de Malacología —_—_—_—_——— Iberus, 15 (1): 113-123, 1997 Análisis parasitológico de gasterópodos acuáticos del delta del Llobregat (Barcelona). Estadios larvarios de trematodos digénidos Parasitological study on aquatic gastropods from the Llobregat delta (Barcelona). Larval stages of digenetic trematodes Mercedes VILLA, Isabel MONTOLIU, Mercedes GRACENEA y Olga GON- ZÁLEZ-MORENO* Recibido el 8-VIII-1996. Aceptado el 20-XT-1996 RESUMEN Se ha estudiado, durante los años 1990 a 1995, el índice de parasitación por estadios larvarios de trematodos digénidos de 6. 184 gasterópodos acuáticos recolectados en el delta del Llobregat, pertenecientes a 4 especies, 2 prosobranquios (Hydrobiidae) y 2 pul- monados (Ellobiidae, Physidae). El hidróbido Mercuria confusa [Frauenfeld, 1863) ha resultado ser el único parasitado por digénidos, mostrando una prevalencia de infestación del 3,22% (168 positivos sobre 5.219 analizados). Se han detectado hasta 5 especies de trematodos en estadio larvario, las cuales siguen dos modalidades de ciclo biológico: a) emisión de cercarias (xifidiocercarias Lecithodendriidae Odhner, 1910 y Microphalli- dae Travassos, 1920, cercarias inermes Notocotylidae Lúhe, 1957; b) ausencia de emer- gencia cercariana (Microphallidae y Heterophyidae -Leiper, 1909- Odhner, 1914). La alta densidad poblacional de M. confusa y su susceptibilidad de ser parasitado por lar- vas de diversas especies de digénidos revelan su importancia en el mantenimiento de los ciclos biológicos de trematodos, ratificando el significativo papel que ejercen los proso- branquios como hospedadores intermediarios específicos en ambientes palustres. ABSTRACT Aquatic gastropods from the Llobregat delta were studied in order to detect infection pre- valence by larval digenetic trematodes in the period 1990-1995. Specimens analysed (6.194) belonged to 4 species, 2 prosobranchs (Hydrobiidae) and 2 pulmonates (Ellobii- dae, Physidae), the hydrobiid M. confusa (Frauenfeld, 1863) being the only infected. Lar- vae of 5 trematode species were detected following two life cycle modalities: a) cercarial emergence (xifidiocercariae Lecithodendriidae Odhner, 1910 and Microphallidae Travas- sos, 1920, unarmed cercariae Notocotylidae Lúhe, 1957); b)without cercarial emergence (Microphallidae and Heterophyidae -Leiper, 1909- Odhner, 1914). The high density of M. confusa population and its susceptibility to infection by several digenetic species show it to be a specific intermediate host in the life cycle of these parasites, thus confirming the signi- ficant role of prosobranchs in the maintenance of trematode life cycles in deltaic zones. PALABRAS CLAVE: delta del Llobregat, Prosobranchia, Hydrobiidae, M. confusa, estadios larvarios de digéni- dos, Lecithodendriidae, Microphallidae, Notocotylidae, Heterophyidae. KEY WORDS: Llobregat delta, Prosobranchia, Hydrobiidae, M. confusa, larval stages of Digenea, Lecithodendriidae, Microphallidae, Notocotylidae, Heterophyidae. * Laboratorio de Parasitología, Facultad de Farmacia, Universidad de Barcelona, Avda. Diagonal, s/n, 08028 Barcelona. 113 Iberus, 15 (1), 1997 INTRODUCCIÓN Las marismas del delta del Llobregat acogen una amplia variedad de gasteró- podos acuáticos y terrestres, distribui- dos en las lagunas, canales y terrenos colindantes, especies que se extienden por toda la planicie litoral en abundan- tes poblaciones y cuya distribución y frecuencia ha quedado reflejada en el estudio de ALTIMIRA (1969), autor que describe hasta 79 especies presentes en diversos hábitats deltaicos, destacando asimismo los tratados malacológicos generales elaborados por Haas (1929), VIDAL ABARCA Y SUÁREZ (1985) y BECH (1990). La importancia parasitológica de este grupo zoológico de invertebrados reside en el destacado papel que osten- tan sus especies como primeros hospe- dadores intermediarios específicos, albergantes de esporocistos, redias y cercarias, de prácticamente todas las especies parásitas de trematodos digéni- dos de ciclos biológicos conocidos; pudiendo representar un doble papel en los ciclos abreviados, al intervenir como primeros y segundos hospedadores intermediarios simultáneamente, alber- gantes además de metacercarias. El proceso natural de eutrofización existente en este tipo de ambiente lagunar, caracterizado por la presencia constante de agua salobre, rica en materia orgánica, y la descomposición de la vegetación helofítica, constituye el factor primordial para el asentamiento de moluscos eurihalinos. Otros peque- ños invertebrados que conviven estre- chamente con los anteriores (crustáceos e insectos) pueden actuar como segun- dos hospedadores intermediarios de estos digénidos, sirviendo, asimismo, como fuente de alimentación de la fauna vertebrada (aves, micromamíferos, peces), hospedadores de las formas adultas del parásito. El conocimiento de la biología de los digénidos en este tipo de ambiente palustre requiere un seguimiento previo de las especies hospedadoras parasita- das, de los biotopos que frecuentan y de su comportamiento. En este sentido, 114 cabe destacar los numerosos trabajos faunístico-ecológicos realizados en el delta del Ebro, basados en las especies vermidianas y sus hospedadores micro- mamíferos (FELIU, TORRES, GÁLLEGO, GOSÁLBEZ Y VENTURA, 1985; GRACENEA, FELIU, MONTOLIU, TORRES Y GÁLLEGO, 1987; FELIU, TORRES, GRACENEA Y MON- TOLIU, 1990), trabajos que han servido como punto de referencia para el estudio de los ciclos biológicos, princi- palmente acuáticos, de digénidos que tienen lugar en dicho enclave (MONTO- LIU, GRACENEA, VILLA Y GONZÁLEZ- MORENO, 1991). De la misma forma, los llevados a cabo en pequeños mamíferos del delta del Llobregat, menos numero- sos y caracterizados por recoger una fauna parasitaria cualitativamente y cuantitativamente más pobre (GRACE- NEA Y MONTOLIU, 1992; GRACENEA, MONTOLIU Y DEBLOCK, 1993), han sido ampliados con el estudio parasitológico de diversas especies de moluscos, hos- pedadoras de larvas de digénidos tanto de ciclo acuático (MONTOLIU, GRACENEA Y DEBLOCK, 1992), como de ciclo terres- tre (GONZÁLEZ-MORENO, GRACENEA, MONTOLIU Y VILLA, 1994). Continuando en esta línea de investigación sobre ciclos biológicos de trematodos parási- tos, este trabajo muestra la diversidad existente en la helmintofauna larvaria asociada a gasterópodos acuáticos adap- tados a hábitats deltaicos. MATERIAL Y MÉTODOS Recolección de caracoles y manteni- miento en el laboratorio: El enclave prospectado, ya señalado anteriormente por GONZÁLEZ-MORENO ET AL. (1994), comprende los márgenes de la laguna de La Ricarda (Biotopo A) y terrenos colindantes del margen derecho parcial- mente inundados por las aguas (Biotopo B) (ver Figura 1). Ambos puntos de muestreo constituyen el hábitat de asen- tamiento de diversas comunidades de moluscos, especies que son capaces de soportar la intensa fluctuación estacio- VILLA ET A4L.: Larvas de tremátodos digénidos en gasterópodos acuáticos del Llobregat BARCELONA NM b 1,5 km RÍO LLOBREGAT PRAT DEL LLOBREGAT 2 es AEROPUERTO Figura 1. Situación geográfica de la zona prospectada en el delta del Llobregat. A: laguna de La Ricarda. B: localización de los biotopos estudiados. Figure 1. Geographic situation of the prospected areas in the Llobregat delta. A: La Ricarda lagoon. B: localization of the study areas. nal del enclave, con aporte constante de agua dulce procedente de los canales y del acuífero superficial e infiltración de agua marina. Biotopo A: la vegetación natural pre- dominante en esta comunidad helofítica son los cañizares que bordean el margen de la laguna, con una especie predomi- nante, el cañizo (Phragmites australis), cuyos tallos parcialmente sumergidos permiten el ascenso de los gasterópodos hacia la superficie. Agua de salinidad variable (3-8%0)y pH entre 7 y 8. Biotopo B: constituido por zonas ane- gadas, poco profundas, cercanas al margen derecho de la laguna, alimenta- das esencialmente por los canales de desagúe y las lluvias; con predominio de la vegetación helofítica asociada a especies halófilas. Junto a las gramíneas (Phragmites) se encuentra la espadaña (Typha angustifolia, T. latifolia), el esparto (Spartina juncea), así como una comuni- dad bentónica (Enteromorpha, Ulva, Chara); dado el menor aporte de agua marina la salinidad es menor (2-5%o) y pH entre 7 y 8. Las especies de caracoles recolecta- das en ambas zonas de muestreo han sido seis: 2 prosobranquios, Mercuria confusa (Frauenfeld, 1863) y Potamopyr- gus jenkinsi (Smith, 1889) (Hydrobiidae) y 2 pulmonados, Ovatella (Myosotella) myosotis (Draparnaud, 1801) (Ellobiidae) y Physa acuta (Draparnaud, 1805) (Physi- dae). En la Tabla I queda reflejado el número de caracoles de cada especie estudiado, el biotopo de prospección y la época de recolección. El estudio ha abarcado un periodo de cinco años (1990-95) con muestreos en primavera y otoño. Para ello, fueron utilizados tamices metálicos con los que se procedía a barrer el fondo fangoso o bien se practicaban pequeñas sacudidas de las partes sumergidas de la vegeta- ción acuática. Una vez en el laboratorio los moluscos eran estabulados, reprodu- ciendo las características específicas de cada zona de captura: con una salinidad 115 Iberus, 15 (1), 1997 Tabla I. Especies de gasterópodos estudiadas: distribución del número de ejemplares según época de recolección y biotopo prospectado (A, B). Table I. Gastropod species analysed: specimen distribution in the prospected biotopes (A, B) and annual variation. 1990 1991 1992 A A A Prosobranchia Mercuria confusa IIS O 27] Potamopyrgus ¡jenkinsi 12 21 9 Pulmonata Ovatella ([Myosotella) myosotis 25 6 17 Physa acuta 4 40 y pH adecuados, dieta constituida por lechuga seca y alimento para peces. Detección y aislamiento de formas larvarias de digénidos: La detección de una posible emisión de cercarias al medio externo se efectuó disponiendo individualmente los gasterópodos en pocillos de placa de cultivo celular con- teniendo agua del biotopo y observán- dolos posteriormente bajo la lupa en búsqueda de cercarias nadando libre- mente. Para la detección de esporocis- tos, redias y metacercarias, se procedió a la disección de los caracoles y a la obser- vación de todos sus órganos. Técnicas microscópicas: Las larvas fueron inicialmente estudiadas in vivo con el microscopio óptico y con la ayuda del colorante vital rojo neutro, proce- diéndose posteriormente a la fijación con el líquido de Bouin, tinción con carmín alumínico y tras cuidadosa des- hidratación se montaron con Bálsamo del Canadá. RESULTADOS El estudio de 5.219 ejemplares de M. confusa ha puesto de manifiesto la capa- cidad de este prosobranquio, el único gasterópodo que ha resultado estar parasitado, para albergar diferentes formas larvarias de digénidos, en total cinco especies caracterizadas por su 116 1993: 1994 1995 Total A B ASNO B E O MO CA O NO 224 40 7 36 : : E Aa 121 OA 1 589 ciclo biológico acuático. Todas las infes- taciones por digénidos han mostrado invasión a nivel del complejo glándula digestiva-gónada, como hábitat de elec- ción para la evolución de las larvas. Las especies de digénidos detecta- das, pertenecientes a 4 familias, presen- tan dos modalidades de ciclos biológi- cos (Fig. 2). En la primera de ellas, es característica la emisión de cercarias al medio externo, bien sea en ciclos trihete- roxenos (con tres hospedadores) o dihe- teroxenos (con dos hospedadores). En la segunda modalidad, ciclo de tipo abre- viado, el gasterópodo se comporta como primer y segundo hospedador interme- diario simultáneamente, no habiendo emisión de cercarias. I. Emisión de cercarias I. a. Ciclos triheteroxenos Cercarias Lecithodendriidae. Lecit- hodendriidae gen. sp.: Cercarias xifi- diocercas (con estilete), distomas, pro- vistas de una cola recta más estrecha que el cuerpo (leptocercas) y virguladas (cuerpo: 150 x 90 um). Originadas en esporocistos sacciformes (100-350 x 50- 100 um) provistos de poro de salida musculoso a través del cual las cercarias emergen según un patrón de emisión predominantemente nocturno. La morfoanatomía de estas cercarias se ajusta a la descrita para la familia Lecithodendriidae (YAMAGUTI, 1975; SCHELL, 1985), destacando su afinidad con xifidiocercarias Lecithodendriidae VILLA ET 4Lz.: Larvas de tremátodos digénidos en gasterópodos acuáticos del Llobregat o 'O) enquistamiento O) anfibios, Cercarias me a e reptiles o EMISIÓN Lecithodendriidae : 22H. 1 quirópteros insectos ( 19) H'D DE CERCARIAS (H. D.) E SLCALIAS enquistamiento O) insectivoros M. feliui ——Y» en crustáceos, 5 (C. russula) E) anfípodos e (H. D.) isópodos (2? H. L) CICLOS TRIHETEROXENOS CICLOS DIHETEROXENOS | — Catia enquistamiento O) Aves a : ——PY» en plantas = aos E Notocotylidae p y > acuáticas conchas del caracol (H. D.) iS enquistamiento 2) o M. fusiformis bip> q = =D> acuáticas : : en esporocistos HD NO EMISIÓN (Microphallidae) (H. D.) peces o reptiles (H. D.) DE CERCARIAS cercarias $ enquistamiento Heterophyidae en el caracol Figura 2. Ciclos biológicos de los digénidos detectados en Mercuria confusa. (—): fases conocidas. (=>): fases presumibles. 2” H. L.: segundo hospedador intermediario. H. D.: hospedador definitivo. 1: penetración activa. 2: penetración pasiva. 3: huevos en heces del H. D. Figure 2. Life cycles of digeneans detected in Mercuria confusa. (—): known phases. (==): likely phases. 2'H. L.: second intermediate host. H. D.: definitive host. 1: active penetration. 2: pasive penetration. 3: eggs in definitive host feces. detectadas en otros biotopos deltaicos emergencia cercariana se produce predo- geográficamente próximos, el delta del minantemente en horas crepusculares. Ebro (MONTOLIU ET AL., 1991) y costa La realización experimental del ciclo francesa mediterránea (DEBLOCK, 1980). biológico, descrito sucintamente por los Las cercarias probablemente infestan autores de la especie, ha permitido su larvas de insectos (segundos hospeda- determinación sistemática. Las cercarias dores intermediarios, en los que se infestan activamente a crustáceos anfí- forma la metacercaria), desarrollándose podos e isópodos (segundos intermedia- el adulto probablemente en anfibios y rios), evolucionando a metacercarias reptiles (LLUCH, ROCA Y NAVARRO, 1986) enquistadas, los cuales son depredados o en quirópteros (ESTEBAN, OLTRA- por el hospedador definitivo, el insectí- FERRERO, BOTELLA Y GRANEL, 1993) por voro Crocidura russula (Hermann, 1780) depredación, aunque sin descartar su en el delta del Llobregat, en el que se posible presencia en micromamíferos originan los adultos a nivel intestinal (GRACENEA ET AL., 1987). (GRACENEA Y MONTOLIU, 1992; GRACE- NEA ET AL., 1993). Cercarias Microphallidae. Mari- trema feliui Gracenea, Montoliu et I. b. Ciclos diheteroxenos Deblock, 1993: Xifidiocercarias, monos- Cercarias Notocotylidae. Notocoty- tomas, anentéricas y leptocercas (cuerpo: lidae gen. sp.: Cercarias de gran tamaño 120 x 68 um), de fórmula excretora (cuerpo: 600-800 ym x 370 um) y gran 2((2+2)+(2+2)= 16 solenocitos. Se origi- opacidad, inermes (ausencia de estilete), nan en esporocistos de aspecto sacci- oftalmocercas (con manchas oculares), forme e irregular, (166-430 x 140-180 monostomas y leptocercas. Se originan um), provistos de poro de salida. La en redias de aspecto fusiforme, provis- 15 Iberus, 15 (1), 1997 tas de faringe subterminal y ciego intes- tinal de gran volumen. Las cercarias acaban de madurar fuera de las redias, emergiendo al medio externo y enquis- tándose rápidamente en la vegetación o sobre la misma concha del caracol, evo- lucionando a metacercarias (cuerpo: 736 x 138 um) confinadas en quistes hemies- féricos (diámetro, 150-300 um). La morfología de las cercarias y el ciclo diheteroxeno con enquistamiento en el medio acuático son propios de la familia Notocotylidae (YAMAGUTI, 1975; SCHELL, 1985). Los adultos se desarro- llan presumiblemente en aves acuáticas deltaicas (hospedadores definitivos) al alimentarse éstos de plantas acuáticas o de moluscos. II. No emisión de cercarias (ciclos abre- viados) Cercarias Microphallidae. Microp- hallus fusiformis Reimer, 1963: Cerca- rias rudimentarias (blastocercarias) de reducidas dimensiones (50-70 x 30-37 ym), inmóviles y constituidas por células indiferenciadas, sin apéndice caudal ni esbozos de otras estructuras. Se forman en esporocistos blanquecinos, transparentes (200-500 x 150 um) y sac- ciformes, en los que evolucionan a metacercarias enquistadas (diámetro, 80-110 x 59-64 um). La metacercaria desenquistada se caracteriza por su pequeño tamaño (140- 160 x 60 um) y cuerpo fusiforme muy espinulado; de carácter progenético, con testículos y glándulas vitelógenas fun- cionales; ovario diestro y metratermo confluyendo a nivel de la pared lateral del atrio genital La morfología de dichas metacercarias, muy similar a la del adulto (MONTOLIU ET AL., 1992). El hos- pedador definitivo en el delta lo consti- tuyen probablemente aves anseriformes (REIMER, 1963). Cercarias Heterophyidae. Hete- rophyidae gen. sp.: Cercarias monosto- mas, leptocercas y oceladas, con órgano de penetración protráctil (cuerpo: 100- 120 x 50-60 ym) y sistema excretor pro- visto de glándula post-vesical. Evolucio- nan a partir de redias de aspecto cilín- drico (158-370 x 52-103 ym), provistas de faringe musculosa y un ciego corto. Las cercarias, tras emerger de la redia, se enquistan dentro del mismo caracol. Las metacercarias están provistas de una doble corona de espinas rodeando a la boca. Son distomas, con testículos homolaterales, vesícula excretora de gran tamaño, encontrándose confinadas en quistes (80x70 um) de doble cubierta. Las aves acuáticas, peces o anfibios podrían actuar como hospedadores defi- nitivos del digénido (YAMAGUTI, 1975). En la Tabla II se encuentran recopila- das las prevalencias de infestación en Mercuria confusa para cada una de las especies descritas anteriormente. DISCUSIÓN Los estudios sobre la helmintofauna larvaria de digénidos de ciclo acuático señalan a los moluscos prosobranquios como los principales hospedadores intermediarios específicos para las espe- cies Digenea de ambientes palustres. Los datos obtenidos en nuestro estudio han mostrado que el hidróbido Mercuria confusa interviene como hospedador intermediario específico de cinco espe- cies de digénidos deltaicos. Este proso- branquio ha sido estudiado parasitoló- (Página derecha). Figura 3. Estadios larvarios de digénidos detectados en Mercuria confusa. A: cer- caria Lecithodendriidae (rojo neutro). B: blastocercaria y quistes metacercarianos intrasporocísticos de M. fusiformis. C: metacercaria enquistada Heterophyidae. D: cercaria de M. feliui (rojo neutro). Escalas 25 um. (Right page). Figure 3. Digenean larval stages detected ¿in Mercuria confusa. 4: cercaria Lecithodendriidae (neutral red). B: intrasporocystic blastocercaria and metacercarial cysts of M. fusifor- mis. C: encysted metacercaria Heterophyidae. D: cercaria of M. feliui (neutral red). Scale bars 25 pm. 118 VILLA ET AL.: Larvas de tremátodos digénidos en gasterópodos acuáticos del Llobregat 119 Iberus, 15 (1), 1997 Tabla II. Parasitación de M. confusa por estadios larvarios de digénidos según épocas de recolección y biotopo prospectado (A, B). N=n" de especímenes estudiados (por emisión y/o por disección). %=prevalencia de parasitación. Table II. Infection of M. confusa with larvae of digenetic trematodes, related to year of collection and prospected biotopes (A, B). N=number of examined snails (emergence andlor dissection). %=prevalence of infection. 1990 1991 1992 A A A % % % 1993 1994 1995 Total % % % % % % Cercarias+Esporocistos (o redias) (emisores de cercarias) N=1215 N=656 N=271 N=989 N=708 N=146 N=1164 N=70 N=5219 Lecithodendriidae gen sp. : 0,40 0,42 - 0,28 - 0,15 Maritrema felivi OASIS ERA SOTA ZONA: - ISAAC O Notocotylidae gen. sp. - OOO O 0,10 Cercarias+Esporocistos (o redias) + metacercarias (ciclos abreviados) N=492 N=482 N=33 N=475 N=210 N=32 N=150 N=0 N=1874 Microphallus fusiformis IESDAE2IO : a - : 2,51 Heterophyidae gen. sp. 325 0,85 gicamente con anterioridad en el delta del Ebro por MONTOLIU ET AL. (1991), detectándose formas larvarias pertene- cientes a 7 especies de digénidos. La ausencia de parasitación en el proso- branquio Potamopyrgus jenkinsi, en el que sólo han sido estudiadas hembras partenogenéticas, podría deberse a un fenómeno similar al observado por LiveLY (1989) para otras especies de Potamopyrgus Stimpson, 1865 en Nueva Zelanda, en las que se demuestra la correlación positiva entre las poblacio- nes sexuadas y la parasitación por microfálidos, no estando nunca parasi- tadas las poblaciones que se reproducen exclusivamente por partenogénesis. En lo que respecta a las especies de pulmo- nados estudiadas, si bien éstas no se encuentraron parasitadas por digénidos, sí que existen numerosas citas de infes- taciones por otras especies de digénidos en ejemplares dulceacuícolas europeos (MOUAHID Y MONÉ, 1988, entre otras). La prevalencia de parasitación por larvas de digénidos mas alta detectada en M. confusa ha correspondido a la familia Microphallidae. Los digénidos de biología conocida de esta familia incluyen a diversos moluscos proso- branquios como hospedadores interme- 120 diarios, tanto en las especies triheteroxe- nas como las diheteroxenas, de ciclo abreviado. En la primera modalidad, para las especies de Maritrema Nicoll, 1907 y Microphallus Ward, 1901, son los prosobranquios pertenecientes a Litto- rina Ferrusac 1822 (BENJAMIN Y JAMES, 1987; IRwIN, MAGUIRE Y SAVILLE, 1990; GALAKTIONOV Y BUSTNERS, 1995) y a Hydrobia Hartmann, 1821 (GARKAVI, 1972; PREVOT Y BARTOLI, 1977; DEBLOCK, 1978; SAVILLE Y IRwIN, 1991) los más fre- cuentemente citados, y más puntual- mente los prosobranquios Bythinella Moquin-Tandon, 1855 (JOURDANE, 1979), Pseudamnicola Paulucci, 1878 (KULKINA Y BELYAKOVA, 1983), Cerithium Bruguiere, 1789, Bittium Leach, 1847 (PREVOT, BARTOLI Y DEBLOCK, 1976; BARTOLI Y Prevor, 1978) y Cerithidea Swainson, 1840 (ABDUL-SALAM Y SREELATHA, 1991). El género Microphallus es el que engloba el mayor número de especies con ciclos abreviados, interviniendo habitual- mente especies de Hydrobia y Littorina y puntualmente de Bittium, incluyéndose únicamente a especies de Hydrobia para el género Maritrema (DEBLOCK, 1977; LAUCKNER, 1984). En lo que se refiere a la familia Lecit- hodendriidae, los prosobranquios vuel- VILLA ET AL.: Larvas de tremátodos digénidos en gasterópodos acuáticos del Llobregat ven a ser citados frecuentemente en los trabajos sobre trematodofauna larvaria, destacando los realizados en especies de Bithynia Leach, 1818 (YAMAGUTL 1975) y Amnicola Gould y Haldemanmn, 1841 (Ca- BLE, 1985). Con respecto a los hospeda- dores intermediarios de notocotílidos, cabe señalar a los prosobranquios Hydro- bia (STUNKARD, 1966; DEBLOCK, 1980), Po- tamopyrgus (BisseT, 1977), Bithynia (Y A- MAGUTI, 1975; VASILEV Y KANEvV, 1984), Littorina (GRANOVICH, MIKHALOVA Y SERGIEVSKIIL, 1987), así como a especies de melaníidos (KHALIFA Y EL-NAFFAR, 1979). En los digénidos heterófidos no son frecuentes los ciclos abreviados como el que tiene lugar en el delta del Llobregat, habiéndose citado a una sola especie de digénido, Metagonimoides ore- gonensis Price, 1931, la cual infesta a di- versas especies de prosobranquios pleu- rocéridos, únicos hospedadores interme- diarios del ciclo (YAMAGUTI, 1975). El análisis cuantitativo de la parasita- ción por larvas de M. confusa muestra diferencias en las prevalencias para cada una de las especies de digénidos halla- dos (Tabla II). Los índices totales más elevados mostrados por los microfálidos (Maritrema feliui - 1,76 %; Microphallus fusiformis - 2,51 %) parecen ajustarse a los detectados en hidróbidos que habitan zonas geográficas litorales próximas al delta del Llobregat. MONTOLIU ET AL. (1991) muestran para M. confusa en el delta del Ebro unos niveles de parasita- ción para Maritrema sp. (1,36 %) muy similares a las del presente trabajo. Asi- “mismo, en el estudio realizado por DEBLOCK (1978) se observan índices de parasitación por microfálidos en especies de Hydrobia que oscilan entre el 2,5 y 0,5% en el litoral atlántico y entre el 6 y 0,5% en el litoral mediterráneo. En otros prosobranquios litorales también muy estudiados del género Littorina, la preva- lencia de estadios larvarios Microphalli- dae es muy alta, del orden del 23,8% en la costa del Mar Báltico (LAUCKNER, 1984) y de hasta el 40-50% en las costas soviéticas (SERGIEVSKIL, 1985). La baja prevalencia por lecitodéndri- dos en el Llobregat no parece ajustarse a las tasas de infestación observadas en M. confusa del delta del Ebro, en el que se alcanzan índices elevados (5,83%) (datos no publicados). Los bajos índices de infestación detectados en el resto de digénidos sí parecen coincidir con los obtenidos para este hidróbido en el Ebro, detectándose el 0,27% para cerca- rias Heterophyidae y el 0,20% para cer- carias Notocotylidae, y con los de DEBLOCK (1978) en las costas francesas para especies de Hydrobia, 0,07-1,67% para heterófidos y 0,20-0,43% para noto- cotílidos. En cuanto a la dinámica del parasi- tismo, las fluctuaciones temporales observadas en el hidróbido no parecen guardar relación directa con su densi- dad poblacional, relativamente cons- tante en todas las prospecciones realiza- das, o con el biotopo de prospección. Ello podría estar relacionado con el comportamiento de los hospedadores definitivos como factor de máxima influencia en dichas variaciones. Mari- trema feliui es la única especie de digé- nido que parece mantenerse relativa- mente constante en el tiempo, hecho que puede explicarse por la presencia regular del insectívoro Crocidura russula en el delta (GRACENEA Y MONTOLIU, 1992). En cuanto a Notocotylidae gen. sp. y Microphallus fusiformis, potencial- mente parásitos de aves deltaicas, su aparición más esporádica en los caraco- les estaría condicionada por el carácter migratorio de sus hospedadores defini- tivos. AGRADECIMIENTOS Los autores agradecen a D. Serge Gofas, malacólogo del Laboratoire de Biologie des Invertébrés Marins et Mala- cologie du Muséum National d'Histoire Naturelle de Paris, la determinación de los moluscos prosobranquios. Asimismo, agradecen a D. 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Keigaku Publishing Co. Tokyo. 590p + 219 pl. 123 Car She EN Modo, Arias > AY A arial o 'auelalord als * ol : cl, to «bismncón (029 nd os vr pl y ene ta «ilu 20 IgE id Erin de vd dl bh b Aca! o sos á ; iaberirs ¡hháb yd Ls A Pas + "0 Pd q q e O Ra! d bes sd ra ¿Y doo hor Yi yiRtltsialo, ra hac AO p MAYA rada A ci A cn e 3 daria cia yde ri ira hb. ' b Se lec e e ¿mon pol eos ddEP8 ra 38 Ñ qutt 1 dé ad) rg bss O Sociedad Española de Malacología ——_—_——— Iberus, 15 (1): 125-130, 1997 On a floating egg mass of the diamond shaped squid 7)»y- sanoteuthis rhombus (Cephalopoda: Thysanoteuthidae) in the western Mediterranean Observaciones sobre una puesta pelágica del calamar losange 7»y- sanoteuthis rhombus (Cephalopoda, Thysanoteuthidae) hallada en el Mediterráneo occidental Angel GUERRA and Francisco ROCHA* Recibido el 14-X1-1996. Aceptado el 3-XII-1996 ABSTRACT This is the second record of a floating egg mass of the diamond shaped squid Thysanoteu- this rhombus in the Mediterranean (37* 11.85” N - 1? 31.15 E). The first one was obser- ved at the Strait of Messina in 1929. The egg mass was a dense, resilient oblong cylinder with rounded tips appoximately 100 cm in length and about 20 cm in diameter. From a small sample, egg capsules and paralarvae (1.85+0.08 mm MIL) are described. Some complementary characters about this species paralarvae, such as the “arm formulae, the presence of an incipient swimming keel-like shaped membrane on some arms, and the mantle chromatophore pattern should assist in their identification. RESUMEN En este trabajo se informa sobre el segundo hallazgo de una puesta pelágica del calamar losange Thysanoteuthis rhombus en el Mar Mediterráneo (37? 11,85" N - 1? 31,15 E). La primera se observó en el estrecho de Mesina en 1929. La masa de huevos consistía en un cilindro oblongo con los bordes romos, denso y elástico, de unos 100 cm de longitud y 20 cm de diámetro. A partir de una muestra pequeña que se pudo obtener se describen las cápsulas ovigeras y las paralarvas (1,85+0,08 mm ML). Se proporcionan algunos caracteres complementarios útiles para la identificación de estas paralarvas: la formula braquial, la presencia de una carena natatoria incipiente aquillada en algunos brazos y el patrón de cromatóforos en el manto. KEY WORDS: Cephalopoda, Thysanoteuthis rhombus, egg mass, paralarvae, Mediterranean Sea. PALABRAS CLAVE: Cephalopoda, Thysanoteuthis rhombus, puesta, paralarvas, mar Mediterráneo. INTRODUCTION The diamond shaped squid Thysano- partially subtropical waters of the World teuthis rhombus Troschel, 1857 is an epi- Ocean including the Mediterranean, pelagic inhabitant of warm tropical and often occurring in pairs or small schools * Instituto de Investigaciones Marinas, C/ Eduardo Cabello 6, 36208 Vigo, España. 25 Iberus, 15 (1), 1997 (NISHIMURA, 1966; CLARKE, 1966; NIG- MATULLIN, ARKHIPKIN AND SABIROV, 1995). It is a large oegopsid squid which reaches up to 85 cm in mantle length (ML) and 24 kg in body weight (NIGMA- TULLIN ET AL., 1995). T. rhombus is one of the fastest-growing squid: in approxi- mately one year, it reaches its maximum ML (NIGMATULLIN ET AL., 1995). This species has high potential fecundity (up to 4.8 million oocytes), but a rather small maximum volume of oviducts (up to 140,000 eggs) and egg masses (35,000 to 75,000), which suggest that T. rhombus is an intermittent spawner with multiple filling and evacuation of oviducts (NIG- MATULLIN ET AL., 1991; NIGMATULLIN ET AL., 1995). T. rhombus is one of the few oegopsid cephalopods in which the spawn is known. Until recently, a total of 21 egg masses had been observed. All were found drifting in the surface water layer of the tropical Atlantic, northwest and southeast Pacific and the Mediterranean (review in SABIROV, ARKHIPKIN, TSYGAN- KOV AND SHCHETINNIKOV, 1987). Egg masses of this species are gelatinous, sau- sage-shaped, 60-180 cm long by 10-30 cm diameter; containing a double spirally arranged row of eggs embedded in the surface layer of the mass (MISAKI AND OKUTANI, 1976; SUZUKI, MISAKI AND OKUTANI, 1979). The egg mass was pho- tographed in natural environment for the first time by SUZUKI ET AL. (1979). The first and unique reference to an egg mass of T. rhombus in the Mediterranean (Strait of Messina) was given by SANZO (1929). Although the occurrence of this species in the Mediterranean is rare (MORALES, 1981; BIaGL, 1982; MANGOLD AND BOLETZKY, 1988) it seems that its presence is increasingly frequent as by- catch in some pelagic fisheries, particu- larly near the coast (EZZEDDINE-NAJAL, 1996). A pair of animals, male and female, were observed by divers in a submarine cave off the coast of Almeria (Southeast Spain) relatively near the place where the sample reported in this paper was collected (GUERRA, 1992). This paper deals with the second record of the egg mass of T. rhombus in 126 the Mediterranean after 67 years. A des- cription of the planktonic paralarvae is given, emphasising several characteris- tics which may be used to identify the early stages of the species. MATERIAL AND METHODS The egg mass was discovered by the vessel “Toftevaag” at 08.27 h on August o MIS ARAS NS ME SS 18 the western Mediterranean (Fig. 1). The reported egg mass was accompanied by . other drifted pleuston, such as jelly-fish. The whole mass was so loose that it could not be taken out of the water. But a sample of gelatinous material contai- ning 2 eggs in early stage of develop- ment, 1 embryo within its egg capsule, 2 paralarvae within the egg capsules, and 32 practically fully developed paralar- vae outside the egg capsules were caught. This sample was fixed in 5% for- malin. The identification was made based on the paralarvae which, alt- hough still embedded in the external gelatinous mass, were near hatching. These paralarvae have similar characte- ristics to those reported by STEPHEN (1992). Egg diameter, dorsal mantle length (ML) and total length (TL) of each paralarvae were measured using a dissecting microscope fitted with an eye-piece graticule. RESULTS The egg mass was a dense, resilient, oblong cylinder with rounded ends (Fig. 2A). The size of the whole mass was about 100 cm in length and about 20 cm in diameter. It was observed that the purple egg capsules lay in two rows, spirally arranged around the cylinder. The diameter of the egg capsules with the well developed embryo ranged from 2.8 to 3.0 mm. The average ML of para- larvae was 1.85+0.08 mm (n= 30) and its TL varied between 2.50 and 2.75 mm. All paralarvae observed had the head inside the mantle cavity, only showing the arms and tentacles exter- GUERRA AND ROCHA: Egg mass of Thysanoteuthis rhombus in Western Mediterranean Atlantic Ocean oVigo Mind Western Mediterranean A Strait of _- Gibraltar Black Sea Ba | N Strait of Messina Mediterranean Figure 1. Thysanoteuthis rhombus. Location of the two floating egg masses collected in the Mediterranean Sea. A: This paper; B: SANZO (1929). Figura 1. Thysanoteuthis rhombus. Localización geográfica de las dos puestas pelágicas encontradas en el Mar Mediterráneo. A: Presente estudio; B: SANZO (1929). nally (Fig. 2B). The mantle is oval, stout, short and blunt posteriorly. The anterior margin of the mantle curves inwards in both dorsal and ventral sides, but is more pronounced ventrally. This may be a result of the preservation process which could had produced the retrac- tion of the head inside the mantle cavity. The fins are subterminal, small and rounded; the fin length 19.5% ML (Figs. 2C, D). The paralarvae have broadly separated, slightly protruding eyes, and funnel locking cartilage (Fig. 2E) with a short, broad, transverse groove and a long, relatively wide, longitudinal groove (sideways Fshaped). The tenta- cles are short (about 33% of the ML), stouter and slightly longer than the longest arm (III); the I and IV pairs of arms are rudimentaries. Brachial formu- lae IMI>II>I=IV. Both, tentacles and deve- loped arms, with small suckers, pro- bably arranged in two rows. On arms II and III, an incipient swimming keel-like shaped membrane was present. Trabe- culate protective membrane was absent in arms and tentacles. The paralarvae show two types of chromatophores: a) large and pale-ochre chromatophores densely concentrated on dorsal, lateral and ventral mantle sides; and b) small, subtriangular dark- red chromatophores arranged in a single row around the anterior margin of the mantle. There is a light area between both types of chromatophores. Slight chromatophores were observed on the dorsal and ventral sides of the head, the tentacles and the arms. The fins lacked chromatophores. DISCUSSION The egg mass reported was captured near the surface in a zone were the in- flow of Atlantic water into the Medite- rranean is high due to the proximity of the Strait of Gibraltar. The water move- ments through this bottleneck are gover- ned by an inflow of surface water into the Mediterranean, and a countercurrent of lesser volume carrying water of hig- her salinity into the Atlantic (MANGOLD AND BOLETZKY, 1988). The egg mass co- llected by SANZO (1929) was in the Strait of Messina where there are strong cu- rrents. Elsewhere egg masses of Thysa- noteuthis rhombus occurred in regions with strong warm currents such as Ku- roshio, Perú countercurrent and the Equatorial countercurrent (YAMAMOTO 127 Iberus, 15 (1), 1997 ARS añ Pe Figure 2. Thysanoteuthis rhombus. Floating egg mass and paralarvae. A: egg mass; B: Egg with non- developed embryo and paralarvae within the egg capsule; C: Funnel locking-cartilage of a newly hatched, 1.85 mm ML; D: Dorsal view of a newly hatched, 1.85 mm ML; E: Ventral view of the same specimen. * Figura 2. Yhysanoteuthis rhombus. Puesta pelágica y paralarva. A: Puesta pelágica; B: Huevo con embrión muy poco desarrollado y paralarva dentro de la cápsula ovígera; C: Cartilago de cierre en el sifón de un recién nacido de 1,85 mm ML; D: Visión dorsal de un recién nacido de 1,85 mm ML; E: Visión ventral del mismo ejemplar. 128 GUERRA AND ROCHA: Egg mass of Thysanoteuthis rhombus in Western Mediterranean AND OKUTANI, 1975; NIGMATULLIN ET AL., 1995). Therefore, as in the Atlantic and the Pacific Oceans, in the Medite- rranean the species seems to spawn in waters with strong currents. NIGMATULLIN ET AL. (1995) indicated that T. rhombus spawns throughout the year in tropical waters, but during the warm season (summer and early autumn) in peripheral regions such as in the Me- diterranean, which agree with the date when the egg mass reported was observed. The egg mass in the report had a shape and a size which coincide with those given for the other egg masses illustrated (SANZO, 1929; MISAKI AND OKUTANI, 1976) and photographed (SUZUKI ET AL., 1979). Considering the dimension of this egg mass, the diameter of the egg capsu- les measured and calculating the surface of the egg mass as a cylinder (6,280 cm?), an estimation gives a figure of about 66,800 eggs. This amount coincides with the total number of eggs in each egg mass calculated by SABIROV ET AL. (1987) which ranged from 32,000 to 76,000 eggs. The embryo and paralarvae found have sizes, shapes and characters which BIBLIOOGRAPHY BIAGI, V., 1982. Sul rinvenimento di un gio- vane esemplare di Thysanoteuthis rhombus Troschel (Cephalopoda - Teuthoidea) in acque Elbane. Bollettino Malacologico, 18 (7- 8): 137-144. CLARKE, M. R., 1966. A review of the systema- tics and ecology of oceanic squids. Advances in Marine Biology, 4: 91-300. 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Notes on the occurence and biology of the oceanic squid, Thysanoteuthis rhombus Troschel, in Japan. Publications of the Seto Marine Biological Laboratory, 13 (4): 327- 349. SABIROV, R. M., ARKHIPKIN, A. I., TSYGANKOV, YU AND SHCHETINNIKOV, A. S., 1987. Egg la- ying and embrional development of dia- mond-shaped squid Thysanoteuthis rhombus (Oegopsida, Thysanoteuthidae). Zoological Journal USSR, 66 (8): 1155-1163. SANZO, L., 1929. Nidamento pelagico, uova elarve di Thysanoteuthis rhombus Troschel. Memorie Reale Comitato Talassografico Italiano, 161: 1-10. 130 STEPHEN, S. J., 1992. Family Thysanoteuthidae Keferstein, 1866. In Sweeney, M. J. et al. (Eds.), “Larval” and juvenile cephalopods: a ma- nual for their identification. Smithsonian Con- tributions to Zoology, 513: 121-123. SUZUKI, S., MISAKI, H. AND OKUTANI, T., 1979. Studies on early life history of decapodan Mollusca - VILA supplementary note on flo- ating egg mass of Thysanoteuthis rhombus Troschel in Japan - The first underwater pho- tography. Venus, The Japanese Journal of Ma- lacology, 38 (2): 153-155. YAMAMOTO, K. AND OKUTANI, T., 1975. Studies on early life history of decapodan Mollusca - V. Systematics and distribution of epipela- gic larvae of decapod cephalopods in the south-western waters of Japan during the summer in 1970. Bulletin Tokai Regional Fis- hery Research Laboratory, 83: 45-96. O Sociedad Española de Malacología ——_—_—_—_——— Iberus, 15 (1): 131-138, 1997 Ontogenetic variation of statolith shape in the short-finned squid lex coindetíz (Mollusca, Cephalopoda) Variacion ontogénica del estatolito de la pota /llex coindetii (Mollusca, Cephalopoda) Ángel E GONZÁLEZ and Ángel GUERRA* Recibido el 27-X1-1996. Aceptado el 10-11-1997 ABSTRACT Changes in statolith morphology of Illex coindetii are described from specimens ranging from 42 to 379 mm mantle length obtained during trawling activities in the North-eastern Atlantic. The growth of the statolith was differentiated in five developmental stages. lt has been observed that the wing of the statolith has two different growth patterns, from the ventral zone to the dorsal one and viceversa. The rostral angle of the statolith varied during its ontogenetic growth from an obtuse angle to a 90? angle. The dorsal zone of the statolith is here named as dorsal dome, and the lateral and ventral zone the lateral dome. RESUMEN En este trabajo se describen los cambios en la morfología del estatolito de /llex coindeti. El estudio se llevo a cabo mediante el análisis de 341 ejemplares con longitudes del manto comprendidas entre 42 y 379 mm. Estos animales se capturaron en la pesquería de arrastre desarrollada en el Atlántico noreste. El crecimiento del estatolito fue diferen- ciado en cinco estadíos de desarrollo. Se observaron dos patrones diferentes de creci- miento del ala del estatolito, desde la parte ventral a la dorsal y viceversa. El angulo ros- tral varía en el crecimiento ontogénico del animal, desde un ángulo marcadamente obtuso a un ángulo recto. El crecimiento en longitud máxima y en anchura del estatolito se enlen- tece al llegar a la maduración de los animales. Se propone una variación en la nomencla- tura del estatolito de esta especie. KEY WORDS: /llex coindetiz, statolith, morphology, North-eastern Atlantic. PALABRAS CLAVE: /llex coindetiz, estatolito, morfología, Atlántico noreste. INTRODUCTION Cephalopods play an important role components of the diet of such top pre- in the trophic web of marine ecosystems dators as marine mammals (XAMPENY as both predators and prey of many ma- AND FILELLA, 1976; CLARKE, 1980, 1986; rine species (AMARATUNGA, 1983). These CLARKE, MARTINS AND PRINCE, 1993; marine molluscs have been cited as GONZÁLEZ, LÓPEZ, GUERRA AND BA- * Instituto de Investigaciones Marinas (Consejo Superior de Investigaciones Científicas), Eduardo Cabello 6, 36208 Vigo, Spain. SA Iberus, 15 (1), 1997 RREIRO, 1994), large teleosteans (BOUXIN AND LEGENDRE, 1936; BELLO, 1991; GUE- RRA, SIMÓN AND GONZÁLEZ, 1993; CLARKE, CLARKE, MARTINS AND SILVA, 1995), sea-birds (RODHOUSE, CLARKE AND MURRAY 1987; CLARKE, CROAXALL AND PRINCE, 1991; FURNESS, 1994; CROA- XALL AND PRINCE, 1996) and cephalo- pods (RASERO, GONZÁLEZ, CASTRO AND GUERRA, 1996, RODHOUSE AND NIGMA- TULLIN, 1996). Beaks, statoliths, chiti- nous sucker rings and, to a lesser extent, gladius, were the main structures which allowed a positive identification to the taxonomic level of species in studies about trophic relationships between cephalopods and other marine animals. These hard structures remain unaltered in the stomach contents of their preda- tors and represent an important source of information on the trophic relations- hips where these animals are involved. Statoliths of cephalopods are small hard paired structures composed by cal- cium carbonate in the form of aragonite. They are situated in fluid-filled cavities termed statocysts inside the cartilagi- nous skulls of the cephalopods belon- ging to the subclass Coleoidea (CLARKE AND MADDOCK, 1988a). The statolith and the macula (statoconia system) constitute the receptor organ for detec- tion of gravity. This is one of the func- tions of the statocysts, in which the level of sophistication is equivalent to the vertebrate vestibular system (YOUNG, 1960, 1989; STEPHEN AND YOUNG, 1982; BUDELMANN, 1978, 1988, 1990). Statoliths are also the structures most frequently used for studies on age and growth. The statoliths of many cep- halopod species show growth incre- ments, which have been shown to have a daily periodicity of deposition in seve- ral species (see JACKSON, 1994). Since teuthoid statoliths are appa- rently species-characteristic and have a greater likelihood of fossilisation than other cephalopod structures, they have become very important for identifi- cation of fossil species (CLARKE AND Frrch, 1975, 1979). The applications of image analysis which have been used in the morphological study of the stato- 132 liths, both recent and fossil, have shed light on certain phylogenetic relations- hips among cephalopods (CLARKE AND MADDOCK, 1988a; 1988b). As was done with shape analysis of fish otoliths (CAMPANA AND CASSELMAN, 1993), the statolith morphology of cephalopods has been used also for stock discrimina- tion (BORGES, 1995). As ontogenetic changes do exist in the statolith (MORRIS AND ALDRICH, 1984; GUERRA AND SÁNCHEZ, 1985; CLARKE AND MADDOCK, 1988b; Bkru- NETTI AND IvANOVIC, 1991), some remains on the reliability of this appro- ach about this subject. From descrip- tions based solely on one statolith from one specimen, the result of the analysis of shape can be altered (LOMBARTE, SÁNCHEZ AND MORALES-NÍN, 1995) and a correct prey identification could also be uncertain. The aim of this study was to deter- mine the changes in the statolith shape for the ommastrephid squid lllex coinde- ti1 (Vérany, 1839) during its ontogenetic growth. MATERIALS AND METHODS 341 specimens of Illex coindetii were collected in the North-eastern Atlantic (Fig. 1) from November 1991 to October 1992. Fishing was carried out at depths ranging from 100 and 350 m over the Galician continental shelf. The animals were sexed, measured to the nearest mm mantle length (ML), weighed (to 0.1 g) and assigned a maturity stage accor- ding to LIPINSKI (1979). The squid ran- ged from 48 to 379 mm ML. The stato- liths were removed from the head and preserved in 96% ethanol. Statolith ma- jor axis and maximum width were re- corded. Then, the statoliths were measu- red using an eyepiece graticule. Terms used in descriptions were assigned fo- llowing the nomenclature established by CLARKE (1978). Measurements were made using an image analysis system (IAS); the equip- ment used is reviewed by Macy (1995). The description of the each develop- GONZÁLEZ AND GUERRA: Growth of /llex coindetiz statolith $ Vigo Miño River , Burela. Celeiro -» z 8 W 1 Figure 1. /llex coindetiz. Fishing area where the samples were obtained. Figura 1. Illex coindetii. Area de pesca donde se obtuvieron las muestras. mental stage of the statolith was made based on microphotographs. RESULTS Statolith morphology: Statolith maximum length ranged from 0.47 (female of 48 mm ML) to 1.66 mm (female of 360 mm ML). No significant differences (p<0.05) between maximum length and maximum width of male and female statoliths of equivalent ML were found at any stage of development. Therefore, sex does not affect the growth of the statolith in /llex coindetii. Figure 2 shows a diagrammatic picture of the different parts of an adult Illex coin- detii statolith in posterior and anterior view. Terms used in descriptions and the measurements made are also shown. Some characteristic features were obser- ved in these statoliths: a) there is conti- nuity between the dorsal dome and the lateral dome. This feature can be obser- ved between the superior and inferior lobes of the lateral dome as well; b) the posterior dome groove is very patent; c) the rostrum is short and an anterior rostral lobe does not exist; d) the wing is very broad; e) the medial fissure is well deve- loped and also has a small posterior inden- tation; £) the dorsal spur is very clear. Statolith development: Although the above description refers to a late developmental stage of the statolith in an adult animal, the way to reach this definitive stage is quite complex. Thus, the statolith has become increasingly complex and passed through different stages which implies growth in different planes. These stages can be described as follows (Fig. 3): Stage I: Statoliths of immature animals ranging from 50 to 80 mm ML. The medial fissure is not yet visible but will be situated under the dorsal dome. This area will be the surface where the wing will connect with the body of the 183 Iberus, 15 (1), 1997 Dorsal dome Dorsal indentation Medial fissure == 2 Wing Ventral indentation Rostral angle Rostrum Superior lobe Lateral lobe Inferior lobe Posterior dome groove Rostrum Dorsal dome / Posterior indentation Wing Maximum length Maximum width Figure 2. Diagrammatic picture of the different parts of an adult /llex coindetiz statolith. Figura 2. Diferentes partes del estatolito de un Vlex coindetii adulto. statolith in a later developmental stage. Another characteristic of the stage 1 sta- tolith is the wide rostral angle (>140"). The primordium of the rostrum is also visible in the ventral zone of the stato- lith. It is virtually impossible to differen- tiate the dorsal dome, the superior and inferior lobes of the lateral dome, being round the general shape. Stage II: This developmental stage includes the statoliths of animals with ML between 90 and 130 mm. The main cha- racteristic of this stage is the growth of the rostrum, and the change in shape of the statolith, which is enlarging the ventral direction. Small dark zones in the ventral zone can be distinguished. This crystalli- sation will form the wing of the statolith. The wing formation is observed in two directions, from the rostrum to the dorsal plane of the statolith and from the dorsal dome to the ventral plane. There is a slight differentiation between superior and medium lateral dome. Stage III: This stage is remarkably dif- ferent from the preceding one. It appears in submature and mature squid ranging from 130 to 200 mm ML. The enlargement of the rostrum continues and the forma- tion of the wing spans from the ventral to the dorsal zone. The medial fissure is small. A zone devoid of crystallisation called the foramen appears for the first 134 time. The foramen runs parallel to the rostrum and will disappear gradually. The rostral angle is getting narrow and at this stage 1t forms almost a 90” angle. Stage IV: Mature animals between 200 and 250 mm ML. The developmen- tal stage is close to definitive conditions. The foramen is partially or totally occlu- ded and the wing is formed along the entire statolith. As the statolith grows, the lobes become more distinct. Practi- cally, the rostral angle is 90". Stage V: This stage describes the sta- toliths of mature specimens bigger than 250 mm ML. There are only minor changes in morphology from stage IV. Crystallisation in the wing is stronger and the foramen is totally occluded. The formation of the wing emphasises the medial fissure. Figure 4 illustrates the relationship between the mantle length and the maximum length and width of the lllex coindetii statolith. DISCUSSION This paper gives a description of dif- ferent developmental stages of Illex coin- detii statoliths, based on observations of the statolith growth. There are impor- tant changes in shape of the statolith GONZÁLEZ AND GUERRA: Growth of /llex coindetíz statolith Figure 3. Stages of development for the statolith of //lex coindetiz. A: Stage 1, maximum statolith length (MSL)= 0.67 mm and maximum statolith width (MSW)= 0.40 mm; B: Stage II, MSL =0.90 mm and MSW= 0.63 mm; C: Stage III, MSL= 1.13 mm and MSW= 0.80 mm; D: Stage IV, MSL= 1.20 mm and MSW= 0.80 mm; E: Stage V, MSL= 1.53 mm and MSW= 1.03 mm. Figura 3. Estadios de desarrollo del estatolito de Ylex coindetii. A: Estadío 1, longitud máxima del estato- lito (LME)= 0,67 mm y anchura máxima del estatolito (AME)= 0,40 mm; B: Estadío 1H, LME= 0,90 mm y AME= 0,63 mm; C: Estadío 11, LME 1,13 mm y AME= 0,80 mm; D: Estadío IV, LME= 1,20 mm y AME= 0,80 mm, E: Estadío V., LME= 1,53 mm y AME= 1,03 mm. through the life cycle of Illex coindeti. Considering the statolith of ommastrep- hids, it is important to note the difficulty in differentiating between the dorsal dome and the superior and inferior lobes described by CLARKE (1978) for teuthoids. This observation agrees with SÁNCHEZ (1981) for Illex coindetii speci- mens from Mediterranean waters and ARKHIPKIN (1990) and BRUNETTI AND IVANOVIC (1991) for Illex argentinus. It was also observed that the rostral angle for statoliths of Illex coindetii juveniles is clearly higher than 140? and progressi- vely it is getting narrower until it reaches 90? in mature animals. Stages I and Il correspond to speci- mens ranging from three to five months of age. The description coincides with stages I and II defined for Illex argenti- nus (BRUNETTI AND IVANOVIC, 1991) and the “definitive stage” noted by Morris and ALDRICH (1984). The shape of the statolith is enlarged to the ventral side and it grows in the ventral zone. The formation of the wing converges from two different directions, from the dorsal zone to the ventral one and vice-versa. Stage III of the statolith of lllex coin- detii corresponds to submature and mature animals between five and eight months of age ranging from 130 to 200 185 Iberus, 15 (1), 1997 Statolith measurements (mm) —» Mantle length (mm) O Maximum length (mm) + Maximum width (mm) Figure 4. /llex coindetii. Relationship between mantle length and maximum length and width . Figura 4. Mex coindetii. Relación entre la longitud del manto y la anchura y longitud máximas. mm ML. The union of the wing with the dorsal dome of the statolith was obser- ved in this stage, which agrees with the observations Of MORRIS AND ALDRICH (1984) for the “juvenile stage” of Illex illecebrosus and stage III of ARKHIPKIN (1990) and BRUNETTI AND IVANOVIC (1991) for Illex argentinus. From this stage onwards, the growth of the stato- lith slows down; this could be related with the process of maturation of the animals as showed in the Figure 4. In stage III appears for the first time the foramen, which was noted as a characte- ristic feature of ommastrephids. This is a lack of crystallisation that runs parallel to the rostrum. It is formed when both planes of the wing grow, connecting over the medial part of the statolith, leaving one distinct zone between this point and the body of the statolith. Stages IV and V are very similar in shape. However, a main difference can be found when the foramen disappears 136 completely in stage V. Stage IV of Illex coindetii corresponded to animals of between 200 and 250 mm ML and ages ranging from eight to ten months. Finally, the statoliths of animals bigger than 250 mm ML and older than ten months are included in the stage V. These statoliths are similar to those des- cribed by MORRIS AND ALDRICH (1984) for the “advanced stage” in statoliths of Illex illecebrosus, to the stage VI observed by BRUNETTI AND IVANOVIC (1991) for Illex argentinus and the statolith descri- bed by SÁNCHEZ (1981) for an adult /llex coindetii specimen from the Mediterra- nean Sea. On the whole, it can be concluded that the shape of the statolith changes gra- dually from the juvenile stage until it reaches a definitive stage of development. 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Datos sobre tres cachalotes Physter macrocephalus L. cap- turados frente a la costas atlánticas de Gali- cia, España (Cetacea, Physeteridae). Misce- lánea Zoológica, 3 (5): 235-242. YOUNG, J. Z., 1960. The statocysts of Octopus vul- garis. Proceedings of the Royal Society of London B, 152: 3-29. YOUNG, J. Z., 1989. The angular acceleration re- ceptor system of diverse cephalopods. Phi- losophical Transactions of the Royal Society of London B, 25: 189-237. NORMAS DE PUBLICACIÓN e La revista IBERUS publica artículos de fondo, notas y monografías que versen sobre cualquiera de los aspectos rela- cionados con la Malacología. Se entiende por artículo un trabajo de investigación de más de 5 páginas de texto, mecanografiadas, incluidas láminas, gráficos y tablas. Las notas son trabajos de menor extensión. Las monografías son trabajos sobre un tema único, de extensión superior a las 50 páginas de la revista y que serán publicadas como un suplemento de IBERUS. Los autores interesados en publicar monografías deberán ponerse previamente en contacto con el Editor de Publicaciones. Se entiende que el contenido de los manuscritos no ha sido publicado, ni se publicará en otra parte ni en su totalidad ni resumido. * Los manuscritos, así como toda la correspondencia relacionada con los mismos, deberán ser remitidos a: Dr. Ángel Guerra Sierra, Editor de Publicaciones, Instituto de Investigaciones Marinas (CSIC), C/Eduardo Cabello 6, 36208 Vigo, España. » El texto del trabajo podrá estar redactado en cualquier lengua culta moderna. * En los trabajos que superen las 20 páginas, cada página de exceso será abonada por el(los) autor(es) a precio de coste. * Los artículos, notas y monografías deberán presentarse mecanografiados sobre DIN A-4, por una sola cara a doble espacio (incluyendo referencias, pies de figura y tablas), con al menos 3 centímetros de margen por cada lado. Los tra- bajos se entregarán por triplicado (original y dos copias). En caso de autoría compartida, uno de los autores deberá hacerse responsable de toda la correspondencia. * Los manuscritos se presentarán de acuerdo al siguiente esquema: Primera página. Deberá incluir un título conciso, pero sugerente del contenido del trabajo, así como una traducción al inglés del mismo (si el artículo no está escrito en inglés). Cuando sea preciso, deberá incluir referencia a familia o táxones superiores. A continuación figurarán, por este orden, el nombre y apellidos completos del autor o autores, las direcciones completas de los mismos, y un resumen del trabajo y su traducción al inglés. Dicho resumen deberá sintetizar, en conjunción con el título, tanto los resultados como las conclusiones del artículo; se sugiere una exten- sión de 100 a 200 palabras. El resumen deberá estar seguido de una lista de palabras clave, también con su versión inglesa para su inclusión en los bancos de datos internacionales. Páginas siguientes. Incluirán el resto del artículo, que debe dividirse en secciones precedidas por breves encabeza- mientos. Siempre que sea posible, se recomienda seguir el siguiente esquema: Introducción, Material y métodos, Resultados, Discusión, Conclusiones, Agradecimientos y Bibliografía. Si se emplean abreviarras no habituales en el texto, deberán indicarse tras el apartado de Material y Métodos. e Las notas breves deberán presentarse de la misma forma, pero sin resumen. * Deberán evitarse notas a pie de página y referencias cruzadas. Deberán respetarse estrictamente los Códigos Internacionales de Nomenclatura Zoológica y Botánica (últimas ediciones). Cuando un táxon aparezca por primera vez deberá citarse su autor y fecha de su descripción. En el caso de artículos sistemáticos, cuando se den las sinonimias de los táxones, éstas deberán citarse COMPLETAS, incluyendo en forma abreviada la publicación donde fueron des- critas, y la localidad tipo si es conocida entre corchetes, según el siguiente esquema (préstese especial cuidado a la pun- tuación): Dendrodoris limbata (Cuvier, 1804) Sinonimias Doris limbata Cuvier, 1804, Ann. Mus. H. N. Paris, 4 (24): 468-469 [Localidad tipo: Marsella]. Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop. Car., 10: 275. Dichas referencias no deberán incluirse en la lista de Bibliografía si es la única vez que se nombran en el texto. Si se incluyen una lista completa de referencias de un taxon inmediatamente tras éste, deberá seguirse el mismo esque- ma (sin incluir en Bibliografía las referencias que no se mencionen en otro lugar del texto). * Sólo los nombres en latín y los de táxones genéricos y específicos deberán llevar subrayado sencillo o preferentemente ir en cursiva. En ningún caso deberá escribirse una palabra totalmente en letras mayúsculas, ni siquiera el Título. Las unidades a utilizar deberán pertenecer al Sistema Métrico Decimal, junto con sus correctas abreviaturas. En artículos escritos en castellano, en los números decimales sepárese la parte entera de la decimal por una coma inferior (,), NUNCA por un punto (.) o coma superior ('). * Las referencias bibliográficas irán en el texto con minúsculas o versalitas: Fretter y Graham (1962) o FRETTER Y GRAHAM (1962). Si son más de dos autores se deberán citar todos la primera vez que aparecen en el texto [Smith, Jones y Brown (1970)] empleándose et al. las siguientes veces [Smith et al. (1970)]. Si un autor ha publicado más de un trabajo en un año se citarán con letras: (Davis, 1989a; Davis, 1989b). No deberá emplearse op. cit. La lista de referencias 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 minúsculas o VERSALITAS. No deberán incluirse referencias a autores cuando éstos aparezcan en el texto exclusiva- mente como autoridades de un taxon. Los nombres de las publicaciones periódicas deberán aparecer COMPLETOS, no abreviados. Cuando se citen libros, dése el título, editor, lugar de publicación, n* de edición si no es la primera y número total de páginas. Deberán evitarse referencias a Tesis Doctorales u otros documentos inéditos de difícil con- sulta. Síganse los siguientes ejemplos (préstese atención a la puntuación): Eretter, V. y Graham, A., 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. Ponder, W. F., 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. En Ponder, W. EF. (Ed.): Prosobranch Phylogeny, Malacological Review, suppl. 4: 129-166. Ros, J., 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Miscelánea Zoolgica, 3 (5): 21-51. í e Las gráficas e ilustraciones deberán ser originales y presentarse sobre papel vegetal o similar, con tinta china negra y ajustadas al formato de caja de la revista o proporcional a éste. Este formato es de 57 mm (una columna) o 120 mm (dos) de anchura y hasta 194 mm de altura, si bien se recomienda utilizar el formato a dos columnas. En caso de pre- parar figuras para que ocupen el total de una página, se ruega ajustar su tamaño para que puedan caber los pies de figura bajo ella. Si han de incluirse gráficas de ordenador, deberán imprimirse con impresora láser sobre papel de buena calidad. Las fotografías, bien contrastadas y sin retocar, deberán ajustarse siempre a los tamaños mencionados. Al componer fotografías sobre una hoja, procúrese que los espacios entre ellas sean regulares y que estén debidamente alineadas. Téngase en cuenta que incluir fotografías de distinto contraste en una misma página conlleva una pobre reproducción final. Las escalas de dibujos y fotografías deberán ser gráficas, y las unidades que se utilicen del sistema métrico decimal. Considérese la reducción que será necesaria a la hora de decidir el tamaño de las escalas o letras en las figuras, que no deberán bajar de los 2 mm. En figuras compuestas, cada parte deberá etiquetarse con letras mayús- culas, el resto de las letras deberán ser minúsculas. No deberán hacerse referencias a los aumentos de una determi- nada ilustración, ya que éstos cambian con la reducción, por lo que debe emplearse una escala gráfica. En su caso, se recomienda la utilización de mapas con proyección UTM. Cada figura, gráfica o ilustración deberá presentarse en hojas separadas y con numeración arábiga (1, 2, 3,...), sin separar “Figuras” y “Láminas”. Los pies de figura, en una hoja aparte, deberán acompañarse de su traducción al inglés. Utilícese el esquema siguiente: Figura 1. Neodoris carvi. A: animal desplazándose; B: detalle de un rinóforo; C: branquia. Las abreviaturas empleadas en las ilustraciones deberán incluirse en la hoja de pies de figura. Los autores interesados en incluir láminas en color deberán abonarlas a precio de coste (30.000 ptas por página). Por lo demás, deberán ajustarse a los mismos requisitos que los indicados para las figuras. + Las Tablas se presentarán en hojas separadas, siempre con numeración romana (1, II, IL...). Las leyendas se inclui- rán en una hoja aparte acompañándose de una traducción al inglés. Deberán evitarse las tablas particularmente com- plejas. Se recomienda reducir el número y extensión de ilustraciones, láminas o tablas al mínimo necesario. * Los artículos que no se ajusten a las normas de publicación serán devueltos al autor con las indicaciones de los cam- - bios necesarios. + El Comité Editorial comunicará al autor responsable del trabajo la fecha de recepción del trabajo y la fecha de envío a revisión. Cada original recibido será sometido a revisión por al menos dos investigadores. El Comité Editorial, a la vista de los informes de los revisores decidirá sobre la aceptación o no de cada manuscrito. El autor recibirá en cada caso copia de los comentarios de los revisores sobre su artículo. En caso de aceptación, el mismo Comité Editorial, si lo considera conveniente, podrá solicitar a los autores otras modificaciones que considere oportunas. Si el trabajo es aceptado, el autor deberá enviar una copia impresa del mismo corregida, acompañada por una versión en disco flexi- ble (diskette), utilizando procesadores de texto en sus versiones de DOS o Macintosh. La fecha de aceptación figura- rá en el artículo publicado. * Las pruebas de imprenta serán enviadas al autor responsable, EXCLUSIVAMENTE para la corrección de erratas, y deberán ser devueltas en un plazo máximo de 15 días. Se recomienda prestar especial atención en la corrección de las pruebas. * De cada trabajo se entregarán gratuitamente 50 separatas. Aquellos autores que deseen un número mayor, deberán hacerlo constar al devolver las pruebas de imprenta, y NUNCA POSTERIORMENTE. El coste de las separatas adi- cionales será cargado al autor. INSTRUCTIONS TO AUTHORS » 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: Dr. Ángel Guerra Sierra, Editor de Publi - caciones, Instituto de Investigaciones Marinas (CSIC), C/Eduardo Cabello 6, 36208 Vigo, 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. When a paper has joint authorship, one author must accept responsability for all correspondence. » Papers should conform the following layout: First page. This must include a concise but informative title, with mention of family of higher taxon when appropriatte, and its Spanish translation. It will be followed by all authors? names and surnames, their full adress(es), an abstract (and its Spanish translation) not exceeding 200 words which summarizes not only contents but results and conclusions, and a list of Key Words (and their Spanish translation) under which the article should be indexed. Following pages. These should content the rest of the paper, divided into sections under short headings. Whenever pos- sible the text should be arranged as follows: Introduction, Material and methods, Results, Discussion, Conclusions, Acknowledgements and References. Unusual abbreviations used in the text must be grouped in one alphabetic sequence after the Material and methods section. e Notes should follow the same layout, without the abstract. * Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical Nomencla- ture must be strictly followed. The first mention in the text of any taxon must be followed by its authority including the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN FULL, including the perio- dical, in an abbreviate form, where they were described, and the type localities in square brackets when known. Follow this example (please note the punctuation): Dendrodoris limbata (Cuvier, 1804) Synonyms Doris limbata Cuvier, 1804, Ann. Mus. H. N. Paris, 4 (24): 468-469 [Type locality: Marseille]. Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop. Car., 10: 275. These references must not be included in the Bibliography list, except if referred to elsewhere in the text. 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 (*). e References in the text should be written in small letters or SMALL CAPITALS: Fretter 8Z Graham (1962) or FRETTER $ 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. ci. The references in the reference list should be in alphabetical order and include all the publications cited in the text but only these. ALL the authors of a paper must be included. These should be written in small letters or SMALL CAPITALS. The references need not be cited when the author and date are given only as authority for a taxonomic name. Titles of periodicals must be given IN FULL, not abbreviated. For books, give the title, name of publisher, place of publication, indication of edition if not the first and total number of pages. Keep references to doctoral theses or any other unpu- blished documents to an absolute minimum. See the following examples (please note the punctuation): Eretter, V. and Graham, A., 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. Ponder, W. F., 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. In Ponder, W/. F. (Ed.): Prosobranch Phylogeny, Malacological Review, suppl. 4: 129-166. Ros, J., 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Miscelá- nea Zoológica, 3 (5): 21-51. + Figures must be original, in Indian ink on draughtsman's tracing paper. Keep in mind page format and column size when designing figures. These should be one column (57 mm) or two columns (120 mm) wide and up 194 mm high, or be proportional to these sizes. Two columns format is recomended. It is desirable to print figures with their legend below, so authors are asked to take this into account when preparing full page figures. If computer generated graphics are to be included, they must be printed on high quality white paper with a laser printer. Photographs must be of good contrast, and should be submitted in the final size. When mounting photographs in a block, ensure spacers are of uni- form width. Remember that grouping photographs of varied contrast results in poor reproduction. Take account of necessary reduction in lettering drawings; final lettering must be at least 2 mm high. In composite drawings, each figure should be given a capital letter; additional lettering should be in lower-case letters. A scale line is recomended to indi- cate size, magnification ratio must be avoided as it may be changed during printing. UTM maps are to be used if neces- sary. Figures must be submitted on separate sheets, and numbered with consecutive Arabic numbers (1, 2, 3,...), without separating Plates' and “Figures”. Legends for Figures must be typed in numerical order on a separate sheet, and an English 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, 300 US$ per page). They should be submitted in the same way that black and white prints. e Tables must be numbered with Roman numbers (1, II, II...) and each typed on a separate sheet. Headings should be typed on a separate sheet, together with their English translation. Complex tables should be avoided. As a general rule, keep the number and extension of illustrations and tables as reduced as possible. + Manuscripts that do not conform to these instructions will be returned for correction before reviewing. * Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the date the manuscript was sent for reviewing. Each manuscript will be critically evaluated by at least two referees. Based of these evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors will receive a copy of the refe- rees” comments. If a manuscript is accepted, the Editorial Board may indicate additional changes if desirable. Accep- table manuscripts will be returned to the author for consideration of comments and criticism; a finalized manuscript must then be returned to the Editor, together with a floppy disk containing the article written with a DOS or Macin- tosh word processor. Dates of reception and acceptance of the manuscript will appear in all published articles. * Proofs will be sent to the author for correcting errors. At this stage no stylistic changes will be accepted. Pay special attention to references and their dates in the text and the Bibliography section, and also to numbers of Figures and Tables appearing in the text. e Fifty reprints per article will be supplied free of charge. Additional reprints must be ordered when the page proofs are returned, and will be charged at cost. NO LATER orders will be accepted. -—. La SocieDAD ESPAÑOLA DE IMALACOLOGÍA Junta directiva desde el 18 de octubre de 1996 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 Ángel Guerra Sierra Instituto de Investigaciones Marinas, c/ Eduardo Cabello 6, 36208 Vigo, España Bibliotecario Rafael Araujo Armero Museo Nacional de Ciencias Naturales, CSIC, c/ José Gutierrez Abascal 2, 28006 Madrid, España Vocales Eugenia María Martínez Cueto-Felgueroso María de los Ángeles Ramos Sánchez Francisco Javier Rocha Valdés Gonzalo Rodríguez Casero 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 1997: Socio numerario (en España): 5.000 ptas. (= 50 U.S. $) (en extranjero): 7.000 ptas (= 70 U.S. $) Socio estudiante: 2.000 ptas. (= 20 U.S. $) Socio Familiar: 500 ptas. (= 5 U.S. $) Socio Protector: 6.000 ptas. (= 60 U.S. $) (mínimo) Socio Corporativo 6.000 ptas. (= 60 U.S. $) INSCRIPCIÓN: 1.000 ptas. (= 10 U.S. $) 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. Members living in foreing countries can deduce 10 U.S. $ if paid before 15 April. Cada socio tiene derecho a recibir anualmente los números de /berus, Reseñas Malacológicas y Noticiarios que se publiquen. 50 Iberus E Revista de la SOCIEDAD ESPAÑOLA DE MALACOLOGÍA tu COMITÉ DE REDACCIÓN EDITOR Ce Ángel Guerra Sierra a Instituto de Investigaciones Marinas, CSIC, Vigo, España AS EDITORES ADJUNTOS * Eugenia M* Martínez Cueto-Felgueroso Universidad de Oviedo, Oviedo, España Francisco Javier Rocha Valdés Instituto de Investigaciones Marinas, CSIC, Vigo, España Gonzalo Rodríguez Casero Universidad de Oviedo, Oviedo, España ComiTÉ EDITORIAL Kepa Altonaga Sustacha Universidad del País Vasco, Bilbao, España Eduardo Angulo Pinedo Universidad del País Vasco, Bilbao, España Thierry Bockeljau Institut Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica Sigurd v. Boletzky Loboratoire Arago, Bonyuls-surMer, Francia Jose Castillejo Murillo Universidad de Santiago de Compostela, Santiago de Compostela, España Karl Edlinger Noturhistorisches Museum Wien, Austria José Carlos García Gómez Universidad de Sevilla, Sevilla, España Edmund Gittenberger Notionaal Natuurhistorisch Museum, Leiden, Holanda Serge Gofas Muséum Notional d'Histoire Naturelle, Paris, Francia Ángel Antonio Luque del Villar Universidad Autónoma de Madrid, Madrid, España María Yolanda Manga González Estación Agrícola Experimental, CSIC, León, España Jordi Martinell Callico Universidad de Barcelona, Barcelona, España Ron K. 0'Dor Dalhousie University, Halifax, Canada Marco Oliverio Universitá di Roma “La Sapienza”, Roma, Italia Pablo E. Penchaszadeh Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina Carlos Enrique Prieto Sierra Universidad del País Vasco, Bilbao, España María de los Ángeles Ramos Sánchez Museo Nacional de Ciencias Naturales, CSIC, Madrid, España Paul 6. Rodhouse British Antarctic Survey, Cambridge, Reino Unido Joandoménec Ros ¡ Aragones Universidad de Barcelona, Barcelona, España María del Carmen Salas Casanovas Universidad de Málaga, Málaga, España Gerhard Steiner Universitát Wien, Austria José Templado González Museo Nacional de Ciencias Naturales, CSIC, Madrid, España Victoriano Urgorri Carrasco Universidad de Santiago de Compostela, Santiago de Compostela, España Anders Warén Swedish Museum of Natural History, Estocolmo, Suecia 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: Dr. Ángel Guerra Sierra, Instituto de Investigaciones Marinas (CSIC), c/ Eduardo Cabello 6, 36208 Vigo, España. Los trabajos se entregarán por triplicado (original y dos copias). Se recomienda a los autores leer cuidadosa- mente 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 dirigirse 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 Electrónico: backhuysCeuronet.nl PORTADA DE /berus Iberus gualterianus (Linnaeus, 1758), una especie emblemática de la península Ibérica, que da nombre a la revista. Dibujo realizado por José Luis González Rebollar “Toza”. LIBRARY Iberus =. HARVA UNI VERS In y REVISTA DE LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA Vol. 15 (2) Oviedo, diciembre 1997 Dep. Leg. B-43072-81 ISSN 0212-3010 Diseño y maquetación: Gonzalo Rodríguez Impresión: LOREDO, S. L. - Gijón PREFACE This volume of /berus comprises a miscellany of various papers and posters presented at the Twelfth International Malacological Congress held at Vigo (Spain) from 3 Septem- ber to 8 September, 1996. This congress was organised by Dr. Ángel Guerra and Dr. Fran- cisco Rocha, members of the Instituto de Investigaciones Marinas (IM) on behalf of Unitas Malacologica and under the auspices of the Consejo Superior de Investigaciones Científi- cas (CSIC), the Sociedad Española de Malacología (SEM) and the Cephalopod Interna- tional Advisory Council (CIAC). Based at the Cultutal Center of the Caixavigo and the adjacent Casa das Artes do Con- cello de Vigo, the congress was attended by four houndred and twenty one specialists on molluscs from different research fields of fifty seven countries. The framework of the Vigo Congress was composed by five symposia, three free lectures and four workshops. These ranged over all Classes of Mollusca, the marine, freswater and land environments, evolu- tion and fossil records, phylogeny and systematics, ecology, medical and applied malaco- logy, the functional morphology of cephalopods, endemisms in the marine realm, and data bases. In addition to the communications most of the time was devoted to informal mee- tings and discussions. The congress was sponsored by over fourteen Governamental and Official Institutions and companies. However, it would not be possible without the enthusiasm and the deter- mination of the Local Organising Committee compossed mainly by people working in the research group of Ecofisiología de Cefalópodos (1IM), the Universities of Vigo and San- tiago de Compostela and the Sociedad Española de Malacología (SEM). Their forsight, kindness and hard work made a very successful congress. We thank all of them and parti- cularly to the director and the manager of IIM, Dr. Ricardo Pérez Martín and Mr. Luis Ansorena, respectively. Our acknowdlegment to Dr. Emilio Rolán, President of the SEM and co-editor of the book of abstracts!, Mrs. María Teresa Fernández, the person in charge for all the secretariat affaires of the congress, and to all the organisers of the different ses- sions, who wisely managed the symposia, workshops and free lectures. We are very grate- ful to all the referees for their expert assistance reviewing these manuscripts. Delay in publi- cation of this volume is basically our responsability, but we wish to point out correspon- dence with some of the authors was not easy. In the oppening address to the Vigo Congress, Dr. Winston E Ponder argued that mala- cology is not doing justice to the importance of molluscs. Molluscs are a large phylum of large bodied, well-known animals with a superb fossil record, excellent model organisms in evolution, genetics, physiology and ecology, economically important (fisheries and cul- tures), and as agricultural pests and carriers of disease. Analysis of several experts revealed, however, that few papers on molluscs are given at international meetings or published in mainstream journals, and instead, most appear in malacological meetings and journals, and much of it is narrowly focussed and trivial, indicating an inward focus and conservatism. If malacology is to improve its perception by the scientific community, the basic research must be focussed on areas of greatest interest, the study of molluscs must contribute to major areas of scientific inquiry and social and economic concern. Our discipline also needs a strong and effective voice in the traditional (books and journals) and the new (electro- nic) media. This, and other issues in non-molluscan specialised journals which now are in press, must be regarded as an effort of the Vigo Congress organisers to reverse the trend of margination of molluscan studies in mainstream biology. This volume has been published under the inestimable collaboration of the associated editors of Iberus D. Gonzalo Rodríguez Casero and Dr. Eugenia Martínez Cueto-Felgue- roso. Nine papers are included in this volume. These articles embrace very different aspects of the malacology and comprise distincts groups of molluscs that belongs to several ecosys- tems. Thus, we find in this book studies which run from the biogeography and demo- graphic response of the snail populations to enviromental conditions to the study of mollus- can evolution, going through inmunology and morphological researches. Finally, we would like to thank the Spanish Ministry of Education and Science, the Education Ministry of the Galician Government, the manager of the 5% Centenary of the Universidad de Santiago de Compostela and the Chancellor of the Universidad de Vigo for their support to organise the “Iwelfth International Malacological Congress and spe- cially the Council of Unitas Malacologica for providing funds for publication of this volume of Iberus. Angel Guerra and Francisco Rocha | GUERRA, A., E. ROLÁN AND F. ROCHA (EDS.), 1995. Abstracts of the Twelfth International Malacological Congress. Vigo, 3th-8th September, 1995. Unitas Malacologica and Instituto de Investigaciones Marinas (CSIC), Vigo, Spain. 530 pp. O Sociedad Española de Malacología — ——_———T— lIberus, 15 (2): 1-11, 1997 Phagocytosis by haemocytes from the Lesser Octopus Ele- done cirrhosa Fagocitosis en hemocitos del pulpo blanco Eledone cirrhosa Shelagh K. MALHAM”, Norman W. RUNHAM' and Christopher J. SECOMBES” Recibido el 8-1-1996. Aceptado el 11-IV-1996 ABSTRACT Haemocytes from Eledone cirrhosa phagocytose formalized bacteria (Vibrio anguillarum). The phagocytic capabilities of E. cirrhosa haemocytes are affected by several factors, inclu- ding the haemocyte culture medium, temperature, duration of the assay, and the bacterial pre-incubation conditions such as haemolymph concentration, temperature and the duration of pre-incubation. Haemocytes will phagocytose in the absence of haemolymph. With a 30min incubation period the number of phagocytosing haemocytes increases as the pre-opsonization concen- tration and incubation temperature increase. However after 2 hours at 15 or 20*C the num- ber of haemocytes phagocytosing unopsonized bacteria is equivalent to the number engul- fing 100% haemolymph opsonized bacteria. RESUMEN Los hemocitos de Eledone cirrhosa fagocitan bacterias formalizadas (Vibrio anguillarum). La capacidad de fagocitar en estas células se ve afectada por varios factores, incluyendo el medio de cultivo de los hemocitos, temperatura, duración del experimento, y las condicio- nes de preincubación de las bacterias, tales como concentración de hemolinfa y tempera- tura y duración de la preincubación. Los hemocitos fagocitan en ausencia de hemolinfa. Con un periodo de incubación de 30 minutos, el numero de hemocitos que fagocitan se incrementa cuando lo hacen la concentración de preopsonización y la temperatura de incu- bación. Sin embargo, tras dos horas a 15 ó 20*C, el número de hemocitos que fagocitan bacterias no opsonizadas es equivalente al de hemocitos que fagocitan bacterias tratadas con hemolinfa al 100%. KEY WORDS: Eledone cirrhosa, haemocytes, phagocytosis, opsonization. PALABRAS CLAVE: Eledone cirrhosa, hemocitos, fagocitosis, opsonización. INTRODUCTION In vivo and in vitro investigations into blood cells or haemocytes are avidly pha- the cellular activities of molluscs have de- gocytic and capable of recognising non- monstrated that, in a number of cases, the self (reviewed by MILLAR AND RATCLIFEFE, “University of Wales, School of Biological Sciences, Brambell Building, North Wales, Bangor, Gwynedd LL57 2UW, United Kingdom. ” Department of Zoology, University of Aberdeen, Tillydrone Ave, Aberdeen, Scotland AB9 2T'N, U.K. Iberus, 15 (2), 1997 1994). The process of phagocytosis invol- ves a number of recognizable stages, which include attraction, attachment, in- gestion and killing of foreign organisms, and is influenced by a number of factors (reviews by RATCLIFFE, ROWLEY, FITZGE- RALD AND RHODES, 1985; MILLAR AND RATCLIFFE, 1994). Variables which have been shown to affect phagocytic rates in molluscs include incubation temperature (FOLEY AND CHENG, 1975), time and pH (ABDUL-SALAM AND MICHELSON, 1980), the size of the particle presented for pha- gocytosis and the nature of the particles (reviewed by BAYNE, 1983). Though pha- gocytosis will take place in the absence of opsonizing agents (RENWRANTZ AND STAHMER, 1983; TUAN AND YOSHINO 1987; FRYER, HULL AND BAYNE, 1989), se- veral experiments have shown that solu- ble humoral factors or opsonins may be instrumental in non-self recognition (PROWSE AND TAIT, 1969) and, or enhan- cement of phagocytosis (reviews by JEN- KIN, 1976; RATCLIFFE ET AL., 1985). The haemocyte culture medium has been shown to influence phagocytosis with, in the case of the Asian clam, Cor- bicula fluminea, the presence of divalent cations being necessary for both opsonin-independent and opsonin- dependent phagocytosis (TUAN AND YOSHINO, 1987). The process of opsoni- zation also appears to be influenced by several other factors. FRYER AND BAYNE (1989), using Biomphalaria glabrata, showed that for this mollusc opsoniza- tion is a time-dependent process. Further, TrIPP (1992), working with Mer- cenaria mercenaria demonstrated that at low temperatures, opsonization caused enhanced phagocytic rates. The octopus Eledone cirrhosa is benthic in habit, ranges in depth from sub-littoral to 770 m and encounters temperatures between 5 and 15”C (BOYLE, 1983). The animal has a closed circulatory system and if wounded pre- vents blood loss by local vasoconstric- tion of the area surrounding the wound. The blood of the octopod does not clot and further blood loss is prevented by allowing seepage of blood through the wou1.d until blood cells eventually plug the wound (WELLs, 1978, 1983; BAYNE, 1983). If the animal loses a large amount of blood a dilution of the respiratory pigment (haemocyanin) occurs which takes up to 2 hours to be reversed (WELLS AND WELLS, 1993). There appears to be only one main type of blood cell or haemocyte in E. cirrhosa. The haemocyte matures in the white body, or haematopoetic organ, of the animal and is released into the closed circulatory system (COWDEN AND Curtis, 1974, 1981). Few cephalopod defense mechanisms have been elucida- ted (FORD, 1992). It is known that E. cirr- hosa haemocytes will phagocytose eryth- rocytes only in the presence of hae- molymph in vitro (STUART, 1968). Also in vivo studies (STUART, 1968; BAYNE, 1973) using different octopods, demonstrate that it is mainly fixed phagocytes in certain organs which clear injected foreign particles, with haemocytes only removing a small fraction of them. This paper investigates whether hae- mocytes from E. cirrhosa are capable of phagocytosing dead bacteria in vitro and whether temperature, time and hae- molymph concentrations influence pha- gocytosis. Additional experiments were also performed to determine whether bacterial pre-incubation (opsonization) at different temperatures, times and haemolymph concentrations affected phagocytic rates. MATERIALS AND METHODS Animals: Octopuses, Eledone cirrhosa (Lamarck) were obtained from crab pots around the North Wales coast. The animals were brought into the aquarium at the University of Bangor and maintai- ned in natural seawater at 10-12*C. After 48 h the animals were weighed, marked using a panjet and assigned to a particular tank. Five octopuses per tank were chosen at random for each set of experiments. Haemolymph: Blood was withdrawn from the branchial blood vessel of each octopus as described by MALHAM, SECOM- MALHAM E7 AL.: Phagocytosis by haemocytes from Eledone cirrhosa BES AND RUNHAM (1995). The blood was centrifuged at 4*C for 5 min at 800g to remove the haemocytes. The resulting hae- molymph from a number of individuals was pooled and frozen at -20*C. Before use the haemolymph was thawed and diluted to a final concentration of 0. 1, 1 or 10% in Sterile Octopus Saline (SOS) (NaCl, 2.367 g/100 ml; Glucose, 1 g/100 ml; CaCL, 0.116 g/100 ml; KH2PO;, 0.0056 g/100 ml; KCl, 0.1089 g/100 ml; MgSO4-H20, 0.503 g/100 ml; MgCL, 0.419 g/100 ml). Haemocytes: From each animal 1 ml blood samples were withdrawn into 10 ml ofice cold Marine Anticoagulant (NaCl, 2.63 g / 100 ml; Glucose, 1.8 g/100 mi; Tri- * Sodium Citrate, 0.088 g/ml; Citric Acid, 0.055 g/100 ml) containing ethylene gly- col-bis(b-aminoethylether) N, N, N”, N', - tetraacetic acid (EGTA) (0.029 g/100 ml). After a blood count the haemocytes were centrifuged at 800 g for 5 min at 4*C, and washed by resuspension in Octopus Rin- ger (NaCl, 2.433 g/100 ml; Glucose, 1.4 g/100 ml; EGTA, 0. 015 g/100 ml; KCl, 0.082 g/100 ml; KH2PO),, 0.004 g/100 ml) containing CaCL (0.0142 g/100 ml), MgCl (0.0524 g/100 ml) and MgSO: (0.0629 g/100 ml). A final haemocyte count was made before the haemocytes were washed for a second time and resuspended in SOS at 1 x 10 haemocytes/ ml. Bacteria: Vibrio anguillarum (MT275) were obtained from the Scottish Office, Agriculture and Fisheries Department, Marine Laboratory, Aberdeen. Formali- zed V. anguillarum were counted, was- hed twice by resuspension in SOS and centrifuged at 13000 g for 10 min before resuspension at 8 x 10% cells/ml in the required treatments. Transmission electron microscope (T.E.M.) preparation: Five hundred yl of blood was withdrawn from the bran- chial blood vessel of the octopus and mixed directly with 500 yl of washed bacteria. After 2h incubation at 15*C the blood was centrifuged and the hae- molymph removed. The pelleted hae- mocytes were fixed for 24 h at 4*C in 2.5% glutaraldehyde (in 0.1M sodium cacodylate buffer at pH 7.4). The hae- mocytes were washed in 0.1M sodium cacodylate buffer and secondarily fixed for 2 h at room temperature in 1% osmium tetroxide before staining en bloc with 2% uranyl acetate over night. The pellet was then dehydrated through ethanol and propylene oxide and embedded in Spurr resin. Cut sections (50 nm) were mounted on 100 mesh pio- loform copper coated grids and stained with lead citrate. Sections were viewed in a GEC Corinth 500 at 60 KV. Phagocytosis assay: Two phagocyto- sis experiments were performed to determine the effect of haemolymph concentration, temperature and time on haemocyte phagocytosis. Five animals were used for each experiment. The first experiment involved incubating hae- mocytes in 16 well tissue culture slides (Nunc) for 2 h at different temperatures, but utilizing one pre-incubation tempe- rature and time for the bacteria. The second experiment involved haemocyte incubations of 30 min only and utilized different temperatures, times and hae- molymph concentrations for bacterial pre-incubations. For the first experiment 50 ml of the haemocyte suspension in SOS was put into each of the 16 well chambers of a tissue culture slide. Fifty microliters of either SOS or haemolymph diluted in SOS was added in duplicate, at half hour intervals, to selected wells. Bacte- ria were resuspended in either SOS or 100% haemolymph for 2 h at 15*C and washed twice before use. Fifty microli- . ters of either SOS treated or hae- molymph treated bacteria immediately followed the haemolymph additions, again in duplicate. Each well of the tissue culture slide therefore contained: 50 ml of haemocytes in SOS, 50 ml of either SOS or haemolymph diluted in SOS to 0.1, 1 or 10% concentration (final concentrations of 0.03, 0.33 or 3.33% res- pectively) and 50 ml of bacteria resus- pended in SOS after treatment. The assays were run at four temperatures (5, 10, 15 and 20%C). After 2 h the tissue Iberus, 15 (2), 1997 culture slides were rinsed in SOS to remove unattached bacteria and the slide fixed by immersion in methanol for 3-5 min. The second experiment involved the addition of 50 ml of haemocytes in SOS at 1 x 10% haemocytes/ml, followed by 50 ml of haemolymph diluted in SOS at 0, 0. 1, 1 or 10% concentrations and 50 ml of the different bacterial preparations added in duplicate to the tissue culture slides. The bacteria were washed and resuspended in haemolymph at concen- trations of 0, 0.1, 1, 10 or 100%, using Phosphate Buffered Saline pH 7.0 (PBS, Gibco, without Ca** and Mg?*) as the diluent. Bacteria were incubated for 1, 10, 60 or 120 min at 5, 10, 15 or 20*C, before being washed twice and used in the assay. The slides were incubated at temperatures of 5, 10, 15 or 20*C. After 30 min the tissue slides were rinsed with SOS and the experiment stopped by immersion of the slide in methanol as previously. All slides were then stained in Giemsa (Sigma), rinsed in Gurr Buffer (BDH pH 6.8) and air dried before mounting using DPX. Statistical analysis: Analysis was performed by random counting of 200 haemocytes in each well. The haemocy- tes were counted under oil using a com- pound binocular microscope at 800x magnification. All slides were numbe- red and randomly selected to reduce observer bias. The number of haemocy- tes which had phagocytosed bacteria was expressed as a percentage of the haemocytes counted in each of the duplicate wells. The results for each of the duplicate wells were averaged and analysis of variance (ANOVA) perfor- med for the 2 experiments using the 5 replicates. In each case P values of < 0.05 were taken as being significant. The replicate means were calculated and Tukey's pairwise comparison was per- formed for each experiment using the calculated confidence interval esti- mation (CI estimation). The CI estimate allows 2 separate means to be statisti- cally compared (Rick, 1988). RESULTS Phagocytosis of the formalized Gram negative bacterium, V. anguilla- rum, by E. cirrhosa haemocytes occurs both in the presence and absence of hae- molymph. Collected haemocytes were incubated with bacteria for 2 h before fi- xation for T.E.M. Sections clearly indi- cate that E. cirrhosa haemocytes pha- gocytose and degrade bacteria (Fig. 1). From Analysis of variance a number of significant conclusions were obtai- ned. Phagocytosis by haemocytes follo- wing pre-incubation of the bacteria in 100% haemolymph was significantly greater than phagocytosis following SOS treatment (F= 594.85, P<0.0001) (Fig. 2). Highly significant values were also obtained for the effect of incubation temperature (F= 155.09, P<0.0001), and also for the duration of the assay (F= 178.9, P<0.0001). The concentrations of haemolymph used in the assay medium did not have a significant effect (E= 0.32, P= 0.814) indicating that the rate of pha- gocytosis was statistically equivalent in assays containing 0, 0.1, 1 or 10% hae- molymph. Cross-wise comparisons of the per- centage of haemocytes phagocytosing opsonized and unopsonized bacteria, temperature and assay duration were also highly significant, (P<0.0001), whereas cross-wise comparisons invol- ving haemolymph concentration in the assay medium, confirmed that the hae- molymph concentrations, in SOS, did not affect phagocytic rates. Hae- molymph concentration was therefore not considered in further analysis, and results at each temperature and time were pooled. Phagocytosis of bacteria pre-incuba- ted in SOS was affected by temperature and time (Fig. 2A). At all temperatures the number of haemocytes engulfing bacteria increased over time. At 20*C there appeared to be fewer haemocytes phagocvtosing than at 15%C, however statistically there was no difference between the means at the 2 temperatu- res. At 109C there was a rapid increase in the number of haemocytes phagocy- MALHAM ET AL.: Phagocytosis by haemocytes from Eledone cirrhosa IS TAE Figure 1. Transmission electron micrograph of an Eledone cirrhosa haemocyte (H) having engulfed a bacterium (Vibrio anguillarum) (B). Scale bar 10 pm. Figura 1. Microfotografía de un hemocito (H) de Eledone cirrhosa tras haber tragado una bacteria (Vibrio anguillarum) (B). Escala 10 ym. tosing bacteria during the first 30 min followed by a slower rate of increase up to 2 h. At both 5 and 10*C significantly lower phagocytic rates were observed than at 15 and 20*C over the 2 h period. Fig. 2B shows the mean number of hae- mocytes phagocytosing bacteria, pre- incubated in 100% haemolymph, over time. The haemocyte phagocytic rate again increased over the 2 h period but there were far smaller differences between the incubation temperatures. The phagocytic rates were again lower at 5%C than at the other temperatures. The maximum increase in phagocytosis at all temperatures occurred within the first 30 min. As with the first experiment, the dif- ferent concentrations of haemolymph in SOS (at 0, 0.1, 1 or 10%) used in the se- cond assay were found to have little ef- fect, so were removed from the pair wise comparison with no appreciable percen- tage error increase (0.027%) and the re- sults pooled at each pre-incubation tem- perature and time. To simplify the pair- wise comparison the assay temperature was not included as a main factor, but was added as an interacting factor. The results from the simplified model show that there were large statistically signifi- cant differences (F= 1083.35, P<0.0001) between the haemolymph pre-incuba- tion concentrations. The pre-incubation temperatures (F= 61.32, P<0.0001), and the pre-incubation times (F= 725.24, P<0.0001) were similarly significantly different. Pre-incubation of the bacteria in PBS alone at different temperatures and time periods caused no significant difference in the phagocytic rate (Fig. 3). Bacteria pre-incubated in 0.1% hae- molymph in PBS at all pre-incubation temperatures and times were phagocyto- sed at a significantly lower rate than in PBS alone. Pre-incubation of the bacteria Iberus, 15 (2), 1997 [o] o [>] o = o 2 8 60 ES o S 50 [5] £=, a E 40 p0) 2 E 30 ——5 EY) —— 10 A 15 10 —= 20 0 0 0,5 1 1,5 2 Time (Hours) 90 80 70 D 3 60 E o =) 3 50 - ( E 40 (5 2 (0) a 30 ——5 20 —==— 10 15 10 —= 20 0 0 0,5 1 1,5 2 Time (Hours) Figure 2. A: phagocytosis of non-opsonized formalized Vibrio anguillarum at 4 temperatures over a 2h haemocyte incubation period. The bacteria were pre-treated with SOS for 2h at 15*C. Tukeys CI estimate= 9.52. B: phagocytosis of opsonized formalized Vibrio anguillarum at 4 temperatures over a 2h haemocyte incubation period. The bacteria were pre-treated with 100% haemolymph for 2 h at 15”C. Tukeys CI estimate = 9.52. Figura 2. A: fagocitosis de bacterias formalizadas Vibrio anguillarum ro opsonizadas a cuantro tem- peraturas sobre un periodo de incubación de'hemocitos de dos horas. Las bacterias fueron pretratadas con SOS durante 2 horas a 15*C. Estimación CI de Tukeys= 9,52. B: fagocitosis de Vibrio anguillarum for- malizado y opsonizado a cuatro temperaturas sobre un periodo de incubación de hemocitos de 2 horas. Las bacterias fueron pretratadas con hemolinfa al 100% durante 2 horas a 15*C. Estimación del inter- valo de confianza de Tukeys= 9,52. in 1% haemolymph showed initially the valent to the values determined in PBS same lowered phagocytic rate as for alone. However, at 10 min following pre- 0.1% pre-incubation. However, pre-incu- incubation at 15 and 20%C more hae- bation of the bacteria in 1% haemolymph mocytes were observed phagocytosing for 10 min at 20C caused an enhanced bacteria than at 5 or 10*C, or at 1 min at phagocytic rate which also occurred at all temperatures. The enhanced pha- all temperatures at 60 and 120 min. Bac- gocytic rate observed using 10% hae- teria pre-incubated in 10% haemolymph molymph is statistically equivalent to for 1 min at 5, 10, 15 and 20*C and for 10 the enhanced rate observed at 1% con- min at 5 and 10*C were statistically equi- centration. MALHAM ET AL.: Phagocytosis by haemocytes from Eledone cirrhosa AE Mb | AN pl il | ] | 0.1% Haemolymph 19) (e) MI pl cO aL 39 3315 25 10 ”n o ES 2 (e) IN) AN 40 AA Si N M NN IN] ! 00% | 100% Haemolymph | ECAARAN 20 i ] a Cross Tabulated Mean Percent Phagocytosis 10% Haemolymph Pre-Incubation Time Figure 3. Phagocytosis of formalized Vibrio anguillarum. The haemocytes were incubated at diffe- rent temperatures for 30 min only. The bacteria were pre-incubated in 0% haemolymph (.e., PBS only), 0.1% haemolymph, 1% haemolymph, 10% haemolymph and 100% haemolymph concen- trations. The bacterial pre-incubation temperatures were 5, 10, 15 and 20*C and the pre-incuba- tion times were 1, 10, 60 and 120 min. Tukeys CI estimate = 3.1. Figura 3. Fagocitosis de Vibrio anguillarum formalizado. Los hemocitos fueron incubados a diferentes temperaturas durante sólo 30 minutos. Las bacterias fueron preincubadas en concentraciones de hemo- linfa del 0% (1.e., sólo PBS), 0,1%, 1%, 10% y 100%. Las temperaturas de preincubación de las bac- terias fueron 5, 10, 15 y 20*C y los tiempos de preincubación de 1, 10, 60 y 120 minutos. Estimación del intervalo de confianza de Tukeys= 3,1. DISCUSSION The results presented here demons- trate that E. cirrhosa haemocytes are capable of recognizing and ingesting the formalized bacterium Vibrio anguillarum. V. anguillarum is a Gram negative com- mensal marine opportunist and was chosen as the experimental bacterium because it has been isolated from, and used in previous studies on wound healing in E. cirrhosa (BULLOCK, PoL- GLASE AND PHILLIPS, 1987). This bacte- rium has also been implicated in causing cephalopod infections when the animals are held in captivity and is a common contributory cause of death at high aquarium temperatures (LEIBOVITZ, MEYERS AND ELSTON, 1977; HANLON, FORSYTHE, COOPER, DINUZZO, FOLSE AND KELLY, 1984; FORD, ALEXANDER, COOPER AND HANLON, 1986; HANLON AND FORSYTHE, 1990). STUART (1968) found that E. cirrhosa haemocytes required haemolymph for in vitro phagocytosis of erythrocytes. The data presented in this paper demonstrate that the presence of hae- molymph is not necessary for ingestion of bacteria. However, this bacterium is smaller with far less surface area than an erythrocyte and as such maybe more easily phagocytosed. It was found by TYSON AND JENKIN (1974) that haemocy- tes from a crayfish (Parachaeraps bicarina- tus) phagocytosed bacteria in the absence of haemolymph, but erythrocy- tes were not phagocytosed unless they were pre-treated with haemolymph (McKay, JENKIN AND ROWLEY, 1969). Further JENKIN (1976), suggested that the concentration of certain recognition Iberus, 15 (2), 1997 molecules on the crayfish haemocyte surface was not sufficient to bind eryth- rocytes, but was sufficient to bind bacte- ria, and a similar explanation could apply to E. cirrhosa haemocytes. Another possibility was demonstrated by BAYNE, MOORE, CAREFOOT AND THOMPSON, (1979), who showed that haemocytes from Mytilus californianus had a greater affinity for yeast cells than human erythrocytes, and suggested that pha- gocytosis of foreign particles was selec- tive. Results from other molluscan species also demonstrate that surface antigenicity of the respective test parti- cles has an effect on phagocytosis by haemocytes (TRIPP AND KENT, 1967; ANDERSON AND GOOD, 1976). Tripp (1966), using the bivalve M. mercenaria, concluded that haemolymph pre-treatment of erythrocytes caused increased phagocytosis. The same expe- riment showed however that if untrea- ted erythrocytes were incubated with haemocytes for longer periods of time, the same levels of phagocytosis were achieved. With E. cirrhosa haemocytes at 15 and 20*—C the phagocytic rate is higher at 30 min for 100% haemolymph treated bacteria compared to SOS treated bacteria, but after 2 h there was no difference in phagocytic rates between the 2 treatments. The data pre- sented here also indicate that a higher percentage of haemocytes phagocytosed haemolymph treated bacteria at 5 and 10*C over 2 h than SOS treated bacteria. Tripp (1992) also showed that the hae- mocytes of M. mercenaria were avidly phagocytic in the absence of hae- molymph, however at low temperatu- res, in the presence of haemolymph there was increased phagocytosis of yeast. ABDUL-SALAM AND MICHELSON (1980), working with Biomphalaria gla- brata, also demonstrated that tempera- ture has an effect on haemocyte pha- gocytosis. A phagocytic activity peak was evident at 30C with inhibition of phagocytosis below 15%C. Low tempera- ture inhibition (4%C) of phagocytic rates has also been demonstrated for the hae- mocytes from the hard clam M. mercena- ria with maximum rates occurring at 22 and 37%C (FOLEY AND CHENG, 1975). With SOS treated bacteria, E. cirrhosa haemocytes demonstrate an activity peak with about 70% of haemocytes phagocytosing after 2 h at 15 and 20*C. At 5*C only 14% of haemocytes contai- ned bacteria, whereas if the bacteria were initially pre-incubated in hae- molymph before addition to the assay the phagocytic rate at 5%C increased to around 47%. The results presented above indicate that the amount of haemolymph present in the bacterial pre-incubation medium has a dramatic effect on the number of haemocytes subsequently engulfing these bacteria within a 30 min period. Haemolymph concentrations of 0.1 and 1% in PBS, resulted in lower numbers of haemocytes phagocytosing compared to PBS alone. This inhibition changes to enhanced phagocytosis, at all higher pre-incubation concentrations. Further comparisons demonstrate that the tem- perature of the pre-incubation medium and particularly the duration of incuba- tion are also important factors. The ob- served trends indicate that increasing the pre-incubation temperature decrea- ses the pre-incubation time needed for enhanced phagocytosis to occur. FRYER ET AL. (1989), working on B. glabrata, si- milarly demonstrated that phagocytosis was inhibited after short pre-incubation periods, whereas longer pre-incubation periods of 1 h resulted in enhanced le- vels. It was suggested by the authors that initial non-specific adsorption of a variety of plasma components (opso- nins) occurred onto, in their case, the ye- ast surface. Longer exposure to the plasma allowed more of the opsonins to bind to the yeast surface. The results from the data presented here for the dif- ferent pre-incubation haemolymph con- centrations and durations of exposure seem to support this hypothesis. In ad- dition it is possible that if the tempera- ture is increased further more of the available plasma components would ad- here onto the surface of the bacterium. When haemocytes from E. cirrhosa were resuspended in SOS, as stated above, there is phagocytosis of the for- MALHAM E7 AL.: Phagocytosis by haemocytes from Eledone cirrhosa malized bacterium V. anguillarum. In buffers containing either EDTA or EGTA, no phagocytosis of the same bac- terium was evident (Malham, unpublis- hed data). SOS contains Ca?”* and Mg?* and it appears likely that the presence of these divalent ions has an effect on pha- gocytosis. FRYER AND ADEMA (1993) sho- wed that manipulated haemocytes from B. glabrata retained some phagocytic ac- tivity, but that addition of excess Ca?* and Mg?* to the haemocytes before the addition of the target particles enhanced their phagocytic rates. E. cirrhosa hae- mocytes were initially drawn into an an- ticoagulant buffer containing EGTA and washed in Octopus Ringer, also contai- ning EGTA, before resuspension in EGTA-free-SOS, all of which could alter haemocyte behaviour and affect pha- gocytosis. Corbicula fluminea haemocytes (TUAN ET AL., 1987) also required extra- cellular Ca?* or Mg?* for both opsonin- dependent and independent phagocyto- sis. The authors suggest that the opso- nin possibly exists as a divalent cation-macromolecular complex due to the loss of enhanced phagocytosis after dialysis against EDTA and EGTA. Furt- her, Mytilus edulis haemocytes phagocy- tosed yeast cells with high efficiency when calcium ions were present in the suspension medium, and gave similar results when haemolymph alone was added, but almost no phagocytosis was recorded with haemocytes in buffered saline (RENWRANTZ AND STAHMER, 1983). When V. anguillarum was resus- pended in SOS, E. cirrhosa haemolymph diluted in SOS, or in PBS alone, there was no change in the haemocyte pha- gocytic rate. However, when V. anguilla- rum was resuspended in haemolymph diluted in PBS (2 1% haemolymph con- centration) or in haemolymph alone, en- hanced phagocytosis was observed. REFERENCES ABDUL-SALAM, J. M. AND MICHELSON, E. H., 1980. Biomphalaria glabrata amoebocytes: as- say of factors influencing in vitro phagocyto- sis. Journal of Invertebrate Pathology, 36: 52-59. Haemolymph lectins have been shown to act as opsonins for haemocyte phagocytosis (e. g., RENWRANTZ, 1983; RENWRANTZ 1986; SMINIA AND VAN DER KNAPr, 1986, VasTA, 1991). Aggluti- nation results from Octopus maya (Fis- HER AND DINUZZO, 1991) further sup- port the role of lectins in recognition of non-self. Studies using the molluscs My- tilus edulis (RENWRANTZ AND STAHMER, 1983) and Lymnaea stagnalis (VAN DER KnNAPr, 1982) have demonstrated that molecules antigenically related to hae- molymph lectins have been found in the cytoplasm and on the surface of hae- mocytes. Lectins, in particular C-type, are found in a number of invertebrates including Octopus vulgaris. These lectins are Ca?* dependent, and these ions are required for ligand binding of the lectin (RÓGENER, RENWRANTZ AND UHLEN- BRUCK, 1986). STUART (1968) suggested a possible link between an opsonic factor and haemocyanin in E. cirrhosa. Also a lectin identified from the haemolymph of O. vulgaris has been shown to be simi- lar to a haemocyanin subunit (ROGENER, RENWRANTZ AND UHLENBRUCK, 1985). The nature of the soluble factor causing enhanced phagocytosis in E. cirrhosa has not been studied, however the factor(s) must be present at a high concentration, since it is effective at a haemolymph concentration of 1% at 15 and 20*C. In conclusion, in vitro phagocytosis of Vibrio anguillarum by haemocytes from E. cirrhosa is aided by a component of haemolymph and is affected by tem- perature, duration of the assay and pre- incubation of the bacterium with diffe- rent haemolymph concentrations. Fur- ther studies to elucidate whether E. cirr- hosa haemocytes are capable of pha- gocytosing and digesting live microor- ganisms in vitro and in vivo are being pursued. ANDERSON, R. S. AND GOOD, R. A,, 1976. Op- sonic involvement in phagocytosis by mo- llusk hemocytes. Journal of Invertebrate Pat- hology, 27: 57-64. 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Lectins in molluscs and arthropods: their occurrence, origin and ro- les in inmmunity. Zoological Society of London, 56: 81-93. MALHAM E7 AL.: Phagocytosis by haemocytes from Eledone cirrhosa RENWRANTZ, L. AND STAHMER, A., 1983. Op- sonizing properties of an isolated hemolymph agglutinin and demonstration of lectin-like recognition molecules at the surface of he- mocytes from Mytilus edulis. Journal of Com- parative Physiology, 149: 535-546. RICE, J. A., 1988. Mathematical Statistics and Data Analysis. Wadsworth and Brook, California, 595 pp- RÓGENER, W., RENWRANTZ, L. AND UHLEN- BRUCK, G., 1985. Isolation and charaterization of a lectin from the hemolymph of the cep- halopod Octopus vulgaris (Lam.) inhibited by a-D-lactose and N-acetyl-lactosamine. Deve- lopmental and Comparative Immunology, 9: 605- 616. RÓGENER, W., RENWRANTZ, L. AND UHLEN- BRUCK, G., 1986. Comparison of a hemolymph lectin from Octopus vulgaris with hemocya- nin. 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Journal of Invertebrate Pathology, 50: 310-319 TYsON, C. J. AND JENKIN, C. R., 1974. Phagocy- tosis of bacteria in vitro by haemocytes from the crayfish (Parachaeraps bicarinatus). Aus- tralian Journal of Experimental Biology and Me- dical Science, 52: 915-923. VAN DER KNAPpP, W. P. W., 1982. The internal defence system of the pond snail Lymnaea stagnalis. PhD Thesis, Vrije Universiteit te Amsterdam, The Netherlands, 139 pp. VASTA, G. R., 1991. The multiple biological ro- les of invertebrate lectins: their participation in nonself recognition mechanisms. In: Warr, G. W. and Cohen, N. (Eds.): Phylogenesis of Im- mune Functions. CRC Press Inc., New York: 74-102. WELLs, M. J., 1978. Octopus: Physiology and be- haviour ofan advanced invertebrate. Chapman and Hall, London, 478 pp. WELLs, M. J., 1983. Circulation in Cephalopods. In Saleuddin, A. S. M. and Wilbur, K. M. (Eds.): The Mollusca. Vol 5. Academic Press, London: 239-290. WELLs, M. J]., AND WELLs, J. 1993. Fluid uptake and the maintenance of blood volume in Oc- topus. Journal of Experimental Biology, 175: 211- 218. IM EN A 04 0% b e SN Wi ida JA LZ » AAA Eo k y de MA Y 0 4 bas AD El ve e FELÓ ñ á "Elo | $ ys + E ma EN E des AN mu 1 AO | e A e EN O O ) a A Mo pue a AN PS Y ES O A ol Ada Un «roll mel me vos AN ASA TO, li 108 8 lr Cort), o peros Mk Mireia p A poseo psd pai e dE ud hd pi99 A y Hot dit UN DS Ae ] O Sociedad Española de Malacología ——__—_—_—_—_—_——— Iberus, 15 (2): 13-24, 1997 New data on the morphology and the distribution of Bul;- mulus corneus Sowerby, 1833 (Gastropoda: Pulmonata: Orthalicidae) in Nicaragua Nuevos datos sobre la morfología y la distribución de Bulimulus cor- neus Sowerby, 1833 (Gastropoda: Pulmonata: Orthalicidae) en Nica- ragua Antonio Mijail PÉREZ*! and Adolfo LÓPEZ* Recibido el 8-[-1996. Aceptado el 22-IV-1996 ABSTRACT Aspects related to the morphology and distribution of Bulimulus corneus Sowerby, 1833 in Nicaragua are presented. Regarding morphology, a complete redescription of the shell and the first description of the genitalia are included. The number of records have been largelly increased; from three localities mentioned in the literature to 53. The previous figu- res have allowed us to draw a preliminary distribution map of the species in Nicaragua, and discuss the presence of the closely related species Bulimulus unicolor Sowerby, 1833 in the country. RESUMEN Se presentan aspectos relacionados con la morfología y la distribución de Bulimulus cor- neus Sowerby, 1833 en Nicaragua. En relación con la morfología, se presenta una redes- cripción de la concha y la primera descripción del aparato genital. El número de registros de la especie en el país ha sido notablemente incrementado de tres a 53 localidades. Las cifras anteriores nos han permitido confeccionar un mapa preliminar de distribución para la especie en Nicaragua, así como discutir la presencia de Bulimulus unicolor Sowerby, 1833, una especie muy relacionada, en el país. KEY WORDS: Bulimulus corneus, Orthalicidae, morphology, distribution, Nicaragua. PALABRAS CLAVE: Bulimulus corneus, Orthalicidae, morfología, distribución, Nicaragua. INTRODUCTION According to BREURE (1979), the distribution of Bulimulus corneus genus Bulimulus Leach, 1814 contains 88 Sowerby, 1833 as from SW Mexico to the species, distributed over the Antilles, central zone of Costa Rica, there being Central America and northern South apparently no records outside of these America. MARTENS (1890-1901) gave the limits. In Nicaragua, previous reports * Universidad Centroamericana, Apartado A-90, Managua, Nicaragua. ' Dirección postal temporal: Universidad del País Vasco, Facultad de Ciencias, Departamento de Biología Animal y Genética, Laboratorio de Zoología, Apartado 644, 48080 Bilbao, España. 13 Iberus, 15 (2), 1997 IE DMAX Figure 1. Bulimulus corneus. Shell measurements. Abbreviations. LONG: length; DMAX: maximum diameter; LEC: height of the body whorl; DME: minimum diameter; LAB: aperture length; AAB: aperture width; LEP: spire length; AEP: spire width. Figura 1. Bulimulus corneus. Medidas de la concha. Abreviaturas. LONG: longitud; DMAX: diámetro máximo; LEC: altura de la vuelta principal; DME: diámetro mínimo; LAB: altura de la abertura; AAB: anchura de la abertura; LEP: altura de la espira; AEP: anchura de la espira. have been from Realejo (Chinandega), San Juan Castillo (sic), El Toro rapids, RAAN (Autonomous Region of the North Atlantic) (MARTENS, 1890-1901) and Bluefields (FLUCK, 1900). Martens stated that this species is closely related to Bulimulus unicolor So- werby, 1833, and this was confirmed by PiLSBRY (1897). None of the authors re- cognized TATE's (1870) reports of B. uni- color from Granada, Mesapa and San Ni- colas, on the Pacific slope of Nicaragua. The internal and external morpho- logy of B. corneus, shell measurements and data on distribution were recently presented for the first time in an abstrac- ted version (PÉREZ AND LóÓPEz, 1995), and are here given in detail. New distri- bution data gathered in the last few months are also presented, together with a commentary on the presence of B. unicolor in Nicaragua. 14 MATERIALS AND METHODS All specimens were hand-collected and live specimens were relaxed in menthol and fixed in 70% alcohol. All individuals considered for the study were fully- grown adult specimens. All localities reported are additions to those previously mentioned in the literature. The list of localities is gi- ven in Table 1. The distribution map was made using the UTM cartographic met- hod with a grid size of 100 Km?. When more than one locality occur on the same UTM 10 Km? quadrat, only the one that appears first in the listis mapped. The ab- breviations w. l. n. and Bib. means wit- hout lot number and bibliographic loca- lity respectively. The variables measured in the shells are (Fig. 1): 1. length (LONG) 2. maximum diameter (DMAX,) 3. height of body whorl (LEC), 4. minimum diameter (DME), 5. PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua aperture length (LAB), 6. aperture width (AAB), 7. spire length (LEP), 8. spire width (AEP). All measurements were made in adult specimens. We calculated various descriptive statistics for the measured variables, in or- der to give a morphological description of the samples. We also used a Principal Component Analysis (PCA) to explore the variability among populations. RESULTS AND DISCUSSION Description: Shell (Fig. 2): Shell thin, spirally striate, corneus to brown, so- mewhat translucent, showing through the dark bands that stipple the mantle. Pro- file bulimoid-conic. Apex obtuse; proto- conch typically bulimoid with sculpture of punctures in an irregular decussate pat- tern; whorls 5.5 to 6. Aperture ovate, mar- gin thin, sharp, umbilicus narrow. Mea- surements taken on the shell are presen- ted in Tables I and II. Genitalia (Fig. 3): Penis with wide she- ath, dilated in its central part, and reaching to more than one half of the phallus. Epip- hallus approximately half as wide as pe- nis. Flagellum thinner, approximately one half the phallus length. Sperm conduct thickened at mid-center, ending at globose spermatheca distally. Vagina more or less fusiform, slightly longer than the penis, and ?/3 the width. We have found that shell dimensions are quite variable within (Table I) and bet- ween populations (Table II), as also men- tioned by PILSBRY (1897). For this reason, and because of the small total sample size (n= 44) studied from all populations (11) we have not considered the taxonomical implications of the variability. However, it should be mentioned that shell length (LONG) and height of body whorl (LEC) display the highest variances of all varia- bles considered (Table II). Shell length is always one of the variables on which des- criptions are based. We recommend cau- tion in the use of either variables for a ta- xonomic characterization of the species. It must be pointed out that THOMPSON (1967) invalidated various subspecies of the closely-related Bulimulus unicolor So- werby, 1833, believing them to be varia- tions related to climatic conditions. In this paper he considered shell length (LONG), maximum diameter (DMAX) and two ot- her variables. In the PCA made from conchological variables it is possible to see the marked scatter of the specimens (Fig. 4). Within the plot, there is a segregation of six indi- viduals from the populations of El Gua- yabo, Granada (1), Xiloá, Managua (2), and Las Lajas, Rivas (3). It is interesting to notice that the other specimen from Ri- vas is located within the cloud of points. The only specimen considered from the Nicaraguan Atlantic slope (6), can be ob- served between the cloud of points and the six individuals previously mentioned. Another three specimens from Ocotal (9) segregate towards the lower right cor- ner of the scatterplot. These specimens, as the previous six, have conchological fea- tures very much like the ones from other populations (see Table 1), although the ones from Ocotal have larger sizes. In Table III, it can be seen the contri- bution made by each principal compo- nent to total variance. Components I (70.77%) and Il (18.15%), comprise the major quantity of total variance (88.92%). The absence of negative signs (Table IV) among the eigen values obtained for component l, also with the larger contri- bution (70.77%), allow us to presume that it is related to size and Il is related to shape. Thus, differences among popula- tions would be apparently due to size rather than shape; and it is known that size is usually influenced by ecological factors, and consequently is highly corre- lated with local environmental condi- tions (BEROVIDES, 1988). In their genitalia (see Figure 3), the individuals from the populations of UCA Campus (Managua Department) and Ocotal (Nueva Segovias Depart- ment) share the same external morpho- logy and show only small differences in size of the structures. However, a more detailed anatomical analysis, including an analysis of the internal anatomy of the genital ducts (v. g. penis), of the Ocotal population is required when fresh material is available. Iberus, 15 (2), 1997 16 PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua Figure 3. Bulimulus corneus. Genitalia. A: specimen from Ocotal; B: specimen from Campus UCA; pr: prostate; p: penis; f: flagellum; a: atrio; (sp: spermatheca)= bc: bursa copulatrix; s. ov: spermo- viduct; sp. d: spermathecal duct; ep: epiphallus; v: vagina. Scale bars 1 mm. Figura 3. Bulimulus corneus. Aparato genital. A: ejemplar de Ocotal; B: ejemplar del Campus UCA; pr: próstata; p: pene; f: flagelo; a: atrio; (sp: espermateca)= bc: bolsa copulatriz; s. 0v: espermoviducto; sp. d: conducto espermático; ep: epifalo; v: vagina. Escalas 1 mm. Distribution: Fifty three localities for B. corneus have been added to those previous recorded. They distributed over 12 Departments in the three natural regions that comprise the country (Fig. 5, Table V). CONCLUSIONS The distribution map gives a clear idea of B. corneus distribution in Nicara- gua. As pointed out by JACOBSON (1968), a fairly continuos distribution can be seen among the samples, suggesting that absence in other areas is due to lack of sampling, and that B. corneus is wides- pread in the country. B. corneus has a very wide ecological tolerance, ocurring from low altitude to more than 2000 m. The species inhabits a remarkable number of different microha- bitats, including soil with herbs, soil with litter, tree trunks, logs, stones, walls of ruined houses, etc. The wide geograp- hical distribution of the species, can pro- (Left page). Figure 2. Bulimulus corneus. Shell morphology. A: Las Canoas (length 12.8 mm, dia- meter 7.0 mm); B: Campus UCA (length 10.6 mm, diameter 6.7 mm), C: Ocotal (length 19.9 mm, diameter 11.35 mm). (Página izquierda). Figura 2. Bulimulus corneus. Morfología de la concha. A: Las Canoas (1 longitud 12,8 mm, diámetro 7,0 mm); B: Campus UCA (longitud 10,6 mm, diámetro 6,7 mm); C: Ocotal (longitud 19,9 mm, diámetro 11,35 mm). 17 Iberus, 15 (2), 1997 Table I. Variables measured considering each sample separately (X= average, S= standard deviation). Abbreviations as in Figure 1. Tabla I. Variables medidas considerando los ejemplares de cada muestra independientemente (X= media, S= desviación standard). Abreviaturas como en la Figura 1. LOCALITIES VARIABLES LONG DMAX LEC DME LAB AAB LEP AEP Xiloá [n= 6) XK 10.93 6.70 A 6.13 SS] 3.87 4.38 4.85 Mín 10.10 5.80 DES) 5.30 Doll 2.95 2.9 ARS Máx 11.70 O 8.6 6.5 6.0 4.7 5.8 5.4 S 0.62 0.61 137 0.46 0.33 0.63 1.29 0.35 Apoyo (n= 10) X 11.62 8.19 8.82 6.31 DL EZ 3.6 4.49 Mín 10.0 6.3 7.8 DS) 4.6 2.4 2.4 37 Máx 13.5 8.0 10.0 YES 6.0 3.9 4.75 DS S 1.04 052 0.76 0.48 0.42 0.42 0.64 0.45 Asososca (n= 4) X 1237 a 9.27 7.0 DS 3.6 3.82 4.9 Mín 12.0 6.5 8.4 DA 45 ZE 399 4.3 Máx 13% 7.8 9.6 7.1 7 3.8 4.1 5.0 S 0.59 0.17 0.42 0.1 0.35 0.21 0.41 0.16 Las Canoas [n= 5) XK 18202 SIS 9.46 6.6 9 3.62 4.33 AED Mín 1285 6.8 9.0 6.3 AZ SZ 4.2 4.7 Máx 13.8 DL, 10.0 EZ 5.8 4.3 4.6 DS S 052 057 0.42 0.35 0.22 0.42 0.17 0.26 El Guayabo (n= 5) XK 10.65 DS DIS 5.95 IDO 4.85 IS 5.05 Mín 10.1 6.0 DS 5.8 Del 4.8 4.6 4.7 Máx 1912 6.3 6.0 6.1 6.0 4.9 6.0 5.4 S 0.78 0.2 035 0.22 0.64 0.22 0.99 0.5 Las Lajas (n= 2) X 14.25 YES 6.82 6.7 6.35 ASIS SY 5.8 Mín 12.9 6.5 6.3 6.0 Dl 3.8 EZ DES) Máx 1576 8.1 SS 7.4 7.0 4.5 00 6.1 S 1:91 1.18 0.74 0.99 0.92 0.5 0.18 0.65 Tepeyac (n= 2) X 11.25 Del 8.75 5.85 4.8 SD) SS) 45 Mín 11.0 6.7 8.6 0) 4.7 2 SS) 4.4 Máx 155 YES 8.9 6.7 4.9 3.9 3.4 4.6 S 0.35 0.42 0.21 1162 0.14 0.5 (0) 0.14 Ocotal (n= 4) XK 1712 9.52 237 8.72 ES, 4.92 5.40 6.16 Mín 13.6 8.0 10.5 8.0 7.4 4.2 3.8 ZO Máx 19:9 11S5 11212 10.0 8.4 EZ 6.9 AZ S ZO, AO) 1.54 0.87 0.48 0.5 1.34 0.80 Campus UCA (n= 3) XK 11.56 AZ 8.78 (3 5.03 3.46 SSZ 4.47 Mín 10.6 6.7 8.3 6.0 4.9 3.0 2.8 4.1 Máx 12.8 7.8 9.55 2 5. 1 3.8 3.8 4.9 S 1.12 0.55 0.63 0.64 0.12 0.42 0.50 0.41 Laurel Galán (n= 3) XK 12.45 7.6 8.95 6.7 6.02 IEA 7 4.6 Mín 15144 AO 8.35 6.0 4.9 3.0 2.8 4.1 Máx 18389 8.1 9.5 YES 6.35 3.6 4.0 5.0 S 0.98 0.55 05% 0.65 0.45 0.12 0.29 0.46 18 PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua Table IL. Variables measured considering all samples pooled. (X= average; Min: minimum value; Max: maximum value; S= standard deviation). Abbreviations as in Figure 1. Tabla 11. Variables medidas considerando los ejemplares de todas las muestras agrupadas. (X= media; Min: valor mínimo; Max: valor máximo; S= desviación standard). Abreviaturas como en la Figura 1. Variables Xx Min Max S LONG 12.40 10 19.90 2.02 DMAX JESZ Date INESS 0.99 LEC 8.78 4.1 14.20 1.89 DME 6.59 DO) 10.00 0.89 LAB 5.64 4.5 8.40 0.87 AAB SINO) DIA DENO) 0.67 LEP AZ ZA 6.90 0.97 AEP 4.86 37 7.20 0.64 Axis ll Axis | Figure 4. Axis I and II of the Principal Component Analysis. Each number represents a sample. 1: El Guayabo; 2: Xiloá; 3: Las Lajas; 4: Asososca; 5: Campus UCA; 6: Loma del Mico; 7: Laurel galán; 8: Tepeyac; 9: Ocotal; 10: Apoyo; 11: La Ceiba. Figura 4. Ejes 1 y 11 del Análisis de Componentes Principales. Los números corresponden a las muestras. 1: El Guayabo; 2: Xiloá; 3: Las Lajas; 4: Asososca; 5: Campus UCA; 6: Loma del Mico; 7: Laurel galán; 8: Tepeyac; 9: Ocotal; 10: Apoyo; 11: La Ceiba. 19 Iberus, 15 (2), 1997 Table III. Percentage of variance explained by each one of the principal components. Abbreviations as in Figure 1. Tabla III. Porcentaje de varianza explicado por cada uno de los componentes principales. Abreviaturas como en la Figura 1. Component Percent of cumulative number Variance percentage LONG 70.77 ANOTA DMAX SMS 88.92 LEC 4.72 93.64 DME EDS, 95.88 LAB 2.06 SISI AAB 0.89 98.84 LEP 0.71 SEDO 100.00 AEP 0.44 Table IV. Eigen values obtained with Principal Component Analysis, considering axes L, Il and III. Abbreviarions as in Figure 1. Tabla IV. Valores propios obtenidos con el Análisis de Componentes Principales, considerando los ejes l, 11 y 1. Abreviaturas como en la Figura 1. Components LONG 03993 Variables | DMAX 0.3726 LEC 0.3072 DME 0.3872 LAB 0.3902 AAB 0.2860 LEP 0.2861 Íl 111 0.1351 0.2987 0.3116 -0.0867 -0.4996 0.0703 -0.2181 -0.2486 0.0308 -0.0825 0.4631 0.7271 O3S339 0.4999 0.2948 0.2258 AEP 0.3756 bably be explained by the numerous mi- crohabitats that it is capable of filling. Considering at the same time the morphological variability and the ecolo- gical range of this species, WOLDA's (1970) stament comes to mind, that va- riation should be understood in terms of the possibilities of survival in natural po- pulations, and not only as a biologically isolated fact about the ecology of species. Regarding the presence of B. unicolor . in Nicaragua, Tate's reports were not re- cognized by MARTENS (1890-1901), or by PiLsBRY (1897), and may have had their 20 origin in the marked variability of B. cor- neus. We think that only B. corneus is found in Nicaragua. PiLsBRY (1897), mentioned Greytown (RAAN: Autonomous Region of the North Atlantic), and later Fluck (1900) gave Bluefields (RAAS: Autonomous Re- gion of the South Atlantic) as localities for B. unicolor in Nicaragua. More re- cently, BREURE (1979) quoted Perico Is- land in the Bay of Panamá as the only lo- cality in Central America. In the last two years we have collec- ted specimens from Bluefields, one of PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua 160 130 Figure 5. Bulimulus corneus. Distribution in Nicaragua, in UT'M notation of 100 Km?. Figura 5. Bulimulus corneus. Distribucion en Nicaragua, en notación UTM de 100 Knr. the localities for B. unicolor, and two ot- her nearby localities (Las Delicias and La Fonseca) (see Table IV). This material agrees well with the description of B. corneus. We are therefore led to think either that in these localities both species live sympatrically, and we have not so far co- llected B. unicolor, or that B. unicolor does not occur there at all. In view of the fact that in the revision of the subfamily BREURE (1979) did not include distribution data for B. unicolor in Nicaragua, and taking into account the morphological variability that THOMPSON (1967) has shown in both B. unicolor, and B. corneus, as mentioned by Pilsbry and also studied by us, we have decided for the moment to accept that the latter species is the only one that oc- curs in Nicaragua. ACKNOWLEDGEMENTS We are grateful to Dra. Ana Isabel Puente (University of Basque Country, Faculty of Sciences, Department of Animal Biology and Genetics, P. O. Box 644, 48080 Bilbao) for critical revision and advice about the paper. Also to Lic. Zamira Guevara (Central America Uni- versity, Department of Ecology and Natural Resources, P. O. Box. 90, Managua, Nicaragua) who collected and processed part of the studied material with us. The great encouragement to our work, given by Ing. Luis Fiallos, Dean of the Faculty of Agricultural Sciences, is also to be mentioned and gratefully acknowledged. The sugges- tions made by two anonymous revie- wers contributed to largely improve the manuscript's scientific quality. 21 Iberus, 15 (2), 1997 Table V. List of new localities for B. corneus Sw., in Nicaragua. S= number of shells, Sa= specimens in alcohol. The samples used for statistical purposes are marked with an asterisk. Bibliographic loca- lities are considered as Bib under “Lot number”. w. l. n. means: “without lot number”. Tabla V. Lista de nuevas localidades para B. corneus Sw., en Nicaragua. S= número de conchas, Sa= especímenes en alcohol. Las muestras usadas con propósitos estadísticos aparecen marcadas con con un aste- risco. Las localidades bibliográficas aparecen con la abreviatura Bib en “Lot number”; w. l. n. significa: “sin número de lote”. Code Lot Localities number Coordinates Geographical BOACO Department 1 92:19 "El Sácal CARAZO Department 2 90:18 La Baronesa CHINANDEGA Department 3 Bib Realejo CHONTALES Department 4 92:14 Punta Mayal 5 88:30 Nueva Guinea ESTELI Department 6 93:09 Estanzuela GRANADA Department 7) 91:26 Tepeyac 8 92:01 Aguas Calientes, Cocibolca 9 w.Ln Isleta de Ken 10. 92:12 Laguna Blanca 11. 95:72 El Guayabo 12 ER933lO Apoyo LEON Department 13 90:05 Asososca 14 90:06 Laguna Monte, Galán 15 88:26 Salinas Grandes MANAGUA Department 16 88:21 Villa Carmen 17 88:28 Las Sierritas 18 90:07 Xiloá 19 SES Asososca ZOO Las Mercedes, Lago Xolotlán ZII S SY Apoyeque 22 94:47 San Francisco Libre ZION Los Placeres, Km 63 24 95:04 Km 66.8 Carr. Matagalpa 12*33'10” N, 85"33'30" W 12%10'N, 8717 W 123927" N, 83918” W 11%52'20" N, 85*26' W 11%43' N, 84*57' W 13%14'04” N, 862216" W 11%52.5' N, 85"59.5' W 11%52' N, 85"55'40" W 11%51'41” N, 85"53'40" W 11%46'15 N, 85%57'45" W 1158" N, 8559 W 1155 N, 85%57'45" W 12%26' N, 86"40' W 12%26' N, 86"40' W 12%16'12” N, 86"30'4” W 1219 N, 8616' W 123 N, 8616' W 12%14' N, 86"46' W 121811” N, 85198" W 12%9'30" N, 8610' W 12%15' N, 8621 W 12*30'12” N, 86%17'40" W 12*33' N, 86"3'41” W 12%29'06" N, 86%04'17" W DD UTM 16PFJ 58 16PFJ 72 16PDJ 78 16PFJ 71 16PGH 79 16PEK 73 16PFJ 01 16PFJ 11 16PFJ 11 16PFJ 10 16PFJ 11 16PFJ 01 16PEJ 37 16PE) 47 16PEJ 25 16PEJ 74 16PEJ 73 16PE) 74 16PEJ 74 16PEJ 84 16PEJ 74 16PEJ 78 16PFJ 08 16PFJ 08 Material examined 6S 125 25 28 25(*) 25 25 4S Ss(*) 105+55a As") 155 6S 10S+4Sa 6S ós(*] 15 25 9S 15 5Sa 1Sa PÉREZ AND LÓPEZ: Morphology and distribution of Bulimulus corneus in Nicaragua Table V. Continuation. * Tabla V. Continuación. Code Lot Localities number 25 95:06 Km42.5 Carr. Matagalpa 26 95:07 Las Canoas LOSA Las Canoas 28 95:26 Mateare ZII Carr. Tipitapa-Masaya, Km 43 SOI La Polvosa 31 95:28 Sierra de San Andrés SS: S Nandayosi 33 95:32 Nandayosi, cerca del rio 34 95:34 LosFilos de Guajachillo SIA 38 San Bartolo SOI El Conchital 37 95:40 Hacienda “El Apante” 38 95:44 Hacienda “El Callao” 39 95:45 Samaria AOS El Tamarindo 41 95:59 El Platanal 42 95:64 Carr.Sur, Km 15.5, INCAE ASI 72 Campus UCA MATAGALPA Department 44 92:83 Ciudad Darío AS 2:88 Fuentepura NUEVA SEGOVIAS Department d6 94:59 Ocotal RIVAS Department AT Z3 9 Río Las Lajas RIO SAN JUAN Department 48 92:24 La Toboba 49 92:25 El Castillo 50 w.ln San Carlos 51 92:26 Laurel Galán Coordinates Geographical 12%21'02” N, 8602'58" W 12%19'00” N, 80%00'07” W 121905" N, 855922" W 121410" N, 86%25'48" W 120948” N, 86%00'07" W 121324” N, 86"24'59” W 12%10'28” N, 86*24'35"” W 12%06'43” N, 86"31'14” W 12%06'54” N, 86"30'16" W 12%08'33" N, 86"24'33 W 11%54'58' N, 86"33'05" W 11%54'18” N, 86"33'43" W 11%57'48” N, 862915" W 12%01'42” N, 8620'07" W MESAUSAN, IRSA A 12%29'37" N, 86"05'01” W 12%27'06" N, 86%05'02” W 10%03'13” N, 86"18'33" W 12%07'30" N, 861613" W 12%43'50" N, 861153" W 1258" N, 8655" W 13%36'35" N, 862818” W 112 N, 85 W 11%8' N, 84*57'50" W 11720” N, 8424 W 12720" N, 86"46' W 12%7' N, 86"46' W RAAS (AUTONOMOUS REGION OF THE SOUTH ATLANTIC) 52 93:23C Las Delicias 53 93:23B La Fonseca 54 94:19 Loma del Mico 55 Bib Bluefields 12%16'12” N, 8352/50" W 12%15'36" N, 83"58'48" W 12%4'24” N, 83%47'42” W 12% N, 834418" W RAAN (AUTONOMOUS REGION OF THE NORTH ATLANTIC) 56 Bib San Juan Castillo 14%24' N, 8354'54" W UTM 16PFJ 06 16PFJ 16 16PFJ 06 16PEJ 65 16PEJ 65 16PEJ 65 16PEJ 64 16PEJ 53 16PEJ 53 16PEJ 42 16PEJ 41 16PEJ 41 16PEJ 52 16PEJ 72 16PEJ 42 16PEJ 98 16PEJ 97 16PEJ 73 16PEJ 74 16PEK 90 16PFK 13 16PEL 51 16PFH 35 16PGH 72 16PGH 81 16PGH 43 16PGH 43 17PJP 17PJP 17PJP 17PJP 17PJR Material examined 7S IS 155 108 108 118 325 35 135 ÁS 7S 155 35 145 28 39 15S 1S 3Sa(*) 135 28 55") 251") 351") 23 Iberus, 15 (2), 1997 REFERENCES BEROVIDES, V., 1988. Orden y diversidad en el mundo viviente. Editorial Cintífico- Técnica, La Habana. 108 pp. BREURE, A.S. H., 1979. Systematics, phylogeny and zoogeography of Bulimulinae. Zoologis- che Verhandelingen, 168: 1-205. FLuck, W. H., 1900. Shell collecting in the Mos- quito Coast. The Nautilus, 14: 94. JACOBSON, M. K., 1968. On a collection of te- rrestrial mollusks from Nicaragua. Nautilus, 81: 114-120. MARTENS, E. V., 1890-1901. Biologia Centrali- Americana. Land and Freshwater Mollusca. Tay- lor and Francis, London. 706 pp. PÉREZ, A. M. AND LÓFPEZ, A., 1995. New data on the morphology and the distribution of Bu- limulus corneus Sowerby, 1833 (Gastropoda: Pulmonata: Orthalicidae). In Guerra, A., Ro- lán, E. and Rocha, F. (Eds.): Abstracts of the Twelfth International Malacological Congress, Vigo. Unitas Malacologica and Instituto de In- vestigaciones Marinas, Vigo: 396-398. 24 PiLsBRY, H. A., 1897. Manual of Conchology. 2nd series, vol. 11, pp. 65-144. TATE, R., 1870. On the land and freshwater mo- llusca of Nicaragua. American Journal of Con- chology, 5: 151-162. THOMPSON, F. G., 1967. The land and freshwa- ter snails of Campeche. Bulletin Florida State Museum, 11(4): 221-256. WOoLDA, H., 1970. Ecological variation and its implications for the dynamics of populations of the landsnail Cepaea nemoralis. Proc. Adv. Study Inst. Dynamics Numbers Popul.: 98-108. O Sociedad Española de Malacología —————— Iberus, 15 (2): 25-34, 1997 Phenological patterns and life history tactics of Helicoidea (Gastropoda, Pulmonata) snails from Northern Greece Patrones fenológicos y estrategias de vida en Helicoidea (Gastropoda, Pulmonata) del Norte de Grecia Maria LAZARIDOU-DIMITRIADOU and Stefanos SGARDELIS* Recibido el 8-[-1996. Aceptado el 31-VIL1996 ABSTRACT The present study mainly concerns with the differences in the biological cycles and strategies adopted by different Helicoidea snail species. In Northern Greece the climatic conditions are not very uniform. Some snails breed during the same period as in Northern Europe but most breed in autumn, as species from Southern Europe do. Breeding may take place in all sea- sons except in winter, and seems to be species-specific. Long-lived snails of big size differ from shortlived species of small size as to the time of their breeding period. The climatic con- ditions affect the time of the breeding season and their whole life cycle and phenologies. En- vironmental variables in Northern Greece are strongly seasonal and thus Helicoidea snails exhibit predictable oscillations in their activity patterns, which can be interpreted by the de- mographic response of the populations. Terrestrial snails seem to follow two different pheno- logic curve types: the semelparous and short-lived species populations show a more stable phenological pattern than the biennial and pluriennial ones, who mature after the first year of their lives, being more plastic trying to face the climatic differences from one year to another. RESUMEN El presente estudio trata de las diferencias en los ciclos biológicos y a las estrategias adopta- das por diferentes especies de Helicoidea. En el norte de Grecia, las condiciones climáticas no son muy uniformes. Algunas especies crían durante el mismo periodo en que lo hacen en el N de Europa, pero la mayoría lo hacen en otoño, como sucede en especies del S del con- tinente. La cría puede tener lugar durante casi todas las estaciones, excepto el invierno, y el periodo parece ser específico para cada especie. Las especies longevas y de gran talla di- fieren de las de pequeño tamaño y vida más corta en la duración de su ciclo de cría. Las condiciones climáticas afectan al momento de la temporada de cría y a todo su ciclo vital y fenología. Las variables ambientales son fuertemente estacionales, así que aparecen osci- laciones predecibles en los patrones de actividad, que pueden ser interpretadas por la res- puesta demográfica de las poblaciones. Las babosas parecen seguir dos tipos de curvas fe- nológicas distintas. Las especies semelpáricas y de corta vida muestran un patrón fenológico más estable que el de especies bianuales y prurianuales, que maduran tras el primer año de vida y son más flexibles a la hora de enfrentarse a las diferencias climáticas interanvales. KEY WORDS: Biological cycle, phenology, Northern Greece, Helicoidea, Helix, Eobania, Helicella, Monacha, Bradybaena. PALABRAS CLAVE: ciclo biológico, fenología, N Grecia, Helicoidea, Helix, Eobania, Helicella, Monacha, Bradybaena. * Departments of Zoology and Ecology, School of Biology, Aristotle University of Thessaloniki, 54006 Thessaloniki, Macedonia, Greece. 25 Iberus, 15 (2), 1997 INTRODUCTION Greece has a Mediterranean climate which is differentiated mainly along a northern-southern gradient. In Northern Greece climate is transient from Mediterranean (mostly coastal areas) to temperate (inland areas). A typical cha- racteristic of this climate type is the coincidence of high temperatures and low precipitations during summer (las- ting from June to October). The wet sea- son is divided by a cold winter which is milder in the coastal areas. Drought is a strong agent controlling population dy- namics and activity of most soil inverte- brates as it imposes a pause in most physiological activities. Low temperatu- res during winter are also important for population dynamics and activity as they impose hibernation in some of the invertebrates e. g. terrestrial gastropods. So observed discontinuities in popula- tion development during the transition from the favourable to the unfavourable seasons and vice-versa may be attribu- ted to environmental thresholds. Although the association between climate and life history phenomena is self evident, it can vary among terres- trial molluscs, even between popula- tions of the same species. Phenology reflects certain aspects of the demo- graphy of a population, that is the timing of its life cycle characteristics in a given environment. The classification of phenological patterns into categories or types (WOLDA, 1988) is better by using phenological models (VAN STRAALEN, 1982; STAMOU, ASIKIDIS, ARGYROPOULOU AND SGARDELIS, 1993). Using a phenolo- gical model, complex phenograms can be classified into types considering their skewness, curtosis, phase and period. The aim of the present study was to find out whether terrestrial gastropods adopt a general phenological pattern if * they are differentiated according to their origin, or their biotopes (inland and coastal areas) or life spans. The present study is a part of an extensive research done on the distribution and ecology of Helicoidea gastropods in Northern Greece (LAZARIDOU-DIMITRIADOU, 1981, 26 1995; LAZARIDOU-DIMITRIADOU AND KATTOULAS, 1981, 1985, 1991; STAIKOU, LAZARIDOU-DIMITRIADOU AND FARMA- KIS, 1988; HATZUOANNOU, ELEUTHERIA- DIS AND LAZARIDOU-DIMITRIADOU, 1989; STAIKOU, LAZARIDOU-DIMITRIADOU AND PANA, 1990; STAIKOU AND LAZARIDOU- DIMITRIADOU, 1990, 1991). MATERIALS AND METHODS Data used derived from monthly quantitative samplings of Helicoidea snails from different parts of Northern Greece. The following species were studied: Family Bradybaenidae, Brady- baena fruticum (Múller, 1774); Family Helicidae, Cepaea vindobonensis (Férus- sac, 1821), Eobania vermiculata (Muller, 1774), Helix lucorum Linnaeus, 1758, Helicella (Xerothracia) pappi (Schút, 1962), Helix figulina (Rossmássler, 1839), Helix pomatia rhodopensis Kobelt, 1906, Theba pisana (Múller, 1774); Family Hygromii- dae, Cernuella virgata (Da Costa, 1778), Monacha cartusiana (Múller, 1774), Xero- lenta obvia (Menke, 1828), Xeropicta arenosa Ziegler, 1836, Xerotricha conspur- cata (Draparnaud, 1801). Sampling lasted two or four years depending on the species and their life span (LAZARI- DOU-DIMITRIADOU, 1981, 1995; LAZARI- DOU-DIMITRIADOU AND KATTOULAS, 1981, 1985, 1991; STAIKOU ET AL., 1988; STAIKOU ET AL., 1990; STAIKOU AND LAZARIDOU-DIMITRIADOU, 1990, 1991). Details regarding the sites and the sam- pling procedures are given in previous studies on these species (LAZARIDOU- DIMITRIADOU, 1981, 1995; LAZARIDOU- DIMITRIADOU AND KATTOULAS, 1985, 1991; STAIKOU ET AL., 1988). Ombrother- mic data for Northern Greece from 1980 to 1990 are given in Figure 1. Data were provided by Mahairas P., Professor of Climatology from the Aristotle Univer- sity of Thessaloniki. Fischer's exact test for independence in 2 x 2 contingency tables (ZAR, 1984) was used for comparisons between the different categories, e. g. snails with autumnal and vernal-estival reproduc- tive periods, long-lived (> 3 years) and LAZARIDOU-DIMITRIADOU AND SGARDELIS: Phenology of Helicoidea in N Greece JF MA M Precipitation (mm) Prec. (mm) A SON D Months Figure 1. Ombrothermic diagram from Northern Greece. Temperature: open squares; Precipitation: solid rhombus. Figura 1. Diagrama ombrotérmico del Norte de Grecia. Temperatura: cuadrados abiertos; Precipitación: rombos sólidos. short-lived snail species (up to 3 years), large (largest shell diameter > 25 mm) and small sized snails. The phenological pattern of Helicoi- dea species was studied by using the phenological model applied for the study of microarthropods (STaAMOU ET AL., 1993). In short, in this model when asymmetries and discontinuities are dis- played the scales of the time-axis were adjusted. Changing time scales results in the definition of a new variable termed ecological time (ET), which is a function of a standard clock time (STAMOU ET AL., 1993). This technique is based upon the following considera- tions: 1) the timing of a population in the field is determined by the sequence of demographic events and/or beha- vioural characteristics (i. e. migratory), and 2) the rate of the demographic events depends on the fluctuations of environmental variables. In this model it is assumed that the phenology of any population inhabiting a seasonal environment can be descri- bed by a symmetric periodic curve: F(ET) = EXP(a-+b x COS(27 x (ET-9/T)) (1) where the independent variable ET, termed Ecological time, is a function of standard clock time (ST), ET = f (ST). In the course of standard time, ecological time is going faster during periods of sharp changes in abundance and slower during periods of abundance stability. Thus, the proposed equation describing the length of the Ecological time unit (AET) as a function of Standard time ST is: AET= f (ST) = EXP(or+b1 x COS(27 x (ST-p1)/T1)) (2) * The model has eight parameters of which the period T and the phase q of the phenogram, as well as period T1 and the phase q1 of the function relating ET to ST, are the most important. The period T and the phase q of the pheno- grams are expressed formally in ET units. For convenience they could be expressed in ST units (as T' and q”) by using equation (2) for the calculation of 27 Iberus, 15 (2), 1997 the Standard time T” or q” which corres- ponds to T or € units in the Ecological time-scale (see STAMOU ET AL., 1993: fig. 1). For the comparison of phenograms two more parameters can be derived: a) an estimation of the sharpness (curtosis) of the phenogram C=(R2-R1)/T”, where (R2-R1) is the time interval around the phase q”, during which the abundance is above overall mean, and b) an esti- mation of the skewness of the pheno- gram S= (0'-Qm)/T' where qmis the time (in ST units) when the abundance of the population is at minimum. Thus, pheno- grams displaying a peak at p'-qm= T'/2 (half period) is symmetric (S= 0.5), phe- nograms displaying a peak soon after the minimum (S< 0.5) are positively skewed, and phenograms with S> 0.5 are negatively skewed. The model was fitted on log-transformed census data. For ETi= f (STi) given as a time vector and for a given set of q and T, the para- meters a and b were estimated by least- square regression. RESULTS The model fitted to census data for snail populations sampled at monthly intervals from different areas are shown in Figure 2. The values of the most important parameters of the fitted phe- nological model are given in Table 1. In all but two examined cases the phenological pattern was strongly sea- sonal (Fig. 2), with a more or less 1 year periodicity apart from Monacha cartu- siana which seems to display a six months periodicity at least during 1984 (Table I). Abundance of Bradybaena fruti- cum and Eobania vermiculata fluctuated almost randomly throughout the year. Population densities of the different species do not exhibit phase synchroni- zation. Even different populations of the same species do not always exhibit a peak density at the same time of the year. Furthermore even the same popu- lation displays a phase instability during successive years of study. For instance Xerolenta obvia from Paleokas- tro peaked either in January or in April 28 (q, in Table I). X. obvia from Karvali peaked in July. Helix lucorum displayed a similar interannual instability. In both cases the shift in phase seems to be asso- ciated with an unexpected change of the weather, an extended dryness (STAIKOU ET AL., 1988: fig. 2) which provoked an overall decline of the population density (Fig. 2). The phase expressed as months after minimum population densities (qm) (Table I) is a measure of the rate of population increase from absolute minimum to peak densities. Qm might have lower values when the time needed for a species to mature is short. Minimum (Qm values were estimated for X. obvia in Paleokastro, the 1st and the third year of study, indicating a rapid population growth which occurs in a period of about 3 months (Fig. 2). In Karvali, where X. obvia gets mature in 2 years, Qm value was larger (Table 1), as was the case with M. cartusiana in 1985. Helicella (Xerothracia) pappi snails that need 2-3 years to mature exhibited larger Qm values, and H. lucorum snails - which need 3-4 years exhibited the largest value of all except in 1983. Species that mature in one year, exhibit a very rapid population growth just after the adverse period, which is winter, and consequently they are posi- tively skewed. Species that mature in 2- 3 years are negatively skewed (Table Il, Fig. 3). In this case the phenological pattern may be positively skewed or symmetrical but the population remains active for longer periods and the pheno- logical pattern is platycurtic (Table II, Fig. 3). Whereas species that mature in one year usually display a leptocurtic phenology (Table Il, Fig. 3). The only species population that showed both a negatively and a positively skewed phe- nological pattern was H. lucorum. The positively skewed phenograms are leptocurtic when they concern snail species populations that mature in one year whereas they may be slightly platy- curtic when they live in regions where favourable conditions last longer as is the case of Xeropicta arenosa Ziegler in Edessa. Pluriennial snail species popula- LAZARIDOU-DIMITRIADOU AND SGARDELIS: Phenology of Helicoidea in N Greece 10 ”,* Helix lucorum 1) Edessa to . Population density (ind./m2) JFMAMJJASONDJFMAMJJASONDJFMAMJJASONDJFM 83 84 14 A a o Monacha cartusiana 42 o E A AN Edessa E 10 / Ñ p N = (o) ¡MORO NS Dos 7 Ñ Y 3 : a / A ; Y 5 Wa 4 ES sS. U ó e ás NS e 5 ó > a) D (e) o A S a o a 5] 2 [aY 0 AA] MJJASONDJ]FMAMJJASONDJFMAMJIJJ 6 Xeropicta arenosa Edessa Population density (ind./m2) JEMAMJJASONDJIJFMAMJJASONDJFMAMJJASONDIJFMA 86 83 84 Months 21 ] Xerolenta obvia os Paleokastro ¿$ Lo 1 40 " Á Mn ¿ LO y X | : o | 204 |! eN l No E so S DA i je] Ea] ES PS ES A , o HAL ONDJFMAMJJASONDJFMAMJJASONDJFMAMIJAS 88 180 160 o Xerolenta obvia 140 / V . / MN Karvali 120 / EN ' Va 10) P y 80 i X . í ye dl 60 la A > 90 A , , » 20 Ñ JÓ $ le O AAXA>->>—- -á ->>--- 91 mM y y AÑES OON D ñ F M A M y J A S 89 90 30 Helicella (Xerothracia) pappi IN Philippi MAMJJASONDJFMAMJJASONDJFMAMJJASONDJ 86 Months Figure 2. Abundance variations of Helicoidea snails from Northern Greece. Line: model estimates; asterisks: observations. Figura 2. Variaciones de la abundancia de caracoles helícidos del Norte de Grecia. Línea: estimaciones del modelo; asteriscos: observaciones. tions usually exhibit platycurtic pheno- logical patterns, too, since their adults diapause and hide in the soil or under plants and do not emerge massively. Negatively skewed and symmetric phe- nograms are usually platycurtic. It seems that there are no negatively skewed-leptocurtic species except for M. cartusiana that is negatively skewed and slightly leptocurtic. Maximum activity duration is about the same for different populations of the same species or for the same population during successive years of study (Table [, MA column). In Northern Greece long-lived species are of big size and short-lived are of small size (x?= 9.983, P= 0.001). Additionally, short-lived species breed in autumn whereas long-lived species may breed in autumn or vernal-estival period (x?= 4.261, P= 0.039) (Table III). In Northern Greece bigger helicids may breed during the vernal-estival or Iberus, 15 (2), 1997 Table I. The estimated parameters of the fitted model in standard time units (months). Tabla 1. Parámetros estimados del modelo ajustado en unidades de tiempo standard (meses). Phase ej Phase (qm Maximum Species Place Year E (Months after (Months after activity period R? Das ps January) minimum dens!y) MA (Monthts) dpi Helicella pappi Philippi 1987 12.6 2.9 5.9 7.1 OIOSIEZS 1988 10.5 3.4 6.8 6.9 0.95 Xerolenta obvia Karvali 1990 12.6 6.7 4.7 4.4 0.79 2 Paleokastro 1988 14.2 0.3 29 42 0.84 ] 1989 10.8 3.5 30) 7.3 0.77 1990 10.4 0.2 2.8 2.9 0.91 Xeropicta arenosa Potidea 1979 12.7 6.1 )) 5.1 0.87 | 1980 11.2 6.2 38) 6.9 0.88 Edessa 1984 12,5 6.6 4.9 6.7 0.69 ] 1985 11223 7.8 5.8 6.9 0.68 Monacha cartusiana Edessa 1984 6.5 2.4 37 3.2 0.53 l 1985 13.0 -1.1 6.2 4.3 0.45 2 Helix lucorum 1983 12.5 4.] 5.0 8.1 0.94 34 1984 12.0 8.4 7.9 8.5 0.84 1985 12.0 8.9 8.5 8.2 0.86 Table IL. Skewness (negatively skewed <0.5; positively skewed >0.5) and curtosis (leptocurtic <0.5; platycurtic >0.5) from the phenograms of Helicoidea snails from Northern Greece). Abbreviations: Phil: Philippi; Pal: Paleokastro area; Kar: Karvali; Pot: Potidea; E: Edessa (the paranthesis means slightly). Tabla 11. Desviación (desviado negativamente <0, 5; desviado positivamente >0,5) y curtosis (leptocúrtico <0,5; platicúrico >0,5) en los fenogramas de caracoles helícidos del Norte de Grecia. Abreviaturas: Phil: Philippi; Pal: Paleokastro area; Kar: Karvali; Pot: Potidea; E: Edessa (los paréntesis significan ligeramente). Species Place and Year deta o Symmetric Leptocurtic Platycurtic Mesocurtic ada Helicella pappi Phil. 1987 (+) + 23 1988 + > Xerolenta obvia Paleo. 1988 + . 1 1989 (+) + 1990 + + Karv. 1990 + + (+) 2 Xeropicta arenosa Potid. 1979 + (+) ] 1980 + + Edes. 1984 + + l 1985 + + Monacha cartusiana Edes. 1983 + (+) 2 1984 + + 1 1985 + (+) + 2 Helix lucorum Edes. 1983 + + 3-4 1984 + + 1985 + autumnal period and small ones breed of the reproductive period is very short mainly in autumn (x= 4.261, P= 0.039) whereas in the inland areas it is variable (Table III). In coastal areas the duration according to the species. 30 LAZARIDOU-DIMITRIADOU AND SGARDELIS: Phenology of Helicoidea in N Greece ps mM Right O.4 Xik] xapl Ss 80 Platy- hi2 a hpf2 a ni3 Ml 60 a Left Skewness Al 4ES mc3 a 30 Kurtosis 20 Lepto- Figure 3. Ordination of phenograms into skewness(S)- curtosis (C) plane (Skewness: negatively ske- wed< 0.5; positively skewed> 0.5. Curtosis: leptocurtic< 0.5; platycurtic> 0.5) of Helicoidea snails from Northern Greece. Numbers denote 1st, 2d or 3d generation. Abbreviations, xap: Xeropicta are- nosa Potidea; xae: Xeropicta arenosa Edessa; hl: Helix lucorum; hpf: Helicella pappi Philippi; mc: Monacha cartusiana; xip: Xerolenta obvia Paleokastro. Figura 3. Ordenación de fenogramas respecto al plano desviación (S)- curtosis (C) (Desviación: negati- va< 0,5; positiva> 0,5. Curtosis: leptocúrtico< 0,5; platicúrtico> 0,5) de babosas de la familia Helicoidea del Norte de Grecia. Los número denotan las primera, segunda y tercera generaciones. Abreviaturas, xap: Xeropicta arenosa Potidea; xae: Xeropicta arenosa Edessa; hl: Helix lucorum; hpf Helicella pappi Philippi; mec: Monacha cartusiana; x1p: Xerolenta obvia Paleokastro. DISCUSSION In Northern Greece the climatic con- ditions are not uniform (HATZIOANNOU ET AL., 1989). The ombrothermic diagram for Northern Greece (Fig. 1) from 1980 to 1990 shows that the dry season is from June to October, whilst the wet season is divided by a cold winter period. Breeding may take place almost in all seasons except during winter. In Northern Greece, snails mainly breed from April to the end of autumn. The strong seasonality of the climate imposes a seasonal pattern of breeding. Consequently, there are two main breeding periods: the autumnal breeding period starts with the first rainfalls and stops with low temperatu- res (LAZARIDOU-DIMITRIADOU, 1981; STAIKOU AND LAZARIDOU-DIMITRIADOU, 1991) and the vernal-estival breeding period which starts when the mean monthly temperature rises over 10%C and stops when the arid period starts (LAZARIDOU-DIMITRIADOU, 1981; LAZA- RIDOU-DIMITRIADOU AND KATTOULAS, 1985; STAIKOU ET AL., 1988). Most of the land snails though, mainly short-lived and small snails, breed during the autumnal period (Table III) as Helicoi- dea species from Southern Europe do (CHATFIELD, 1968; REAL AND REAL- TESsTUD, 1983; HELLER, 1982; SACCHI, 31 Iberus, 15 (2), 1997 Table II. Life cycle characteristics from Helicoidea snails from Northern Greece. Abbreviations: D: largest shell diameter. Ehinos is found in Rhodope area, Edessa and Thessaloniki in North Central Macedonia, Philippi and Karvali near Kavala, and Paleokastro and Potidea in Chalkidiki. Abbreviations, Y. years up to maturity; SL: short lived < 3 years; LL: long lived > 3 years; SS: small sized D < 25 mm; LS: large sized D > 25 mm; V: vernal-estival reproductive period; A: automnal reproductive period. Tabla III. Características del ciclo de vida de los caracoles helícidos del Norte de Grecia. Abreviaturas: D: mayor diámetro de la concha. Ehinos se encuentra en el área de Rhodope, Edessa y Thessaloniki al Norte de Macedonia, Phillipi y Karvali cerca de Kavala, y Paleokastro y Potidea en Chalkidiki. Abreviaturas, Y: años hasta la madurez; SL: vida corta < 3 años; LL: vida larga > 3 años; SS: pequeña talla D < 25 mm; LS: gran talla D > 25 mm, V: periodo reproductivo estival; A: periodo reproductivo otoñal. Species Locality Longitude Helix pomatia rhodopensis Ehinos 249 58' 34" Helix lucorum Edessa DINA Monacha cartusiana (rarely ) Bradybaena fruticum Edessa Cepaea vindobonensis Edessa Helix figulina Thessaloniki 22% 57' 29” Theba pisana Thessaloniki Xerotricha conspurcata Thessaloniki Eobania vermiculata Thessaloniki Helicella (Xerothracia) papi Philippi 249 15' 48” Xerolenta obvia Paleokastro MUDA Karvali 2423011 Xeropicta arenosa Potidea DNA Edessa IAS Cernuella virgata Potidea ESSE Latitude Y SL LL SS LS A 419 16' 50” 3 + + 409 47 47" 3-4 + + 2 + + + 1 + + 2 + + 2 + + 419 24' 26” 2 + + + 2 + + + | + + + 2 + + + 419 7'26' 2-3 + + + 409 24' 59" | + + + 409 59 44” 2 + + + 4091124” 1 + + + 407 47' 47” 1 + + + 401124" | + + + 1971; BONAVITA AND BONAVITA, 1962; DEBLOCK AND HOESTLANDT, 1967) and only some breed during the vernal period as land snails from Northern Europe do (POLARD, 1975; WOLDA AND KREULEN, 1973). M. cartusiana, which is of Northern origin, breeds in both seasons (STAIKOU AND LAZARIDOU-DIMI- TRIADOU, 1990). Estival breeding period happens in places with a wet climate during summer months (STAIKOU ET AL,, 1988). In areas where different species coexist (as in Logos area in Edessa) alt- hough the climatic conditions are the same the species do not breed during the same period provoking less antago- nistic intraspecific reactions to their hat- chings (STAIKOU ET AL., 1988). Semelpa- rous species with an r-strategy synchro- nize their breeding period with the favourable period which is October-mid 32 November (LAZARIDOU-DIMITRIADOU, 1981, 1995; LAZARIDOU-DIMITRIADOU AND KATTOULAS, 1985). There is also a marked difference between the snail species living along the sea shore and the inland ones. Cou- pling and laying of eggs is more or less synchronous for the populations living along the seashore and do not last more than a week each. On the contrary bree- ding lasts about a month for the inland species (LAZARIDOU-DIMITRIADOU, 1981; STAIKOU AND LAZARIDOU-DIMITRIADOU, 1991). The climatic conditions under which the land molluscs live do not only affect the time of the breeding season but also their whole life cycle and phenologies. X. obvia needs two years to mature in coastal or semi-coastal areas and one year on the mountains (e. g. Paleokastro, LAZARIDOU-DIMITRIADOU AND SGARDELIS: Phenology of Helicoidea in N Greece Central Chalkidiki, unpublished data). Similarly, M. cartusiana which is of nort- hern origin needs two years to mature in the south instead of one, because it has to face the summer aestivation, alt- hough 15% of its population in Edessa tends to be semelparous (STAIKOU AND LAZARIDOU-DIMITRIADOU, 1990), as in Northern Europe (CHATFIELD, 1968). Species like B. fruticum and E. vermi- culata which are iteroparous and univol- tine species, living for a few years, exhi- bit no seasonal trend of abundance but fluctuate almost randomly during the year. These show rather stable patterns despite the oscillations of environmental parameters. All the rest of the studied Helicoidea species can encounter seaso- nality by adjusting the timing of their vi- tal activities. They respond to the ad- verse period of the year, which is winter time for Northern Greece, by displaying asymmetric (positively or negatively skewed phenograms) seasonal patterns of abundance variation. These patterns reflect a differential response of the spe- cies to tolerance against stress which se- ems not to be region-specific but species- specific. The annual or biennial r-strate- gists (X. arenosa, X. obvia) show a positively skewed phenological pattern due to the rapid development of juveni- les soon after the adverse winter period. These juveniles have been hatched in au- tumn but stayed buried in the soil du- ring winter. Their growth stops before summer dryness during which the geni- talia and the gonad maturation take place and the snails are ready to lay eggs before their death in autumn. On the contrary, populations of pluriennial spe- cies are characterized by low abundance during the onset of the adverse period and a negatively skewed or symmetric phenciogical pattern. It seems that the growth rate of juveniles is much slower than that of annual species. The only species population that showed both a negatively and a positively skewed phe- nological pattern was H. lucorum. Howe- ver, a positively skewed pattern in 1983, that is a rapid growth of population den- sity was probable just after the adverse period since a massive emergence from hibernation of adult snails took place be- cause of good weather conditions which started earlier than usually (STAIKOU ET AL., 1988: fig. 2). Additionally, this spe- cies has a low net reproductive rate (Ro = 0.9) and a low annual turnover ratio (P/B = 1.24), the snails live up to 12 ye- ars and they mature after the third year of their lives (STAIKOU ET AL., 1988). Negatively skewed and leptocurtic phenograms could not be characteristic of a snail species since it would mean that this population would grow slowly and steadily during the favourable pe- riod of the year and would introduce ra- pidly growing immature snails just be- fore the adverse period. This would be possible only if immature snails were more resistant and tolerant to the stress. Such a phenology has not also been re- corded in acari or collembola (SrTamou ET AL., 1993; SGARDELIS, SARKAR, ASIKI- DIS, CANCELA DA FONCECA AND STAMOU, 1993). However, M. cartustana is a case of negatively skewed and slightly lepto- curtic phenology. In this population 15% of juveniles mature in one year as it happens in Northern Europe. So, in 1984 the dry season (summer time) was inte- rrupted by a wet period (STAIKOU ET AL,, 1988: fig. 2) and this 15% of juveniles, which had already matured, managed to lay eggs before the majority of the snails which were mature and ready to copulate and lay eggs in autumn. This population, though, comes from Nort- hern Europe and it is found in the sout- hern limits of its distribution. To sum up, environmental variables in Northern Greece are strongly seaso- nal and thus Helicoidea snails exhibit predictable oscillations in their activity patterns, which can be interpreted by the demographic response of the popu- lations as it has been found with soil microarthropods (STAMOU ET AL., 1993; SGARDELIS ET AL., 1993). The semelpa- rous and short-lived snail species popu- lations show a more stable phenological pattern than the biennial and plurien- nial ones, who mature after the first year of their life, and they are more plastic trying to face the climatic differences from one year to the other. 33 Iberus, 15 (2), 1997 REFERENCES BONAVITA, A. AND BONAVITA, D., 1962. Con- tribution a létude écologique d'Euparypha pisana Múller des rivages Mediterranéens de la Provence. Note preliminaire. Pubblicazioni della Stazione Zoologica di Napoli, 32 (suppl.): 189-204. CHATFIELD, J. E., 1968. The life history of the he- licid snail Monacha cantiana (Montagu), with reference also to M. cartusiana (Muller). Pro- ceedings of the Malacological Society of London, 38: 233-245. DEBLOCK, R. AND HOESTLANDT, H., 1967. Don- nées biologiques sur le Gastéropode littoral Theba pisana Múller aux limites septentrio- nales de son extension. Compte rendu de 1” Academie des Sciences, Paris, 265: 893-896. HATZHIOANNOU, M., ELEUTHERIADIS, Ñ. AND LAZARIDOU-DIMITRIADOU, M., 1989. Contri- bution a la reproduction de l'escargot co- mestible Helix pomatia rhodopensis Kobelt (1906) vivant dans la Grece du Nord-Est. Ha- liotis, 19: 137-141. HELLER, J., 1982. Natural history of Theba pisana in Israel (Pulmonata: Helicidae). Journal of Zoology, 196: 475-487. LAZARIDOU-DIMITRIADOU, M., 1981. Contribu- tion a l'étude biologique et écologique des es- cargots Cernuella virgata (Da Costa) et Xero- picta arenosa Ziegler (Gastropoda, Pulmo- nata) vivant sur les microdunes de Potidea, Chalkidiki (Grece du Nord). Atti del V” Con- vegno della Societa Malacologica Italiana, Pa- via: 73-83. LAZARIDOU-DIMITRIADOU, M., 1995. The life cy- cle, demographic analysis, growth and se- condary production of the snail Helicella (Xe- rothracia) pappi Schútt (1962) (Gastropoda: Pulmonata) in E. Macedonia (Greece). Mala- cologia, 37: 1-11. LAZARIDOU-DIMITRIADOU, M. AND KATTOU- LAS, M., 1981. Contribution a l'étude de la biologie et de la croissance des escargots commercialisés en Grece: Eobania vermicu- lata (Múller) et Helix aspersa Múller. Haliotis, dEN29 137: LAZARIDOU-DIMITRIADOU, M. AND KATTOULAS, M., 1985. Contribution a l'étude biologique et écologique d'Eobania vermiculata (Gasté- ropode, Helicidae). Biologica Gallo-Hellenica, 10: 132-137. LAZARIDOU-DIMITRIADOU, M. AND KATTOULAS, M., 1991. Energy flux in a natural population of the land snail Eobania vermiculata (Múller) (Gastropoda Pulmonata Stylommatophora) in Greece. Canadian Journal of Zoology, 69: 881-891. POLARD, E., 1975. Aspects of the ecology of He- lix pomatia L. Journal of Animal Ecology, 44: 305-329. 34 REAL, G. AND REAL-TESTUD, A. M., 1983. Re- partition en France d'Helicella itala (L., 1758) (Gasteropode, Pulmone Terrestre). Haliotis, 13: 155-161. SACCHI, C. F., 1971. Ecologie comparée des Gas- téropodes Pulmonés des dunes Mediterran- nées et Atlantiques. Natura, 62: 277-358. SGARDELIS, S. P., SARKAR, S., ASIKIDIS, M. D., CANCELA DA FONCECA, J. P. AND STAMOU, G. P., 1993. Phenological patterns of soil mi- croarthropods from three climate regions. European Journal of Soil Biology, 29 (2): 49-57. STAIKOU, A. AND LAZARIDOU-DIMITRIADOU, M., 1990. Aspects of the life cycle populations dy- namics, growth and secondary production of the snail Monacha cartusiana (Muller, 1774) (Gastropoda: Pulmonata) in northern Greece. Malacologia, 31 (2): 353-362. STAIKOU, A. AND LAZARIDOU-DIMITRIADOU, M., 1991. The life cycle, population dynamics, growth and secondary production of the snail Xeropicta arenosa Ziegler (Gastropoda: Pulmonata) in northern Greece. Zoological Journal of Linnean Society, 101: 179-188. STAIKOU, A., LAZARIDOU-DIMITRIADOU, M. AND FARMAKIS, N., 1988. Aspects of the life cycle, population dynamics, growth and secondary production of the edible snail Helix lucorum Linnaeus, 1758 (Gastropoda, Pulmonata) in Greece. Journal of Molluscan Studies, 54: 139- 155% STAIKOU A., LAZARIDOU-DIMITRIADOU, M. AND PAna, E., 1990. The life cycle, population dy- namics, growth and secondary production of the snail Bradybaena fruticum (Muller, 1774) (Gastropoda Pulmonata) in northern Gre- ece. Journal of Molluscan Studies, 56: 137-146. STAMOU, G. P., AsikiDIS M. D., ARGYROPOU- LOU, M. AND SGARDELIS, S. P., 1993. Ecologi- cal time versus standard clock time: the asym- metry of phenologies and the life history strategies of some soil arthropods from Me- diterranean ecosystems. Oikos, 66: 27-35. VAN STRAALEN, N. M., 1982. Demographic analysis of arthropod populations using 2 continuous stage-variable. Journal of Animal Ecology, 51: 769-783. WoLDA, H., 1988. Insect seasonality: Why? An- nual Revue of Ecological Systems, 19: 1-18. WOLDA, H. AND KREULEN, D. A., 1973. Ecology of some experimental populations of the land snail Cepaea nemoralis (L.). IL. Production and survival of eggs and juveniles. Netherlands Journal of Zoology, 23 (2): 168-188. ZAR, J. H., 1984. Biostatistical Analysis. Pren- tice-Hall International Inc., United King- dom, 718 pp. O Sociedad Española de Malacología Iberus, 15 (2): 35-50, 1997 Fragmented knowledge on West-European and Iberian Caudofoveata and Solenogastres Conocimiento fragmentado de los Solenogastros y Caudofoveados de Europa occidental y Península Ibérica Luitfried von SALVINI-PLAWEN* Recibido el 8-1-1996. Aceptado el 4-X-1996 ABSTRACT A basic problem in our knowledge of the aplacophoran molluscs, viz. the Caudofoveata and the Solenogastres, is the poor availability of faunistic samplings. This lacunarity even concerns the European waters; in the present contribution, particular attention is paid to the gap in the records along the French and Iberian shelf regions. This is underlined by presenting an updated geographic distribution of eight caudofoveate and thirteen soleno- gastre species. Benthos investigators are called upon to focus more intensively on sam- pling the smaller marine fauna from mobile bottoms of the West-European shelf regions. RESUMEN Un problema esencial para el conocimiento de los Caudofoveados y Solenogastros (moluscos aplacóforos) es la insignificante disponibilidad de material recogido en diferen- tes muestreos faunísticos. Esta carencia todavía afecta al Atlántico europeo y particular- mente concierne a la falta de muestras en la plataforma continental de Francia y de la Península Ibérica. Esta situación se pone en evidencia con la recopilación actualizada de la distribución geográfica de ocho especies de Caudofoveados y trece de Solenogastros. Se hace una invitación especial a los investigadores del bentos para que intensifiquen su atención por la pequeña fauna marina de sustratos blandos en la plataforma occidental europea. KEY WORDS: Caudofoveata, Solenogastres, Aplacophora, new records, distribution, Europe. PALABRAS CLAVE: Caudofoveados, Solenogastros, Aplacophora, nuevas citas, distribución, Europa. INTRODUCTION This contribution is restricted to a very simple, but momentous problem: the dearth of faunistic information with all its consequences in both classes of aplacophorous molluscs, the Caudofo- veata (formerly Chaetodermomorpha) and the Solenogastres (formerly Neome- niomorpha). Members of the Caudofoveata and the Solenogastres live predominantly in marine offshore habitats below 50 meters depth and are in general not really rare members of benthic biotopes (see SAL- VINI-PLAWEN 1990). The Caudofoveata (average size 2-15 mm) are micro-omni- vores burrowing within muddy sedi- * Institut fúr Zoologie, Universitat Wien, Althanstraffe 14, A-1090 Wien IX, Austria. 99 Iberus, 15 (2), 1997 ments, whereas the Cnidaria-vorous So- lenogastres (average size 2-20 mm) are bound to clay, secondary hard bottoms or enidarian colonies. Our scarce know- ledge about species diversity, biology and zoogeography of the representatives of both groups is in part due to the intri- cate, high-effort and expensive sampling methods for benthic meiofauna (ships, cable winches, benthic sledge-dredges). Due to these technical and financial diffi- culties, investigation of marine meio- fauna is generally restricted to the “home turf” of marine biological stations or fis- hery institutes as typified by Ply- mouth/UK or Naples/Italy. For more than a century, this has resulted in an un- balanced biogeographic and systematic knowledge of small fauna, even in Euro- pean waters, restricting, the informative data predominantly to animals of the North and Mediterranean Seas. Surpri- singly, there are only poor records from the shelf region off France and the Iberian peninsula (delimited in Figure 1 by the 200 m isobath). Therefore, a special invitation is ad- dressed to all Spanish, Portuguese, and French colleagues who perform benthic offshore investigations to include in their projects the sampling of meiofauna from mobile bottoms. It is only with the help of such cooperative collecting work that examination, determination and research on small benthic animals such as Caudo- foveata and Soleno-gastres can be satis- factorily carried out. This cooperation is vital to adequately enlarge our know- ledge about the organization, biology and biogeography of these primitive mo- lluscan groups that still bear calcareous bodies instead of a shell. Furthermore, this knowledge is essential to also under- stand Mollusca in general. Finally, it must be underlined once more that the determination of members of both classes requires detailed examina- tions (see SALVIN-PLAWEN, 1975 versus ODHNER, 1921, for Caudofoveata; serial sections for most Solenogastres). The nature of the problem becomes more evident when one examines the con- crete documentation. With respect to the presently-known caudofoveate and sole- 36 nogastre fauna in Northeast-Atlantic (Eu- ropean) waters, there is a distinct gradient between the Scandinavian-British records (cf. SALVINI-PLAWEN, 1975; SEAWARD, 1982, 1991) and the West-European reports. Ex- cept for a recent collection from four sites off the coast of Galicia (in connection with the project “Fauna Ibérica”; in prepara- tion), other knowledge of aplacophoran representatives from French and Iberian shelf regions is restricted to a few random findings. On the other hand, investigations and records of both Caudofoveata and So- lenogastres are again available from the Mediterranean Sea. For Europe asa whole, this results in an almost “bipolar” pattern of intraspecific distribution. Clearly, in contrast to the regular sampling in Scan- dinavian, British and Mediterranean seas (cf. SALVINI-PLAWEN, 1972, 1975, 1977a, b, 1988; SEAWARD, 1991), no purposeful offs- hore samplings have been conducted on the Iberian and French shelf in the past to obtain at least an overview of West-Euro- pe's small benthic fauna including Cau- dofoveata and Solenogastres (the French BIOGAS and POLYGAS samplings lie be- yond the European shelf region). The overall knowledge on Caudofo- veata and Solenogastres has significantly increased during the last decades, but is still very poor when compared with ot- her Mollusca (for an organisational and structural overview see SALVINI-PLAWEN, 1985b, and SCHELTEMA, TSCHERKASSKY AND KUZIRIAN, 1994, respectively; their phylogenetic status is analysed in SAL- VINI-PLAWEN AND STEINER, 1996). The poor, random information on their occu- rrence in West-European waters also ne- gatively affects our knowledge on the full range of organisation (systematics, com- parative anatomy) as well as on biologi- cal conditions and circumstances. The above-mentioned “bipolarity” in intraspecific distribution, with the inter- vening West-European gap, becomes ob- vious when considering all aplacophoran representatives known from both nort- hern and southern waters; these are do- cumented below. Other species with an up to now purely Mediterranean or North-European distribution have a po- tential West-European occurrence (Lusi- SALVIN-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres : Of, Gibraltar Figure 1. Section of the West-European Atlantic demonstrating the ofÉshore shelf region down to the 200 m isobath (followed by the continental decline to 3000 m depth). Figura 1. Sección del Atlántico europeo mostrando la zona de la plataforma continental, hasta la isóba- ta de 200 m (seguida de la zona del talud continental hasta los 3000 m de profundidad). tanic region and / or Bay of Biscay ); exam- ples given below are the caudofoveates Psilodens tenuis and Chaetoderma strigis- CAUDOFOVEATA The Caudofoveata burrow within mobile bottoms and have adapted a ver- miform body with reduced pedal sole quamatum as well as the solenogastres Bi- serramenia psammobionta and Anamenia gorgonophila. (midventral fusion of the lateral mantle rims). Generally, sampling in muddy bio- topes (sledge-dredges, grabs) success- 37 Iberus, 15 (2), 1997 fully yields specimens. This group (for- merly Chatodermomorpha) was separa- ted from the Solenogastres and elevated to class rank due to the paraphyletic sta- tus of its aplacophorous organisation (see SALVINI-PLAWEN AND STEINER, 1996). It includes 98 named species classified into three families (Limifossoridae, Prochae- todermatidae, Chaetodermatidae). Se- venteen European representatives have been described so far, six of which occur in the Mediterranean including three en- demic species (cf. SALVINI-PLAWEN, 1990). All species of the West-European shelf region along with the Iberian waters of the Mediterranean will be documen- ted. Thus, among the Prochaetodermati- dae, the deep-sea species Prochaetoderma yongei Scheltema, P. clenchi (Scheltema), and P. (Chevroderma) turnerae (Scheltema) from the BIOGAS-cruises are not consi- dered. These three species also inhabit the basin of the Bay of Biscay (2* 10' - 92 W> at depths of 1175-2006 m (P. yongei), 1913-2430 m (P. clenchi) and 2124-4760 m (P. turnerae) (see SCHELTEMA, 1985; for ta- xonomy cf. SALVINI-PLAWEN, 1992). Besides Falcidens aequabilis, other spe- cies of Falcidens (Chaetodermatidae) are likewise of biogeographical interest: at le- ast among the known species which are provided with a slender, tail-like poste- rior body, each appears to inhabit a well- defined, non-overlapping geographic re- gion. Thus, E gutturosus (Kowalevsky, 1901) is Mediterranean (endemic), while F. crossotus Salvini-Plawen has a Scandi- navian-British distribution. A third “tai- led” species, Falcidens vasconiensis Sal- vini-Plawen, comes from the Gulf of Gas- cogne (SALVINI-PLAWEN, 1996), and future (not yet recorded) Lusitanic repre- sentatives may well belong to yet another species. Family LIMIFOSSORIDAE Scutopus ventrolineatus Salvini-Plawen, 1968 Known distribution (Figure 2A): Scan- dinavian coast (Skagerrak to Tromso0), North Sea, West-Scotland, Irish Sea, southern Bay of Biscaya, Alborán Sea (off Vélez-Málaga), Gulf of Lion (off Banyuls, off Marseille), SE Africa (off Durban); 40-1248 m. Remarks: The occurrence of this very slender and often coiled species has been summarised in SALVINI-PLAWEN (1975, 1977b). Supplementary records come from the North Sea (Hartley, 1984) and from off Barcelona / Catalonia with the cruises RETRO 1 (41? 08' 07” N, 022 04” 32” E, 510 m) and ESPERMA 89 (41? 0437" N, 019 5933" E, 600'm) carried out by Luis Dantart; a recent finding comes from off Vélez-Málaga (4* 03” W) at 400 m. This species is of special inte- rest insofar as it has also been recorded from off Southeast Africa, which indica- tes a distribution along all East-Atlantic. Scutopus robustus Salvini-Plawen, 1970 Known distribution (Fig. 2B): Off the Norwegian coast with larger gaps from Oslofjord to North of Trondheimsfjord, scattered in the Western Mediterra- nean Sea to 9? East; 50-3542 m. Remarks: There are no additional records referring to this slender, up to 10 mm species beyond the occurrence summarised in SALVINI-PLAWEN (1975, 1977a). Psilodens tenuis Salvini-Plawen, 1977 Known distribution (Fig. 2C): Lusita- nic Atlantic S of Cap Sao Vicente; 2500 m. 38 Remarks: There is a single record only, as communicated in SALVINI- PLAWEN (1977a). SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres Family PROCHAETODERMATIDAE Prochaetoderma raduliferum (Kowalevsky, 1901) Chaetoderma radulifera Kowalevsky, 1901, Archs. Zool. exp. gén., sér. 3, 9: 264. Known distribution: Endemic in the Mediterranean Sea, known from the Sea of Marmara in the East to off the Alge- rian coast in the West; 30-2415 m. Remarks: To date, this species is known only from the Mediterra-nean Sea (see map Abb. 5 in SALVINI-PLAWEN, 1977b). Unlike most other members of the Prochaetodermatidae, P. raduliferum is not a true deep-sea species. As is demonstrated by Adriatic and lonic samplings (30-215 m) as well as by records from off Banyuls (60-275 m) summarised in SALVINI-PLAWEN (1977b), it is quite regularly found on muddy offshore bottoms. In accordance with this, there are new records from off the West coast of Malta at 120-160 m (MrirsuD, 1996; the specimen photograp- hed by Mirsub Fig. 2, however, is a broken Falcidens gutturosus, see below), from off Barcelona /Catalonia by Luis Dantart (four stations at 41? 04' 37”-41? OIM AN AOS ASS OLA SE SOS 680 m), and most recently from off Vélez-Málaga to off Málaga (80-300 m). Family CHAETODERMATIDAE Falcidens gutturosus (Kowalevsky, 1901) Chaetoderma gutturosum Kowalevsky, 1901, Archs. Zool. exp. gén., sér. 3, 9: 281. Known distribution: Endemic in the Mediterranean Sea, known from off Palestine and from the Sea of Marmara in the East to off Málaga in the West; 40- 866 m. Remarks: Falcidens gutturosus is a fairly common species characterised by a slender, tail-like posterior body with an orange-red terminal tassle. Beyond the already known, purely Mediterranean distribution (see map Abb. 2 in SALVINI-PLAWEN, 1977b), there are new samplings from off the West coast of Malta at 120-160 m (Mirsub, 1994, 1996), by L. Dantart from off Barcelona (see SALVINI- PLAWEN, 1996) and by A. Zenetos from the Gulf of Korinth as well as from the Gulf of Petalión (56 m; Greece); most recently, specimens were recorded from off Vélez Málaga (40 m) and off Málaga (211 m). Falcidens vasconiensis Salvini-Plawen, 1996 Known distribution: Gulf of Gas- cogne; 141-170 m. Remaks: Up to present there is a single record only from off the Cap Breton in the southeastern Bay or Biscaya (SALVINI- PLAWEN, 1996). Its distribution throug- hout the shelf region of the Gulf of Gas- cogne is to be expected. Falcidens aequabilis Salvini-Plawen, 1972 Known distribution: Endemic in the Mediterranean Sea, ranging from the Aegean Sea to the western Mediterra- nean deep-sea bottom as far as the Greenwich meridian; 132-3542 m. Remarks: This species appears to in- habit deeper and/or far offshore bot- toms. Because of the technical effort in- volved, it is consequently less fre- quently recorded than F. gutturosus or 39 Iberus, 15 (2), 1997 Prochaetoderma raduliferum, but is well- documented from the West-Mediterra- nean deep-sea (Campagne Polymede, cf. SALVINIEPLAWEN, 1977a and 1977b: map Abb. 2). There is a new record by Luis Dantart from off Barcelona / Catalonia (RETROTATAOI406AN 02 03 OLE 350-426 m). Chaetoderma (?) strigisquamatum Salvini-Plawen, 1977 Known distribution (Fig. 2C): Basin of Alborán (W-Mediterranean Sea); 1491 m. Remarks: There is a single record of this chaetodermatid species, whose generic classification needs confirmation (radula SOLENOGASTRES The Solenogastres (= “those with a belly furrow”) include narrowed apla- cophoran molluscs that still bear a pedal groove to glide upon (formerly Neome- niomorpha). A total of some 190 species has been described which, in accordance with integumentary characters, is grou- ped into four orders (Pholidoskepia, Ne- omeniamorpha, Sterrofustia, Cavi- belonia). To date, 46 species are known in European waters, of which 26 (inclu- ding 19 “endemics”) are represented in the Mediterranean Sea. However, little is known about their biogeography: most species have been recorded only once, and inaccurate descriptions (see e. g. Wirenia, below) additionally contri- bute to difficulties in classification, re- sulting in insufficient faunistic informa- tion. Thus, the poorly described Nemato- menia (?) corallophila (Kowalevsky, 1881), recorded from off La Calle / Algeria at 73-183 m (37? N, 8* 30” E) as living epi- zoically on Corallium rubrum (Linné), apparatus; cf. SALVINI-PLAWEN, 1977a). Ba- sed on the known distribution of Caudo- foveata in the Mediterranean Sea in ge- neral (SALVINI-PLAWEN, 1977b), a Lusita- nic occurrence of this species might be expected. could only be recognised in the future and re-described if it is rediscovered again on a red coral (its alledged finding in the Bay of Rosas/Costa Brava is a mistranslation by Mars, 1965, from Ma- LUQUER, 1917). Most species documented here, toget- her with a few others, belong to the small number of representatives found several times. Records of these findings are very much tied to sampling methods and habitat. For example, the well-known Neo- menta carinata was never recorded by means of sledge-dredges (muddy bottoms), as predominantly used by the author and his group. In another example, those Solenogastres living on thecapho- ran Hydrozoa or Octocorallia (e. g. Nema- tomenia and Anamenia, below), are more often sampled from secondary hard bottoms (e. g. with Agassiz-trawls) or by workers studying cnidarians. Thus, all these circumstances help explain our frag- mented biogeographic knowledge. Order PHOLIDOSKEPIA Family DONDERSIDAE Nematomenta flavens (Pruvot, 1890) Dondersia flavens Pruvot, 1890, Archs. Zool. ex. gén., sér. 2, 8: XXIL Known distribution (Fig. 2D): Off Banyuls - Costa Brava, Shetland Islands; 45-167 m. 40 Remarks: This slender, up to 4 cm long species has a showy lemon- yellow colour. It is not rare along the SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres rocky (French and Spanish) Catalan coast, feeding upon Hydrozoa-The- caphora at 45-90 m (Pruvor, 1891; MALUQUER, 1917; Mars, 1965). Ano- ther record refers to the Shetland Islands at 167 m, epizoic upon Lafoea dumosa SARS; the anatomical examina- tion revealed the presence of a vesti- gial radula and the necessity for a family reclassification (SALVINI-PLA- WEN, 1978: 39-40). Recently, the Irish Sea Survey (MACKIE, OLIVER AND REES, 1995: 192) collected eleven samples of N. banyu- lensis (see below); however, there is no information about the exact methos of determination applied. The sampled material had been fixed in formalin and then preserved in alcohol (MACKIE ET AL., 1995: 15-16); the specific body- colour of both N. banyulensis (red) and N. flavens (yellow) is no longer visible after such treatment. Therefore, it might well be that some of the N. ban- yulensis-records in reality belong to the externally very similar N. flavens. Only an accurate histological determination (serial sections) can provide the exact specific classification of these speci- mens. Nematomenta banyulensis (Pruvot, 1890) Dondersia banyulensis Pruvot, 1890, Archs. Zool. ex. gén., sér. 2, 8: XXIL Nematomenia banyulensis var. norvegica Odhner, 1921, Bergens Mus. Aarb. 1918-19, Naturvid. reekke 3: 43. Muyzomenia Simroth, 1893, Zeitschr. wiss. Zool., 56: 324. Known distribution (Fig. 2E): Off Dal- matia, Gulfs of Naples and Salerno, Cóte Vermeille, off Roscoff, Plymouth Sound to Irish Sea to W-Scotland, off Northumberland, Trondheimsfjord-Fill (amfjord; 31-300 m. Remarks: This well-known species likewise lives epizoically upon Hydro- zoa-Thecaphora. Its slender body reaches a length of up to 3 cm and is red (as are also two other Mediterranean species). Its distribution is summarised in NIERSTRASZ AND STORK (1940) and more recently in SEAWARD (1982, 1991) for the British waters. It has also recently been found several times by the Irish Sea Survey (MACKIE ET AL., 1995: 192), but compare the above remarks with N. flavens. Geographically new records include samples from off Sebe- nico /Sibenik (Adriatic Sea) at 57 m, 61 m and 67-68 m (see also SALVINI- PLAWEN, 1986) and from the Fill (an)fjord = north-eastern Hitra Island off Trondheimsfjord (Mus. Uppsala). A comparative examination of Medi- terranean and Norwegian (syntype) individuals, particularly with respect to the mantle scales, revealed no differen- ces which would vindicate a separation of the Norwegian specimens (as varia- tion or subspecies proper). Stylomenia salvatori Pruvot, 1899 Known distribution: Off Banyuls, (?)Costa Brava; about 60-80 m. Remarks: This species had been found together with Rhopalomenia agla- opheniae (q. v.) in an aquarium filled with benthic material from off Banyuls- sur-Mer; based on the presence of Rh. aglaopheniae, this indicates an original depth of about 60-80 m (see PrUVOT, 1891: 721). MALUQUER (1916: 244, 1917: 37-38) reports finding animals similar to S. salvatori from the Bay of Rosas and off Llansa (Costa Brava). Even if the occurrence of this species is to be expec- ted there, the record needs to be confir- med because no accurate determination (histological examination) was perfor- med. 41 Iberus, 15 (2), 1997 Family LEPIDOMENIIDAE Tegulaherpia myodoryata Salvini-Plawen, 1988 “Species D” in Salvini-Plawen, 1968a, Sarsia, 31: 132. Tegulaherpia celtica Caudwell, Jones and Killeen, 1995, Journ. Conch. (London), 35: 258. Known distribution (Fig. 2F): Off Li- vorno, off Banyuls-sur-Mer, southern Bay of Biscay (North of Asturias, THALASSA- Stat. W-415)?, Celtic Sea, area around Ber- gen (Raunefjord, Hjeltefjord), area around Trondheim (Fill (an)fjord, Trondheimsf- jord); 75-470 (75-860 / 1150?) m. Remarks: This Mediterranean species is likewise native to Northern Europe. In the course of examining more compre- hensive Solenogastres material from the North Atlantic, the already communica- ted “Species D” (SALVINI-PLAWEN, 1968a) and T. celtica (CAUDWELL, JONES AND KI- LLEEN, 1995), according to histological exa- mination by series sections, turned out to be conspecific with T. myodoryata from the Western Mediterranean Sea as described in SALVIN-PLAWEN (1988). Moreover, se- veral other Norwegian individuals (Hjel- tefjord, 200 m, Fill (an)-fjord, Trond- heimfjord, 470 m) likewise belong to this species. A single specimen from the THA- LASSA-Cruise (Stat. W-415, 439 55' 06” N, 06” 11 18” W; 860-1150 m), forwarded in 1971 by F. Monniot (Paris) to the author, possibly also represents T. myodoryata, since the mantle scales fully fit into the range of shape, outline and size of the myo- doryata-scales. However, the animal was useless for histological examination, and the record from a depth between 860 and 1150 m lends doubt to a conspecificity as long as no bathymetrically interbridging and /or additional samples are taken. Family WIRENIIDAE Wirenia argentea Odhner, 1921 Aesthoherpia glandulosa Salvini-Plawen, 1985, The Mollusca (Academic Press), 10: 94. “Species B” in Salvini-Plawen, 1968a, Sarsia, 31: 131. Aesthoherpia glandulosa Salvini-Plawen and “Species D” Haszprunar, 1986, Zool. Anz., 217: 345-360. Known distribution (Fig. SA): Area around Bergen/Norway, Hardanger- fjord, area of Trondheimsfjord, Adriatic Sea, Aegean Sea; 95-700 m. Remarks: A most recent examination of the hitherto missing type material of Wirenia argentea Odhner (now in the Naturhistoriska Rijkmuseet, Stockholm) revealed that Aesthoherpia glandulosa Salvini-Plawen is conspecific with it. Despite some inaccurate and insufficient presentations by ODHNER (1921: 31-34; foregut, no radula, and so on), which led to the description of Aesthoherpia (see PLAWEN, 1988: 383-384), Wirenia has nomenclatorial priority. The organisation of the species and its presently known geo- graphic distribution are communicated (as Aesthoherpia glandulosa) in SALVINI- PLAWEN (1988). Some additional findings come from recently examined Norwegian samples: area Northwest of Bergen (Hjel- tefjord, 280 m; Herdlafjord, 200 m; Man- gerfjord, 350 m) and Fill (an)fjord-Trond- heimsfjord (95 m, 185 m, 320 m, 470 m and 490-500 m). The record of “Wirenia argen- tea” by HARTLEY (1984) from the North Sea needs specific confirmation. Family MACELLOMENIDAE Macellomenta palifera (Pruvot, 1890) Paramenia palifera Pruvot, 1890, Archs. Zool. exp. gén., sér. 2, 8: XXIIL 42 SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres ¡IS 7 Ñ N DN Figure 2. A-C. Caudofoveata. Known European distribution. A: Scutopus ventrolineatus Salvini- Plawen, 1968; B: Scutopus robustus Salvini-Plawen, 1970; C: Known records of Psilodens tenuis Salvini-Plawen, 1970 (black circle) and of Chaetoderma(?) strigisquamatum Salvini-Plawen, 1971 (asterisk). D-F: Solenogastres. Known distribution. D: Nematomenia flavens (Pruvot, 1890); E: Nematomenia banyulensis (Pruvot, 1890); E: Tegulaherpia myodoryata Salvini-Plawen, 1988. Figura 2. A-C. Caudofoveados. Distribuciones conocidas. A: Scutopus ventrolineatus Salvini-Plawen, 1968; B: Scutopus robustus Salvini-Plawen, 1970; C: Citas conocidas de Psilodens tenuis Salvini- Plawen, 1970 (círculo negro) y de Chaetoderma(?) strigisquamatum Salvini-Plawen, 1971 (asterisco). D-F: Solenogastros. Distribuciones conocidas. D: Nematomenia flavens (Pruvot, 1890); E: Nematomenia banyulensis (Pruvot, 1890); F: Tegulaherpia myodoryata Salvini-Plawen, 1988. 43 Iberus, 15 (2), 1997 Known distribution (Fig. 3B): Cóte Vermeille, Irish Sea (?); 80-120 m. Remarks: This species, with its parti- cular calcareous mantle-bodies, was ori- ginally recorded with a single specimen North of Port Vendres (Cóte Vermeille; southeastern France) on muddy bottom at 80 m. Two individuals recently sampled from the Irish Sea at 80 m and 120 m come very close to M. palifera (CAUDWELL ET AL., 1995). In view of the “bipolar” occurrence of other species (demonstrated herein), there is a high probability that the species are identi- cal. However, as the British animals have not been investigated anatomically (series sections), true conspecificity ramains uncertain. Order NEOMENIAMORPHA Family NEOMENIIDAE Neomenta carinata Tullberg, 1875 Solenopus nitidulus Koren and Danielssen, 1877, Arch. Math. Naturvid. (Kristiania), 2: 124. Solenopus affinis Koren and Danielssen, 1877, Arch. Math. Naturvid. (Kristiania), 2: 127. Neomenia grandis Thiele, 1894, Zeitschr. wiss. Zool., 58: 223. Known distribution (Fig. 3C): Nor- thern Kattegat and Bohuslán (W-Swe- den), Norwegian coast between Oslo- fijord and Sognesjóen/Sogne-fjord, Romsdalsfjord, Trondheimsfjord, South of Lofoten, Iceland, Shetland Islands, British Isles, off Roscoff, Costa Brava, Gulf of Genova, Gulf of Naples, off Mes- sina; 10-565 m. Remarks: This up to 3 cm long species has a stoutish shape and is well docu- mented along the coast of Scandinavia and around the British Isles (KOREN AND DANIELSSEN, 1879; WIRÉN, 1892; ODHNER, 1921; Muus, 1959; SEAWARD, 1982, 1991) including Strindfjord / Trondheimsfjord (Mus. Copenhagen) and the Hebrides (Mus. Leiden). The Mediterranean records include N. affinis (Koren and Danielssen) which, according, to certain, minor anatomical differences (pers. obs.), can be classified as a subspecies only (SALVINI-PLAWEN 1986); the same holds true for N. grandis Thiele (NIERSTRASZ AND STORK, 1940). A remarkable record refers to Iceland (KNUDSEN, 1949), a region in which Neomenia dalyelli (Koren and Danielssen) is generally found. Order CAVIBELONIA Family PARARRHOPALIDAE Eleutheromenia sierra (Pruvot, 1890) Paramenta sierra Pruvot, 1890, Archs. Zool. exp. gén., 8: XXUL Known distribution (Fig. 3D): Costa Brava, Bretagne; Irish Sea; Trondheim region; 40-128 m. Remarks: PRUVOT (1891) typifies the species from a single specimen (Cap Creus / Costa Brava; 80 m) and in 1897 he reports another finding from off Roscoff at about 40 m (Pruvor, 1897). A single spe- cimen of typical appearence (lobed dor- somedian keel) from Stjórn (North or Trondheim / Norway), despite the geo- graphical distance, after serial section reve- 44 aled to be Eleutheromenia sierra. Conse- quently, the questioned presence or this species in the southwestern Cartigan Bay (Irish Sea, 52 m; see HARTLEY 1984, SEAWARD, 1991: 14) as well as the speci- mens from nine samples or the Irish Sea Survey referred to E. sierra (CAUDWELL ET AL., 1995: 266; MACKIE ET AL., 1995: 192; not documented in Fig. 3D) are indirectly confirmed to really belong to this species. On the other hand, the Pararrhopalii- dae represent a fairly diverse group of sys- SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres Figure 3. Solenogastres. A: Known distribution of Wirenía argentea Odhner, 1921; B: Records of Ma- cellomenia palifera (Pruvot, 1890) (black circles) and of Meromenia hirondellei Leloup, 1949 (asterisk); C: Known distribution of Neomenia carinata Tullberg, 1875; D: Records of Eleutheromenia sierra (Pruvot, 1890), see text; E: Known distribution of Biserramenia psammobionta Salvini-Plawen, 1968; E: Known European distribution of Rhopalomenia aglaopheniae (Kowalevsky and Marion, 1887). Figura 3. Solenogastros. A: Distribución conocida de Wirenia argentea Odhner, 1921; B: Citas de Macel- lomenia palifera (Pruvot, 1890) (círculos negros) y de Meromenia hirondellei Leloup, 1949 (asterisco); C: Distribución conocida de Neomenia carinata Tullberg, 1875; D: Citas de Eleutheromenia sierra (Pru- vot, 1890), véase texto; E: Distribución conocida de Biserramenia psammobionta Salvini-Plawen, 1968; F: Distribución europea de Rhopalomenia aglaopheniae (Kowalevsky y Marion, 1887). 45 Iberus, 15 (2), 1997 tematically very difficult representatives (see SALVINI-PLAWEN, 1978); several geo- graphically close records of Pararrhopa- liidae may represent different species (or even genera). Thus, another record from the Irish Sea (Pruvotina sp. in CAUDWELL ET AL., 1995: 265-267) clearly does not belong to E. sierra; the same can be said about two THALASSA-specimens from off Galicia and off Asturias (Bay of Biscay) Family SIMROTHIELLIDAE Biserramenta psammobionta Salvini-Plawen, 1968 Known distribution (Fig. 3E): Irish Sea, Plymouth area, Bretagne, Galicia; 8-30 m. Remarks: In addition to the type mate- rial from off Roscoff at 8-10 m (SALVINI- PLAWEN, 1968b; see also MONNIOT, 1965), several individuals have recently been recorded in Plymouth Sound at 9-11 m (50% 20” 43” N, 4? 09 05” W; see also KIKINGER AND SALVINI-PLAWEN, 1995). Moreover, a single specimen has been sampled by lan Killeen during the Irish Sea Survey from the Cardigan Bay / Wales (Stat. 46, 52* 19% 12” N, 04? 37" W) at 30 m, provided by Cathy Caudwell to the author (see also CAUDWELL ET AL., 1995). Finally, the 12 Solenogastres refe- rred to as “Lepidomenia sp.” by Celia Bes- teiro in her Ph. D. thesis (1986) from Galicia / Spain (Ría de Ferrol, “Bajo de la Palma”; 43? 27' 59” N, 08* 16' 23” W; 14 m) also represent Biserramenia psammo- bionta. They all come from coarse sand or shell gravel bottoms and at least those from Roscoff, Plymouth and Galicia are interstitially living animals (cf. SALVINI- PLAWEN, 1985a; also OTT AND BOcH- DANSKY, 1991, for the Plymouth animals). The histological examination of these specimens revealed some details beyond the original description (SALVINI- PLAWEN, 1968b). First, the characteristic circular musculature around the spaw- ning ducts and the posterior mantle cavity is not yet elaborated in juveniles. The slender pericardioducts with a tiny lumen still open from dorsal into the spawning ducts close to their rostral ends, these ducts being paired throug- hout with a wide lumen. Further diffe- rentiation thus includes a curving elon- gation of the rostral portion of the spaw- ning ducts, which results in the adult spawning ducts bending dorso-poste- riorly (as described in 1968). Here, the pericardioducts join their ends not axially but ventrally, thus causing a bulgy enlargement or even a slight bend in the continuous lumen. This bulged enlargement is the site of sperm storage, thus functioning as receptacula seminis; well-defined, set-off seminal pouches (vesiculae seminales, as described earlier), however, are not present. The paired lateral pouch of the ventrorostral mantle cavity is well differentiated only in fully-grown individuals and often merely represents two simply lobulated, more or less distinct sacculations opening medially through a short duct or pore into the pallial space. Rather than being paired in the sense of two sin- gular, separate ganglia, the cerebral ganglia are fused together in the middle third of their extension. In some speci- mens a small dorsoterminal sense organ could be detected at the rear of the body. Family AMPHIMENIIDAE Meromenia hirondellei Leloup, 1949 Known distribution (Fig. 3B): Nort- hern Bay of Biscay; 166 m. Remarks: A single fragment of this species had been recorded from the con- tinental platform in the northern Bay of 46 Biscay (46” 27” N, 4* 09% 45” W) at 166 m depth. Due to the unknown organisa- tion of the anterior body, the generic classification is uncertain (LELOUP, 1950: 21-23; SALVINI-PLAWEN, 1972: 224-225). SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres 20 o y Figure 4. Solenogastres. A: Known European distribution of Anamenia gorgonophila (Kowalevsky, 1880); an additional record refers to the Azores; B: Known distribution of Dorymenia sarsii (Koren and Danielssen, 1877). Figura 4. Solenogastros. A: Distribución europea de Anamenia gorgonophila (Kowalevsky, 1880); una cita adiccional se refiere a las islas Azores; B: Distribución conocida de Dorymenia sarsii (Koren y Daniels- sen, 1877). Family RHOPALOMENIDAE Rhopalomenia aglaopheniae (Kowalevsky and Marion, 1887) Rhopalomenia eisigi Thiele, 1894, Zeitschr. wiss. Zool., 58: 269. Known distribution (Fig. 3F): Off Cap Matapan/Tainaron (South-Peleponnes), Gulf of Naples, off Marseille, Cóte Ver- meille, off Roscoff, British Isles; 50-137 m. Remarks: This well-known species li- ves upon Hydrozoa-Thecaphora, almost exclusively upon Lytocarpia myriophyllum (Linné). The distribution is compiled in NIERSTRASZ AND STORK (1940), SALVINI- PLAWEN (1972) and SEAWARD (1982, 1991). The identification of several specimens from off Monrovia / Liberia (THIELE, 1906: 324) needs re-examination and / or confir- mation. Family STROPHMENIDAE Anamenia gorgonophila (Kowalevsky, 1880) Proneomenia nierstraszi Stork [in Nierstrasz and Stork], 1940, Zoologica (Stuttgart), 99: 57. Anamenia heathi Leloup, 1947, Bull. Mus. roy. Hist. Nat. Belgique, 23 (26): 1-11. Known distribution (Fig. 4A): Gulfs of Naples and Salerno, off La Calle (eas- ternmost Algeria), off Marseille, Sea of Alborán, Gorringe-Bank (WSW of Cap Sao Vicente), Azores; 65-845 m. Remarks: The records of this species have beenrevised by SALVINI-PLAWEN (1972). The investigation of numerous Solenogastres recorded more recently from the SW of the Isle of Alborán (see TEMPLADO, GARCÍA- CARRASCOSA, BARATECH, CAPACCIONI, JUAN, LÓPEZ-IBOR, SILVESTRE AND MASSÓ, 1986: 101-102) revealed that they in part belong to A. gorgonophila, and the known distribution of the species supports the assumption of its presence in Lusitanian waters as well. This species lives upon Gor- gonaria, predominantly upon Paramuricaea clavata (Risso) = P. chamaeleon (Koch), but also upon Eunicella spp. and others. 47 Iberus, 15 (2), 1997 Family PRONEOMENIIDAE Dorymenia sarsil (Koren and Danielssen, 1877) Simrothiella sarsi Auctt. (see Opinion 1185 ICZN) Known distribution (Fig. 4B): Trond- heimfjord, Sognefjord, Bergen area, Os- lofjord, Skagerrak, Gorringe Bank (off Cap Sao Vicente); North Atlantic-Arctic Ocean outside Troms0?; 183-681 m (1134 m?). Remarks: The up to 7 cm long, slender species was redescribed by ODHNER (1921) and is externally characterised by a dis- tinct (dorso-)terminal extension of the body; a photograph is given in SALVINI- PLAWEN (1968a: Abb. 23). Some recently examined material from Scandinavian col- lections extends the known distribution (see Fig. 4B); the North Atlantic-Arctic spe- cimens (71 25 N, 15” 41' E, 1134 m; see ODHNER, 1921, and JAECKEL, 1954) only doubtfully belong to D. sarsii based on the geographic and bathymetric distribution. In addition to the polystichous radula, a pair of copulatory stylets and the presence of a dorsoterminal sense organ typical for the genus, histological investigations un- derline two particular specific characters in mature animals: the anterior portion of the pericardioducts bears small pockets serving as vesiculae seminales, and the posterior portion of the spawning ducts (“lower gametoducts”) in front of their fu- sion each elaborate a ventral enlargement or voluminous sacculation (pocket). Both these characters allow this particular Dory- menta species to be identified in SCHEL- TEMA ET AL. (1994: Figs. 22 E and 24 G) as D. sarsii (Koren and Danielssen). The ge- ographic distribution of this species is thus enlarged to the Gorringe Bank of the Ibe- rian shelf region (36? 50' N, 9* 15 W, 681 m, Cf. SCHELTEMA ET AL., 1994: 18). Family LEPIDOMENIIDAE Lepidomenia ? spp. Lepidomenia hystrix Auctt. non Marion and Kowalevsky, 1886 Lepidomenia (?) swedmarki Salvini-Plawen, 1985, Stygología, 1 (1): 103. Remarks: Some records of small, mesop- sammic Solenogastres from off Marseille, Bretagne and off Belfast / Northern Ireland have been systematised as Lepidomenia hystrix Marion and Kowalevsky (see SALVINI-PLAWEN, 1985a; SEAWARD, 1991). Referring to the discussion in SALVINI- ACKNOWLEDGEMENTS The updating of the geographical distribution in part comes from the Scandinavian material in elaboration by REFERENCES CAUDWELL, C. M., JONES, A. M. AND KILLEEN, [. J., 1995. Three Solenogastres from the Irish Sea, new to the British marine area. Journal . of Conchology (London), 35: 257-269. 48 PLAWEN (1985a), the classification of these animals (not having been investigated ana- tomically) as L. hystrix is unwarranted and misleading. The specimens should be determined by means of serial sections rather than simply naming them based on accessory features. Dr. Claudia Handl (Wien) supported by the grant P 9075-B of the Austrian Science Foundation FWFE to the author. HARTLEY, J. P., 1984. The benthic ecology of the Forties Oilfield (North Sea). Journal of Expe- rimental Marine Biology and Ecology, 80: 161- 195: SALVINI-PLAWEN: West-European and Iberian Caudeofoveata and Solenogastres HASszPRUNAR, G., 1986. Feinmorphologische Untersuchungen an Sinnesstrukuren ur- sprúnglicher Solenogastres (Mollusca). Zoo- logischer Anzeiger, 217: 345-362. JAECKEL, G. A., 1954: Aculifera Aplacophora und Aculifera Placophora im Gebiet der Nord- und Ostsee. Kieler Meeresforschungen, 10: 261-271. KIKINGER, R. AND SALVINI-PLAWEN, L. v., 1995. Morphogenesis from polyp to Stauromedusa in the genus Stylocoronella (Cnidaria: Scyp- hozoa). Journal of the Marine Biological Asso- ciation of the United Kingdom, 75 (4): 899-912. KNUDSEN, J., 1949. Amphineura. The Zoology of Iceland, 4 (59): 1-11. KOREN, J. AND DANIELSSEN, D. C., 1879. Des- criptions of new species belonging to the Ge- nus Solenopus, with some observations on their organization. Annals and Magazine of Natural History (ser. 5), 3: 321-328. LELOUr, E., 1950. Solénogastres provenant des campagnes du Prince Albert ler de Monaco. Résultats des Campagnes scientifiques par Albert ler de Monaco, 110: 1-38. MACKIE, A. S. Y., OLIVER, P. G. AND REES, E. I. S., 1995. Benthic Biodiversity in the southern Irish Sea. BIOMOR Reports (Natural Museum of Wales), 1: 1-263. MALUQUER, J., 1916. Treballs oceanografics en la costa del Empurda. Anuari Junta Ciencies Naturales Barcelona, 1: 221-261. MALUQUER, J., 1917. Notes per a l'estudi dels So- lenogastres (Molluscos Amfineures) de Ca- talunya. Treballs de la Institució catalana d'- Historia Natural, 3: 9-53. Mass, P., 1965. Faune marine des Pyrénées- Orientales (5): Mollusques Aplacophores, Polyplacophores, Scaphopodes et Bivalves. Vie et Milieu, Suppl. 15 (4): 1-156. MirSUD, C., 1994. Falcidens gutturosus (Kowa- levsky, 1901) (Mollusca, Caudofoveata). New to the Maltese circalittoral fauna! La Conchi- glía, 26 (270): 11-12. MirsubD, C., 1996. Living Mollusca from circa- littoral coastal muds, off Western Malta. La Conchiglia, 28 (suppl. to 279): 23-41. MONMNIOT, F., 1965. Ascidies interstitielles des cótes d'Europe. Mémoirs du Muséum Natio- nal d'Histoire Naturelle (Paris), Sér. A, 35: 1- 154. MUuus, B., 1959. Skallus, Soteender, Bleeksprut- ter. Danmarks Fauna, 65: 1-239. NIERSTRASZ, H. AND STORK, H., 1940. Mono- graphie der Solenogastren des Golfes von Nea- pel. Zoologica (Stuttgart), 99: 1-99, Taf. fXV. ODHNER, N., 1921. Norwegian Solenogastres. Bergens Museums Aarbok 1918-19, Naturvi- denskabelig reekke, 3: 1-82. OPINION 1185: ICZN 1981. Simrothiella Pilsbry, 1898 (Mollusca, Solenogastres): Designation of a type species. Bulletin of Zoological No- menclature, 38 (3): 185-187. Orr, J. AND BOCHDANSKY, A., 1991. Meiofauna of marine sediments. PAL-Videofilm (Pro Mare, Vienna). Pruvor, G., 1891. L'organisation de quelques Néoméniens des cótes de France. Archives de Zoologie expérimentale et générale, sér. 2, 9: 699- 810. Pruvor, G., 1897. Essai sur les fonds et la faune de la Manche occidentale (Cótes de Bretag- ne) comparés a ceux du Golfe du Lion. Archi- ves de Zoologie expérimentale et générale, sér. 3, 5: 511-660. SALVINI-PLAWEN, L. v., 1968a. Uber einige Be- obachtungen an Solenogastres (Mollusca, Aculifera). Sarsia, 31: 131-142. SALVINI-PLAWEN, L. v., 1968b. Neue Formen im marinen Mesopsammon: Kamptozoa und Aculifera. Annalen des Naturhistori- schen Museums Wien, 72: 231-272. SALVINI-PLAWEN, L. v., 1972. Revision der monegassischen Solenogastres (Mollusca Aculifera). Zeitschrift fúr zoologische Syste- matik und Evolutionsforschung, 10: 215-240. SALVINI-PLAWEN, L. v., 1975. Mollusca: Cau- dofoveata. Marine Invertebrates of Scandina- via, 4: 1-55. SALVINI-PLAWEN, L. v., 1977a. Caudofoveata (Mollusca) des Forschungsprojektes Poly- mede. Bulletin du Muséum Nationale d'His- toire Naturelle (sér. 3), No. 447, Zoologie Vol. 310: 413-421. SALVINI-PLAWEN, L. v., 1977b. Caudofoveata (Mollusca), Priapulida und apode Holo- thurien (Labidoplax, Myriotrochus) bei Ban- yuls und im Mittelmeer allgemein. Vie et Milieu, 27A: 55-81. SALVINI-PLAWEN, L. v., 1978. Antarktische und subantarktische Solenogastres. Zoolo- gica (Stuttgart), 128: 1-315. SALVINI-PLAWEN, L. v., 1985a. New intersti- tial Solenogastres (Mollusca). Stygologia, 1: 101-108. SALVINEPLAWEN, L. v., 1985b. Early evolu- tion and the primitive groups. The Mollusca (Academic Press), 10 (Evolution): 59-150. SALVINI-PLAWEN, L. v., 1986. Caudofoveata e Solenogastres del Mediterraneo. Bollettino Malacologico, 22: 189-196. SALVINI-PLAWEN, L. v., 1988. Einige Soleno- gastres (Mollusca) der europáischen Meio- fauna. Annalen des Naturhistorischen Mu- seums Wien, 90B: 373-385. SALVINI-PLAWEN, L. v., 1990. The status of the Caudofoveata and the Solenogastres in the Mediterranean Sea. Lavori della Societa Ita- liana di Malacologia, 23: 5-30. SALVINI-PLAWEN, L. v., 1992. On certain Cau- dofoveata from the VEMA-Expedition. In Gitteenberger, E. and Goud, J. (Eds.): Pro- ceedings of the Ninth International Malacolo- gical Congress. Unitas Malacologica, Lei- den: 318-333. 49 Iberus, 15 (2), 1997 SALVINI-PLAWEN, L. v., 1996. Falcidens vasco- niensis spec. nov. (Mollusca, Caudofoveata) du plateau continental du golfe de Gascogne. Bulletin de la Société Zoologique de France, 121: 339-345. SALVINI-PLAWEN, L. V. AND STEINER, G., 1996. Synapomorphies and plesiomorphies in hig- her classification of Mollusca. In Taylor, J. (Ed.): Origin and evolutionary radiation of the Mollusca. Oxford University Press, Oxford (U. K.): 29:51. SCHELTEMA, A. H., 1985. The aplacophoran fa- mily Prochaetodermatidae in the North Ame- rican Basin, including Chevroderma n. g. and Spathoderma n. g. (Mollusca, Chaetoder- momorpha). Biological Bulletin, 19: 484-529. SCHELTEMA, A. H., TSCHERKASSKY, M. AND KUZIRIAN, M., 1994. Aplacophora. Micros- copic Anatomy of Invertebrates, 5 (Mollusca I): 13-54. 50 SEAWARD, D. R., 1982. Sea Area Atlas of the ma- rine molluscs of Britain and Ireland. Nature Conservancy Council, Shrewsbury, 53 pp., 746 maps. SEAWAROD, D. R., 1991. Distribution of the marine molluscs of North West Europe. Nature Con- servancy Council, Peterborough, 114 pp. TEMPLADO, J., GARCÍA-CARRASCOSA, M., BA- RATECH, L., CAPACCIONI, R., JUAN, A., LÓPEZ- IBOR, A., SILVESTRE, R. AND MAssóÓ, C., 1986. Estudio preliminar de la fauna asociada a los fondos coraliferos del mar de Alborán (SE de España). Boletín del Instituto Español de Oceanografía, 3 (4): 93-104. THIELE, J., 1906. Archaeomenia prisca n. g. n. sp. Wissenschaftliche Ergebnisse der Deutschen Tief- see-Expedition Valdivia, 9: 317-324. WIRÉN, A., 1892. Studien úber die Solenogas- tren II. Kunglige Svenska Vetenskaps-Akade- miens Handlingar, 25 (6): 1-100. O Sociedad Española de Malacología —_—_—_—_—_—_—_———— Iberus, 15 (2): 51-66, 1997 Molluscs as evolving constructions: necessary aspects for a discussion of their phylogeny Los moluscos como construcciones en evolución: aspectos necesarios para una discusión sobre su filogenia Karl EDLINGER* and Wolfgang Friedrich GUTMANN! Recibido el 8-[X-1995. Aceptado el 19-X-1996 ABSTRACT A model for the evolution of molluscs is presented. The reconstruction is based on the Frank- furt Theory which conceives organisms as energy transforming hydraulic units which are subject to evolutionary transformation according to the constructional principles ruling orga- nisation. Evolution is reconstructed as a process of constructional transformations in which all stages are explained as explicitly viable constructions; the irreversible transformation phases are also rationally explained by referring to constructional properties of the orga- nismic machines. lt is maintained that the predecessors of molluscs must have been elongate worm like animals which were internally tethered by muscles in a way that creeping on the flattened ventral “foot” surface became possible. Only organisms controlling the cross-section by a densely spaced muscle system could start creeping on hard substrate. The establishment of the radula is shown to have been dependent on the adhesive creeping movements which allowed anchorage of the construction to the substrate during rasping. The formation of the shell elements was rendered possible by concentration of motility to the ventral side while the dorsal body wall was held undeformed as a precondition for shell formation. Formation of the muscle grid of the foot on the ventral side of the body and the stabilisation by skeletal elements of the dorsal side caused a shift of the inner organs into a dorsal hump. From the model for the primitive molluscs with segmented shells the continuation into the conchiferan constructions with fused shells and the constructional lineages into the major mollusc cons- tructions are given in captions parallelizing sequences of visualisations of the constructional stages with the major alterations. RESUMEN Se presenta un modelo sobre la evolución de los moluscos basado en la Teoría de Frankfurt, que concibe a los organismos como unidades hidráulicas transformadoras de energía suje- tas a transformaciones evolutivas de acuerdo con los principios constructivos que regulan la organización. La evolución se reconstruye como un proceso de transformaciones construc- cionales en el que todos los estados se explican como construcciones explícitamente viables; las fases de transformación irreversibles también se explican racionalmente refiriéndolas a propiedades construccionales de las máquinas orgánicas. Se defiende que los predecesores de los moluscos deben haber sido animales alargados, tipo gusano, que contaban con una malla muscular interna de tal manera que fuese posible arrastrarse sobre la superficie aplanada ventral del “pie”. Sólo aquellos organismos que controlasen su sección mediante un sistema muscular densamente espaciado podían empe- zar a arrastrarse sobre un sustrato duro. * Naturhistorisches Museum Wien, 3. Zoologische Abteilung, Burgring 7, A-1014 Wien, Austria. Prof. Gutmann died on April 1997. Sl Iberus, 15 (2), 1997 La aparición de la rádula se muestra como un proceso dependiente de los movimientos de reptación que permitieron el anclaje de la estructura al sustrato durante el raspado. La for- mación de elementos conchíferos fue posible por la concentración de la motilidad en la cara ventral mientras que la pared dorsal del cuerpo se mantenía sin deformación, paso previo para la formación de una concha. La aparición de la malla muscular del pie en la cara ven- tral del cuerpo y la estabilización de la cara dorsal mediante elementos esqueléticos pro- vocaron el traslado de los órganos internos hacia una ¡joroba dorsal. A partir del modelo de moluscos primitivos con conchas segmentadas se presenta el desarrollo hacia construc- ciones conchíferas con conchas fusionadas y hacia las principales estructuras construccio- nales de moluscos, mediante encabezamientos de sequencias paralelas de estados de de- sarrollo, incluyendo las principales alteraciones. KEY WORDS: Molluscs, hydraulic constructions, phylogenetical reconstruction, metamery, radiation. PALABRAS CLAVE: moluscos, construcciones hidráulicas, reconstrucción filogenética, metamería, radiación. INTRODUCTION Evolution can never be directly obser- ved. It must be reconstructed in models describing the sequences of transforma- tional steps. All stages of evolution neces- sarily have to be represented by explicitly viable organismic constructions. The the- oretical concept and the methodology which allows such reconstructions is the Frankfurt Evolution Theory (FET) (GUTMANN, 1974, 1989; BONIK, GRASSHOFF AND GUTMANN, 1977; GUTMANN AND EDLINGER, 1994a, b, c, d; EDLINGER, 1989a, b; EDLINGER, GUITMANN AND WEINGAR- TEN, 1991; VOGEL, 1991). In accordance with the demands and results of other bio- logical disciplines, this theory conceives of organisms as energy transforming units and as autoformative constructions. Reconstructions of the evolutionary changes in the sense of the Frankfurt Evo- lutionary Theory are supported by the insight into the constructional properties of all animal soft body systems which function as hydroskeleton apparatuses and hydraulic units. Before reconstruc- tions are attempted the principles and physical laws governing the organismic entities must be known. ORGANISMS: ENERGY TRANS- FORMING ENTITIES On the basis of constructional expla- nations metazoan animals must be des- De cribed and explained as self-sustaining and energy transforming systems. Basi- cally, they function as machines. They are capable of actively acquiring matter and energy from their environment. By transforming the chemical energy thus obtained into mechanical force the wor- king activity of the body construction is generated. Energy transformation in the cons- -tructions is determined by the structure of specific macromolecular components, mainly in muscles and cilia. To become effective as driving engines the energy transforming structures have to be inte- grated into an energy cascade of a mecha- nically coherent structural whole in which the forces are transmitted to mechanically working units. A chain of force transmit- ting structures connects the energy trans- forming sites with the working units of the animal body. This chain must never be interrupted because interruption would lead to dysfunction and failure of the orga- nismic construction. The internal activity is also dependent on the form determi- ning structural order and on the effective suppression of useless motoric deforma- tions by restraining structures. After the chemo-mechanical energy transformation on the macromolecular level the energy is finally utilised in the machinery for the generation of form, locomotion, behaviour, and also for reproduction. EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions The constructional concept of orga- nisation conceives of organismic units as hydraulic entities that are composed of enclosing membranes and flexible integumental walls. The form of every living body must be enforced by tethe- ring and or bandaging structures which suppress the tendency of all living (hydraulic) units to assume a spherical shape. In animal constructions the form is entirely or mainly determined by the tensile force of muscles and by the mesh of connective tissue structures which are organised in highly ordered arrays. In the course of evolution all transforma- tion stages must be shown to comply with the laws of form-enforcement in hydraulic units. The internal construc- tional principles and law-like principles determine the directionality and the irreversibility of evolutionary change. The capability of living construc- tions to function as energy transforming apparatuses and the ability to obtain an input of matter and energy are determi- ned by the structural order of the energy transforming and working construction. All stages of evolutionary transforma- tion series must never lose their structu- ral order and can only undergo gradual transformation in a way that the prece- ding constructional stages open up the organismic options of the subsequent alterations. These alterations lead into constructional alleys with further trans- formation sequences. Evolution which is driven by the activity of the organisms themselves and the insuppressible generation of non-directional variation has to follow very specific internal principles; the step by step alterations must be forced on alleys of ordered constructions and on sequences of non-fortuitous stages. Because the principles and laws of organisation can be elucidated in extant organisms, the transitional sequences can be reliably reconstructed. So models can be formulated which are rational in respect to the methodology applied. Con- sequently they provide valid explanations of the intermediate constructional stages. In all organismic constructions the course of evolution, mainly of the gross morphological level, can be shown to de- pend on the constructional organisation outlined above. Consequently all minor physiologically, cytologically, and histo- logically based functions must be un- derstood as subservient and dependent on the conditions given by the construc- tion as a whole. Morphological features and so called morphological characters in the traditio- nal sense, at all levels of the organisms, must be seen as enforced by mechanical structures and as energy transforming structures. Constructional morphology and the reconstruction of evolutionary transfor- mation do not allow the employment of traditional methodologies which prescribe the dismantling and disintegration of living constructions into morphological patterns or an array of distinct characters. Such a procedure is advocated by cladis- tic “methodologies”. Selection and deli- mitation of features automatically des- troys the coherence of organismic systems and the basic hydraulic constitution. In living machines constructional coherence, the hydraulic properties, the order of form- enforcing structures, and the cataract of energy transforming structures of orga- nisms are not accessible to character analy- sis and comparison of form as advocated by traditional morphology. Only cons- tructional alterations based on the recons- truction of constructionally and functio- nally viable stages are of interest. Superficial aspects of similarity of form and so called homologies in the sense of traditional morphology are also of no rele- vance because constructional explanation has to follow lines of analysis that are com- parable to engineering procedures. Such principles preclude the depic- tion of fortuitous alterations and arbi- trary morphoclines and allow the deter- mination and the reconstruction of the transformation alleys and the establish- ment of the sequences of stages in the course of evolution. All evolutionary alterations must be shown to be delimi- ted by the internal constructional pro- perties of the living machines (Fig. 1). Thus, the constructional alterations of evolution have to be reconstructed as 9 Iberus, 15 (2), 1997 REMOVAL OF WASTE haa METABOLIC === A GENE EXPRESSION AND ACTIVITY GENETIC INTERACTIONS IONIC COMPOSITION ¿== 2 VOLUME REGULATION PROPERTIES OF MEMBRANES (SOLIDITY, PERMEABILITY) CELL SHAPE AND ARRANGEMENT OR TISSUES POLYMERS AND CROSSLINKING PROTEINS ENERGY MECHANICAL PROPERTIES AND ACTIVITIES (VISCOSITY, STIFFNESS, CONTRACTILITY) CYTOSKELETAL PATTERN RYTHM AND EXTRACELLULAR MATRIX AND AUTODEFORMATION ADHESIVE FORCES A PERMANENTLY ACTIVE ORGANISMIC CONSTRUCTION MODULATING PARTIAL SYSTEMS MEA Figure 1. Interrelations between the different levels and parts of organisms and between organisms and their environment (after BEREITER-HAHN, 1991 and EDLINGER, 1995). Figura 1. Interrelaciones entre los diferentes niveles y partes de organismos y entre los organismos y su medio ambiente (tomado de BEREITER-HAHN, 1991 y EDLINGER, 1995). internally guided and directed and cannot be understood as adaptational processes or as ruled by environmental factors in the sense of Darwinian thinking. Evolution in the sense of the FET, which is certainly the most radical variant of post-Darwinian concepts assumes a new status and is based on the properties of the evolving organismic entities. Therefore, all Darwi- nian tenets are excluded from the recons- tructions as they are not conducive to the constructional principles which rule orga- nismic constructions and evolutionary transformation. The theoretical tenets and the met- hodology of the reconstruction procedu- res were elaborated by a group of authors (BONIK ET AL., 1977; GUTMANN, 1974, 1989; EDLINGER, 19894, B, 199la, b, 1992a, b, 1994a, b; EDLINGER ET AL., 1991; VOGEL, 1991) Numerous models provide corroborating evidence for the successful application of the methods and for the validity of the non-Darwi- nian evolutionary theory. On the basis of the just outlined con- cept ontogenetic and phylogenetic deve- lopment of molluscs (like that of other “bauplans” of animals) must be recons- tructed as transformational sequence of organismic constructions over viable intermediate constructional forms. The 54 sequence of evolutionary alterations must be ruled by the intrinsic laws and constructional principles which are res- ponsible for the continuation of energy transformation and motoric activity in all stages. MOLLUSGCS AS A CASE STUDY In the following the evolution of the mollusc constructions is presented as a case study. The stages of mollusc evolu- tion have to be described and figured out as tethered systems which in most cases allow flattening of the foot. In this way they can exert sufficient control of body shape by tethering muscles while the shell structures serve only as defor- mation suppressing elements in the dorsal parts of the coherent whole. THE DERIVATION OF THE MO- LLUSCS: THE KEY TRANSFORMA- TION STEPS In reconstructions based on cons- tructional morphology the morphocli- nes and all aspects of directionality of evolutionary transformation are derived from the biomechanical princi- EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions ples of the living organisms. Thereby, the arrow of time is firmly supported by irreversible constructional explana- tion. On the basis of the constructional analysis only the derivation of molluscs from annelid-like forerunners can be consistently explained (GUTMANN, 1974; EDLINGER, 1991a). The basic mollusc construction must certainly have been a creeping creature with a flattened foot and a dorsal shell or a sequence of shell elements. The problem posed by this basic and only sketchily figured out construction can be formulated in the following way. How can such a constructional constitution arise in skeleton free hydraulic forerunners of whatever kind of primitive metazoan organisation? Which constructional organisation might have been the incipient stage for the emergence of the basic properties? The key changes can only be explai- ned if one sets the starting point of the transformation sequence in an annelid like segmented soft body construction which was capable of controlling the cross-section of the body in a way that creeping on a flattened ventral side became possible. Flattening must have been the first stage of adhesive creeping. The initiation of creeping on a flatte- ned ventral side was a realistic option of annelid-constructions with an internal tethering by dissepimental and other muscles which traversed the cross section of the elongate apparatus. In conjunction with the development of the adhesively creeping foot and its anchorage at the substratum the radula developed in a well demarcated head region. The use of the radula as a rasping organ was dependent on the concomitant formation of the adhesive creeping foot apparatus. This apparatus allowed the radula indirectly to be pressed against the substratum during feeding. Simultaneously the shells could come into existence as dorsal stabilising structures in the non deformed dorsal portions of the body, while an intensifi- cation of motility for peristaltic adhesive creeping occurred on the ventral side. This constructional constitution is in some way still existing in extant flatte- ned chitons. After the establishment of this cons- truction with its segmented sequence of skeletal structures in polyplacophoran- like organisms new paths of evolution were viable. They led to worm-like constructions in a step by step loss of the shell. In this way some recent forms of worm-like molluscs, the Ventroplicata and the Caudofoveata, are easily deriva- ble from chiton-like predecessors. Fusion of the shell-plates occurred in the other major branch. Fusion of the segmented shells, in accordance with specific changes of the soft body led to the formation of the conchiferan cons- truction with its typical unified shell. One consequence of this fusion of the conchiferan shell was that, in con- junction with the formation of the fused shell a narrow waist developed between the cephalopodium and the dorsal body portion while the internal organs were restructured in the frame of a visceral hump. The basic conchiferan organisation is to a certain extent represented in the still extant Tryblidiaceans which display clear metamerism of some organs. The basic Conchiferans provided the cons- tructional basis for a radiative explosion that generated most molluscan cons- truction types of the conchiferan level. The narrowing waist caused and even enforced reductions especially of the of the number of' dorsoventral muscles, gills, and kidneys and the com- pensatory enlargement of the few per- sisting organs. The major steps of transformation are described and depicted in the illus- trations. The complete explanation of the transitions leading to the mollusc constructions cannot be presented here. It was elaborated in a sequence of papers which may be consulted by the interested and critical reader. In the following the major steps are depicted in the illustrations. The ensuing expla- nation is formulated as a caption in respect to the illustration (Fig. 2). It should be stressed that the pictures are no typical morphological illustrations. IS Iberus, 15 (2), 1997 They present visualisations of the array of form enforcing structures and have to be conceived as constructional models. (1.1.) A worm-like predecessor (GUT- MANN, 1974; EDLINGER, 1991a) with me- tameric coelomic pouches, dissepiments, and other segmental tethering elements. Flattening is made possible by dorsoven- tral and other muscles on the dissepiments. All non-longitudinal muscles are helpful in controlling the cross-section and allow flattening of the worm-like body in the in- cipient stages of creeping on hard subs- trate. Metamerism which allowed flattening of the ventral side of the body is manda- tory in the precursor stages of molluscs. Flattening of the body is only possible by internal tethering structures such as mus- cle bearing dissepiments and internal mus- cles transcending the coelomic chambers which were capable of suppressing the tendency to assume a circular shape. Lack of cross sectional tethering as observed in non-metameric worms would result in a circular cross-section of the soft body and preclude even the beginning creeping on hard substrate. In an early stage of evolution the mouth became equipped with chitinous teeth which were located around the sto- modeum and in the foregut. (1.2.) Transition to the basic mollusc construction required the formation of a thick ventral muscle lattice forming a flat and highly deformable foot which was able to follow the contours of the bottom. This enabled the organisms to creep ad- hesively and start rasping at the substrate. In the course of the incipient stages of transformation the coelom and the other inner organs were shifted to the dorsal side. In the dorsal part of the animals co- elomic metamerism was retained and even further demarcated by the newly develo- ping shells. (1.3.) As muscular activity in the form of peristaltic waves travelling over the foot was concentrated in the ventral portion of the pre-mollusc constructions the dorsal parts of the body were held mostly un- 56 deformed. In this situation serial shell ele- ments developed as economising struc- tures; they replaced energetically expen- sive structures such as muscles and con- nective tissue structures. The newly developed skeletal structures served as stabilising elements in the deformable and still worm-like hydraulic apparatus. As the shells formed a sequence of indepen- dent elements bending movements of the body were still possible; they are remnants of the worm-like bending and peristaltic activity of the predecessors. The dorsally situated serial plates were mechanically connected to the foot by strong muscles which continue to demarcate the prece- ding metamerism of the annelid-like pre- decessors. In accordance with the mechanical re- quirements for the flattening of the foot the pairs of muscles were arranged as den- sely spaced pairs of vertical, transverse, and oblique bundles under each plate. These muscles project into the dense mus- cular grid of the foot. Transversely orien- ted muscles cross from one side to the ot- her. In this way the form of the whole ani- mal is under control of the tethering structures. Lateral extensions of the shells and their roof-like position allowed the formation of lateral grooves. In these gro- oves the metameric gills were established as projections of the soft body wall. They were indispensable for the developing mo- lluscs because the adhesion of the foot to the bottom and the covering dorsal shells reduced the body surface usable for res- piration. Only the lateral parts of the soft body continued to be exposed to the su- rrounding medium. The newly develo- ping gills were forced into the old meta- meric constellation because the vessels had to pass between the pre-existing me- tameric muscle bundles. Around the shell a spicule bearing girdle consisting of connective tissue and musculature was established. This girdle served as a protection for the lateral gro- oves with the gills. Pari passu with the rearrangement of the musculature the large coelomic cavi- ties of the annelid like ancestors were res- trained to a narrow dorsal strip. While the posterior part of the coelom was altered EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions El 2 DS e =/=05/2 y Figure 2. The evolution of chiton-like molluscs out of annelid-like metameric ancestors. An early branching into chiton-like and conchiferan constructions occurs. Figura 2. Evolución de los moluscos con forma de quitón a partir de ancestros metaméricos tipo anélido. Tiene lugar una pronta separación entre construcciones conchíferas y con forma de quitón. into the pericardium around the effecti- vely pumping heart the other parts ser- ved as cavities containing the gonads. In conjunction with adhesive creeping the structures around the mouth were per- fected into a radula by shifting of the chi- tinous teeth into a ventral pouch of the fo- regut. The establishment of the radula must be seen in conjunction with the cre- eping mode of locomotion. The scraping, radula could only become effective when the body remained firmly attached to the substrate in a way that exertion of pres- sure by the radula onto the substrate would not push the animal body away from the underlying surface. From this fo- llows that the radula could only develop in strict interdependence with the cree- ping performance of the foot. (1.4.) Reconstructed predecessors of Conchiferan and Polyplacophoran cons- tructions. In the anterior portion of the body the coelom was gradually reduced. The gut formed lateral pouches in the form of the midgut-glands, which, besides their SÍ Iberus, 15 (2), 1997 Figure 3. Evolutionary transformation of chiton-like predecessor into Placophora, Caudofoveata and Ventroplicata. Figura 3. Transformación evolutiva del predecesor tipo quitón en Placophora, Caudofoveata y Ventroplicata. digestive function, served as newly formed fluid filled entities in the frame of the form enforcing structures and as a substantial part of the filling of the inner spaces of the body. The gut pouches were also helpful in holding the gut in its position. As the coelomic space was restrained by the mus- cle construction and receded to narrower dorsal cavities glandular kidney structu- res had to develop as extensions of the metanephridia. They collected excretory material from the extra coelomic spaces mainly in the muscle grid. Such an alte- ration of the metanephridial excretory or- gans was necessitated by the enlargement of the extracoelomic fluid fillings of the muscular grid. The coelom inevitably lost 58 its function as the sole fluid filling unit of the body. There can be no doubt that the evolutionary transformation of the mus- cle apparatus and the restructuring, of the excretory system were coupled and mu- tually dependent. (2.1.) The Polyplacophoran like tran- sitional stage. The model represents an early organisational constellation in mo- lluscan evolution. These forms retained the metameric organisation of muscle system, shell arrangement, gills, and kidneys (Fig. 3). (2.1.1.) The fully developed Polypla- cophoran constructions represent a side EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions Figure 4. The radiation of conchiferan constructions resulting in diverging organismic types which are capable of occupying different environmental conditions. Figura 4. Radiación de construcciones conchíferas que resulta en tipos de organismos divergentes capaces de ocupar diferentes condiciones ambientales. branch of primitive molluscs. Flattening of the shells became more pronounced. The serial shell plates continue to preserve the metameric arrangement of the dorsoven- tral musculature. In the course of flattening, the circulatory and the gill systems were restructured in a way allowing the non- metameric array of the gills and decou- pling of internal metamerism and gill-po- sition. This is unique in Polyplacophorans and not representative for the transition to the molluscs with fused shells. (2.1.2.) Evolution of the Caudofovea- tan construction is part of a radiative differentiation of the polyplacophoran organization. This lineage must have started from polyplacophoran forms (EDLINGER, 1989b) which underwent re- duction of the shells. Concomitantly with shell reduction the lattice-like mus- cle foot was only retained in the anterior portion of the worm like body. The gut was also considerably altered; in the rear portion the intestinal canal was su- rrounded by the midgut-glands, which held the gut in its position. In these bu- rrowing constructions only one poste- rior pair of gills was retained in the hind part of the body. The resulting worm- like form developed independently of the Ventroplicatan constructions. (2.1.3.) Evolution of the Ventroplicatan construction is also a branch of Polypla- cophoran radiation: loss of shell structu- res in conjunction with an enlargement of the girdle resulted in the formation of a worm-like body. The foot-complex per- sisted as a narrow groove running along the ventral side of the whole body length. The groove remains internally tethered by the dorsoventral muscle bundles which are certainly rudiments of the serial shell related retractors. In contrast to the bu- rrowing Caudofoveata the Ventroplicata are capable of climbing in “tree-like” en- vironments. To corroborate the derivation of the worm-like forms from Polyplacop- horans it should be kept in mind that some extant Polyplacophorans as Cryptoplax show an observable tendency to reduce the shells. (2.2.) In the evolutionary alley to the Conchiferan constructions (Fig. 4) the Mo- noplacophorans demarcate a strategic ' intermediate stage. 39 Iberus, 15 (2), 1997 (2.2.1.2.) Figure 5. Transformation of monoplacophoran-like ancestors into scaphopods by extreme dorso- ventral elongation. Figura 5. Transformación de ancestros tipo monoplacóforo en escafópodos mediante una elongación dor- soventral extrema. The unified shell was formed by fu- sion of the segmental skeletal elements. This process ensued in the deepening of the groove between the cephalopodium and the shell-covered visceral hump. The waist became more pronounced and the- reby the cephalopodium was rendered more flexible in respect to the shell. In the Monoplacophora the metameric soft body organization of muscles, gills, nephridia and portions of the coelom is still retained. Metamerism is obviously fading from the rostral portion of the body (GUTMANN, 1974; EDLINGER, 1991a). Noticeable is the concentration of the an- cestral structures, coelom, sac bearing nephridia, and gills in the rear part of the body. This situation is indicative of the transition to the other conchiferan cons- tructions which sprang from a radiative divergence of constructions. (2.2.1.) The fully developed Conchife- ran Constructions. In all conchiferan constructions with fused shells the separation of the two body-portions, the cephalopodium and the dorsal hump, by a waist allows free movement of the foot and the undeformed posture of the visceral sack with the stiff shell frame far from the substrate. The for- mation of the waist enforced the reduction of the number of gill-pairs in all derived conchiferan constructions. 60 (2.2.1.1.) Neopilinida. The Neopilinida are flattened recent Monoplacophoran constructions with clear remnants of metamerism in the arrange- ment of muscles, nephridia and gills. So they must be early representatives of the transition phase to the Conchiferans. Ho- wever, clear indications of reduction of metamerism from the anterior portion of the body become evident; the anterior gills are reduced and the muscle bundles fused (GUTMANN, 1974; EDLINGER, 1991a). (2.2.1.2.) Evolution of Scaphopod cons- tructions. Scaphopods are derived from a Mo- noplacophoran stage with a posterior slit in the shell for the expulsion of faeces. A dorsoventral elongation of the construc- tion caused the reduction of the number of muscles and gill pairs and the forma- tion of a tube-like shell. The foot changes its form by a rearrangement of the inter- nal muscle-lattice and becomes a burro- wing organ. The posterior slit of the shell became an elongate hole. This hole enabled the animals to ventilate their mantle cavity when they penetrated into the substrate. The water is sucked in from the anterior region and expelled through the poste- rior hole. The total reduction of the gills is enforced by the narrowing of the man- tle cavity when the elongate organisms developed (EDLINGER, 1991b) (Fig. 5). EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions (2.2.1.3.) Figure 6. The evolution of the gastropod construction by torsion. Torsion occurs after the forma- tion of a waist between the cephalopodium and the visceral hump and reduction of most of doso- ventral muscles. Figura 6. La evolución de la estructura de un gasterópodo mediante torsión. Ésta ocurre tras la forma- ción de un estrechamiento entre el cefalopodio y el asa visceral, y la reducción de la mayoría de los mús- culos dorsoventrales. (2.2.1.3.) Evolution of Gastropods (ED- ceral hump after a very narrow waist had LINGER, 1988a, 1988b, 1989a). formed and most of the gill pairs had been The most significant event of gastro- reduced. Torsion led to an anterior posi- pod evolution was the torsion of the vis- tion of the partially reduced mantle ca- 61 Iberus, 15 (2), 1997 (2.2.1.4.) (2.2.1.4.2.) Figure 7. The evolution of cephalopods characterized by the appearance of gas-filled spaces at the top of the shell and by a radical rearrangement of the musculature of the cephalopodium with strong muscular arms and suckers. Figura 7. Evolución de los cefalópodos, caracterizada por la aparición de cámaras llenas de gas en la parte alta de la concha y una redistribución radical de la musculatura del cefalopodio con fuertes ramas mus- culares y ventosas. vity with the remaining gills and the for- mation of the chiasma of the lateral nerve cords. The precondition for torsion lies in a step by step reduction of dorsoventral muscle bundles of a Monoplacophoran ancestor and by narrowing of the waist between the cephalopodium and the vis- ceral hump. This process is connected with a step by step spiralling up of the shell. The persistence of only one pair of crossing obliquely transversal muscles could cause the torsion. Totally bilaterally organised symmetric Bellerophontacean shells are representative of this process (Fig. 6). (2.2.1.3.1) Radiation of Gastropod cons- tructions: All gastropod lineages are de- rived from a Bellerophontacean stage with 62 a bilateral and helical shell, which posses- sed slits or a series of holes in the poste- rior part of the shell for the expulsion of faeces. Only one pair of parallel dorso- ventral muscles persisted. Asymmetry could arise by the partial or total reduction of one of the dorsoventral muscles and the change of the planispiral shell of Be- llerophon-like ancestors to an asymme- tric helical form. In most cases the gills be- came also unequal or one gill was entirely reduced (EDLINGER, 1988a, 1988b, 19894a). Flattening and secondary reduction of the coil can result in the formation of a cup- like shell. Flattening must cause a change in the muscle arrangement. The dorso- ventral muscles and their scares on the inner of the shells were enlarged to form EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions Figure 8. The evolution of bivalves by a rearrangement of the dorsoventral muscles and a division of the shell to a bivalve one. Figura 8. Evolución de los bivalvos por medio de una redistribución de los músculos dorsoventrales y una división de la concha hacia una concha bivalva. horseshoe-like insertions. The gills can persist as in Fissurella. In this case the re- tractor muscle will form a homogeneous horse-shoe-like entity. If the gills are also reduced and substituted by secondary gill-like structures the muscles are splitted up into a Monoplacophoran-like situation of Patella (EDLINGER, 1988a, 1988b, 1989a; HASZPRUNAR, 1988). (2.2.1.4.) Evolution of Cephalopod constructions (BONIK, GRASSHOFE, GUT- MANN AND KLEIN-RÓDDER, 1977). The de- velopment of gas-filled chambers in the apical part of the shell rendered the orga- nisms more buoyant. The foot with its de- formability and the adhesive capabilities was altered into a system of muscular arms with suckers which could grasp prey. Retraction of the soft-body into the shell inevitably caused expulsion of a water- current. In the course of evolution im- provement of this mechanisms was alte- red into effective jet propulsion when the hind part of the foot was modified to form a narrow funnel through which the water expelled from the mantle cavity was con- centrated to enhance the jet effect. This constructional situation can be observed in all cephalopods (Fig. 7). An early bi- furcation or diphyletic development gave rise to the branches of endocochlean and exocochlean Cephalopod construction (2.2.1.4.1, 2.2.1.4.2). (2.2.1.5.) Evolution of Bivalve Cons- tructions (VOGEL AND GUTMANN, 1980). The origin of Bivalves can be understood as a rearrangement of the dorsoventral muscles in a high chambered Monopla- cophoran ancestor. A considerable por- tion of the dorsoventrally and obliquely arranged muscles in the foot were shif- ted into a horizontal position connecting not the shell and the foot as in the former stages but the two laterally bent down plates of an evolving bivalved shell. Contraction of the muscles brought the flanks of the shells together in protective behaviour. Most of the vertical (dorso- ventral) portions of the musculature re- tained their original retractile function in respect to the foot but the number of re- tractors was reduced. Consequently most of the gills disappeared with one pair remaining. The remaining pair of gills are utilised as filterfeeding devices because lateral cephalic lobes bridged the gap between the mouth and the gills and formed conveyer belts for the transport of food from the gills to the mouth (Fig. 8). 63 Iberus, 15 (2), 1997 DISCUSSION AND CONCLUSION Itis very obvious that in the perspec- tive of constructional morphology and organismic evolution character-analysis and the establishment of sequences of organismic forms in the sense of homo- logies are inconclusive and arbitrary. They pose problems but do not provide answers because description of form and character analysis do not lead to cons- tructional insight or an understanding of form determination. Constructional alte- ration in evolution can not reasonably be derived from genetic or other molecular features (GHISELIN, 1988; WAGELE, 1994; WAGELE AND WETZEL, 1994; EDLINGER, 1995). The understanding of organisms as constructions leaves no doubt that the constructional configuration determines evolutionary change of the living machi- nery and prescribes the sequence of constructional stages and of irreversible steps. There is little freedom for contin- gency in the order of the “bauplan” con- figurations and no encouragement for simple description and form compari- son. Nothing useful can come from such traditional approaches which are blind to causal aspects and insensitive for ex- planatory principles. When organisms are conceived as energy transforming constructions evo- lution must be ruled by constructional principles and not by subjective “gestalt” properties or subservient molecular me- chanisms. As could be shown in the fore- going context strict obeisance of the cons- tructional principles allows the rejection of constructionally impossible or impro- bable alternatives. From the methodology applied and the mode of reconstruction of the evolutionary transformations just ad- vocated follows that traditional phyloge- netic concepts, form sequences, and cla- dograms based on usual procedures are not considered valid. Therefore, traditio- nal hypotheses are bypassed and left out of consideration when the criteria of the Frankfurt-Theory are applied. If some- body feels the need to adhere to the idea that Plathelminths, non-segmented worms, or Nemerteans might be the pre- cursors of mollusc he or she should pre- 64 sent a continuous model with a strict ex- planation of the intermediate construc- tional steps. The genes and other molecular me- chanisms which are subservient in relation to the living constructions and their struc- tures have to comply with the construc- tional requirements and must follow cons- tructional modifications of the machinery in the course of structural reorganisation. Taken as separate features molecular and physiological mechanisms are not useful for the elucidation of constructional change in evolution. Therefore, constructional morphology neglects all traditional sug- gestions as to the affinity of organismic groups based on form similarities in the sense of homologies and on subservient and constructionally dependent molecu- lar and physiological properties. Itis not possible to give a list of all the published sequences of forms which were figured out to represent what all the aut- hors tried to suggest as stages of mollusk evolution (SCHELTEMA, 1978; GÓTTING, 1980a, 1980b; HAas, 1981; BANDEL, 1983; LAUTERBACH, 1983a, b; HASZPRUNAR, 1988, 1992a, b). Many contributions were for- mulated by paleontologists (RUNNEGAR AND POJETA, 1974; RUNNEGAR, POJETA, NOEL, TAYLOR, TAYLOR AND MCCLUNG, 1975; MAREK AND YOCHELSON, 1976; Yo- CHELSON, FLOWER AND WEBERS, 1973; Po- JETAJR., 1987) who started from one or the other fossilstructure. Most morphoclines presented since the last century are highly contradictory, however, all of them have in common an access that arbitrarily selects some features of skeletal or soft body struc- tures. Organisms as constructionally res- trained and not deliberately transforma- ble entities are not even expected. In none of the published morphoclines is the di- rectionality of evolutionary transforma- tion elaborated or explained as irreversi- ble. Organisms are misconceived as sha- ped and freely modifiable pieces of art. Fruitful discussion can only start when alternative models with continuous ex- planations and well supported transfor- mational polarities are given. In the pre- sent situation the discussion of all the morphoclines would require the sacrifi- EDLINGER AND GUTMANN: Phylogeny of molluscs as evolving constructions cium intellectus. 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AND WETZEL, R., 1994. Nucleic acid sequence data are not per se reliable for inference of phylogenies. Journal of Natural History, 94: 749-761. YOCHELSON, E. L., FLOWER, R. H. AND WEBERS, G. F., 1973. The bearing of the new Late Cam- brian genus Knightoconus (Mollusca: Mono- placophora) upon the origin of the Cepha- lopoda. Lethaia, 6: 275-310. O Sociedad Española de Malacología Iberus, 15 (2): 67-74, 1997 Mollusc fauna of the medium high mountain ranges of the Hungarian Holocene: a zoogeographical research Fauna de moluscos de media y alta montaña del Holoceno de Hun- gría: una investigación zoogeográfica Levente FÚKOH* Recibido el 30-X-1995. Aceptado el 15-11-1997 ABSTRACT An attempt is made to complete the zoogeographical studies of the mollusc fauna of the medium-high mountain ranges of the Hungarian Holocene by analysing twenty-eight chrono- logically and biostratigraphically known faunae. Eighty four species are classified in nine fau- nalcentres and four biozones (Vallonia costata, Clausillidae, Granaria frumentum and Helici- gonia faustina — Acicula polita). A brief discussion is made on the abundance of several species of each faunal-centre. The picture drawn from the fauna agrees with the geographical position and geomorphological conditions of Hungary (Carpatian Basin, Central-Europe) RESUMEN Se pretende completar los estudios zoogeográficos de la fauna de moluscos de media y alta montaña del Holoceno de Hungría mediante el análisis de ventiocho faunas conocidas tanto cronológica como estratigráficamente. Ochenta y cuatro especies fueron clasificadas en nueve “centros faunísticos” y cuatro biozonas (Vallonia costata, Clausillidae, Granaria fru- mentum and Helicigonia faustina — Acicula polita). Se hace una breve discusión sobre la abundancia de varias especies en cada “centro faunístico”. La representación que se obtiene de la fauna coincide con la posición geográfica y las condiciones geomorfológicas de Hun- gría (planicie de los Cárpatos, Europa central). KEY WORDS: Molluscs, Zoogeography, Holocene, Hungary. PALABRAS CLAVE: Moluscos, Zoogeografía, Holoceno, Hungría. INTRODUCTION The paleoecological and biostratigrap- hical studies of the mollusc fauna of the medium-high mountain ranges of the Hun- garian Holocene (Fig. 1) has been comple- ted in the last few years. (FUKOH, 1991, 1992a, 1992b, 1993a, 1993b). Although during these estudies, zoogeographical examinations were carried out for certain faunae (FUKOH, 1983, 1989; BABA AND FUKOH, 1984), a comprehensive view is lacking. The aim of this paper is to complete the lack of information about this subject. The analysis of twenty-two chronolo- gically and biostratigraphically known faunae (FUKOH, KROLOPP AND SÚMEGI, 1995) is completed in this work (Fig. 2), in- creasing the number of species previously cited from 81 (Table 1) to 84 (Table III). * Mátra Museum, H-3200 Gyóngyós, Kossuth u. 40, Hungary. 67 Iberus, 15 (2), 1997 SS 8 MEM(12%) MA(4%) AMe(1%) PM(30%) Figure 1. Map showing the position of the localities of the Biozones stratotypes and the zoogeo- graphical distribution of the Holocene mollusc fauna in the Hungarian medium high mountain area. A: Búkk Mts.; B: Aggtelek karst; C: Bakony Mts. Faunal centres, PM: Ponto-mediterranean; SA: Siberian-Asiatic; MEM: Middle-European-Mountain; HM: Holomediterranean; AM: Adriato- Mediterranean; EM: European-Mountain; MA: Middle-Asiatic; CS: Caspi-Sarmatian; AMe: Atlanto-Mediterranean. Biozones, 1: Vallonía costata biozone; 2: Clausiliidae biozone; 3: Granaria frumentum biozone; 4: Helicigona faustina — Acicula polita biozone. Figura 1. Mapa mostrando la posición de las localidades de los estratotipos de las biozonas y la distribu- ción zoogeográfica de la fauna de moluscos de media y alta montaña del Holoceno de Hungría. A: Montes Búkk; B: karst de Aggtelek; C: Montes Bakony. “Centros faunísticos”, PM: ponto-mediterráneo; SA: sibe- riano asiático; MEM: media montaña europea; HM: holomediterráneo; AM: adriatico-mediterráneo; EM: montaña europea; MA: medio asiático; CS: caspiano-samartiano; AMe: atlanto-mediterráneo. Biozonas, 1: biozona de Vallonia costata; 2: biozona de Clausiliidae; 3: biozona de Granaria frumen- tum; 4: biozona de Helicigona faustina — Acicula polita. MATERIAL AND METHODS distribution of the species. This metho- dology was chosen because, according to the preliminary examinations and calcu- lations, it is more suitable when resear- ching fossil materials. The methodology used to carry out the classification presented in this paper is the same employed to situate zoogeographi- cally the species of the Middle-European faunae. These methods can be divided into two main groups as follows: 1. Methods based on recent distribu- tion of the species (KERNEY, CAMERON AND JUNGBLUTH, 1983; FLASAR, 1971; KÓRNIG, 1983; ALEXANDROWITZ, 1983, 1984; FRANK, 1988, 1990, 1992a, 1992b) 2. Derivative method (BABA, 1982). This method is based on recent and fossil 68 RESULTS AND CONCLUSSIONS Table II shows the distribution in num- ber of the 84 mollusc species found in the studied area by faunal-centres and biozo- nes. Figure 3 illustrates the zoogeograp- hical distribution in number of the Holo- cene and recent mollusc species. FEÚKOH: Mollusc fauna of the medium high mountain ranges of the Hungarian Holocene Boreal Atlantic Sub-Boreal Sub-Atlantic 8 7 6 5 4 3 2 1 0 1000 radiocarbon year BP. (1950) Vallonia costata Clausiliidae Granaria frumentum Helicigona faustina- zone zone zone Acicula polita zone Muflon 1/9-6 Mutflon 1/5-2 | Muflon 1/1 Muflon 11/3-2 Kajla-bérc 3 Kolyuk 11/17-12 Kajla-bérc 2-1 | Kólyuk 11/10-1 Rejtek 1/11 Rejtek 1/II Szentléleki-v. Horvatí-h. 4-3 Szalajka 11 Szalajka 5 Szalajka 3a Szalajka 3b-1 Monosbél Petényi H5 Petényi H3 Kis-kohát 4 Csunya-v 1/2 Csunya-v 111 Nagyoldal 6 Nagyoldal 5-4 Szentgál Háromágú Rigó-h Baradla Figure 2. Biostratigraphic identification of significant terrestric Holocene localities of Hungary (twenty- eight chronologically and biostratigraphically known faunae). Biúikk Mts.: Muflon”-cave, “Kajla- bérc”-cave, “Kólyuk IT”-cave, “Rejtek ”-cave, “Horvéti”-hole, “Petényi”-cave, “Csunya”-valley rock shelter, “Csunya”-valley I=rock shelter, “Háromágu”-cave, “Kajla-bérc”-cave, “Szalajka”-valley-tra- vertine, “Monosbél”-travertine, “Szentléleki”-valley-rock shelter, “Kálmán-rét”-shaft-cave, “Kiskóhát”- shaft-cave. Aggtelek karst: “Baradla”-cave, “Nagy-oldal”-shaft-cave. Bakony Mts: Szentgál, Mecsek- hill, “K6-lik”-cave, “Rigo”-hole. Lines represent the biostratigraphic and chronologic extension and/or connection of localities (shown in Figure 1). Figura 2. Identificación bivestratigráfica de las localidades terrestres más significantes del Holoceno de Hungría (ventiocho faunas conocidas cronológica y bivestratigráficamente. Montes Búkk: cueva “Muflon”, cueva “Kajla-bérc”, cueva “Kólyuk II”, cueva “Rejtek I”, sima “Horvéti”, cueva “Petényi”, abrigo rocoso “Csunya”-valley L, abrigo rocoso “Csunya”-valley IL, cueva “Háromágu”, cueva “Kajla-bérc”, valle “Sza- lajka” travertino, “Monosbél” travertino, valle “Szentléleki” abrigo rocoso, pozo “Kálmán-rét”, pozo “Kiskóhát”. Karst Aggtelek: cueva “Baradla”, pozo “Nagy-oldal”. Montes Bakony: Szentgál, Mecsek- hill, cueva “Kó-lik”, sima “Rigo”. Las líneas representan la extensión y/o conexión bioestratigráfica y cro- nológica de las localidades (mostradas en la Figura 1). 69 Iberus, 15 (2), 1997 Table I: Distribution of 81 molluscs species found the medium-high mountain ranges of the Hungarian Holocene fauna, according to several authors. Authors, 1: Verney, Cameron and Jungbluth; 2: Flasar; 3: Alexandrowicz; 4: Kórnig; 5: Frank; 6: Bába. Abbreviations, a: Alpian; adm: Adriatic-Mediterranean; e: European; h: Holarctic; hm: Holomediterranean; k: Carpatian; ksz: Capian-Sarmatian; m: middle; ma: Middle-Asiatic; med: Mediterranean; merid: meridional; p: Palearctic; po: Pontomediterranean; s, n, o, w, the four cardinal points; sza: Siberian-Asiatic; ws: West-Sibiric; sz: Siberian. Tabla 1. Distribución de 81 especies de moluscos encontradas en media y alta montaña de la fauna del Holoceno de Hungría, de acuerdo con distintos autores. Autores, 1: Verney, Cameron and Jungbluth; 2: Flasar; 3: Alexandrowicz; 4: Kórnig; 5: Frank; 6: Bába. Abreviaturas, a: alpino; adm: adriatico-medi- terráneo; e: europeo; h: holártico; hm: holomediterráneo; k: carpatiano; ksz: capiano-samartiano; m: medio; ma: medioasiático; med: mediterráneo; merid: meridional; p: paleártico; po: pontomediterráneo; s, n, o, w: los cuatro puntos cardinales; sza: siberiano-asíatico; ws: sibírico oeste; sz: siberiano. Species 1 2 3 4 5 6 Achantinula aculeata (0. F. Miller, 1774) wp WSZ Acicula polita (Harimann, 1840) e-0 me me me me-a po Aegopinella minor (Stabile, 1864) some me-se soe some po Aegopinella pura (Alder, 1830) e e e e WSZ Bradybaena fruticum (0. F. Mller, 1774) moe-a e e-a 05Z Bulgarica vetusta (Rossmíssler, 1836) s08 po Carychium minimum (0. E. Múller, 1774) e-S7 e-sz e e-S7 057 Carychium tridentatum (Risso, 1826) e se e e hm Cecilioides acicula (0. E. Múller, 1774) med-we wme Cepea vindobonensis (Férussac, 1821) soe soe soe ksz Chondrina clienta (Westwrlund, 1883) s08-0 s08-0 08-a po Chondrula tridens (0. E. Múller, 1774) msoe pm msoe hm Clausilia cruciata (Studer, 1820) ne-a ba b-a moe-a e Clausilia dubia Draparnaud, 1805 me me me me me po Clousilia pumila C. Pfeiffer, 1828 moe me oe ome moe po Cochlicopa lubrica (0. E. Miller, 1774) h h ws h h h Cochlicopa lubricella (Porro, 1837) h h h h ma Cochlodina cerata (Rossm(ssler, 1836) k wk me Cochlodina laminata (Montagu, 1803) e e ws e e me Cochlodina orthostoma (Menke, 1830) moe 0€ me Columella edentula (Draparnaud, 1801) h h h 0-Sz Daudebardia brevipes (Draparnaud, 1805) mse mse po Daudebardia helenae Fúkúh, 1985 me Daudebardia rufa (Draparnaud, 1805) mse med-me sme po Discus perspectivus (Múhlfeld, 1816) a-ok a-ok a-ok po Discus rotundatus (0. E. Miller, 1774) wme wsm WS wme adm Discus ruderatus (Férussac, 1821) p p p p 0-52 Ena montana (Draparnaud, 1801) me-k-a me me me-k e Ena obscura (0. E. Múller, 1774) e e e hm Euconulus fulvus (0. E. Miller, 1774) h h h h h Evomphalia strigella (Draparnaud, 1801) me ome ome ome Ksz Granaria frumentum (Draparnaud, 1801) med merid moe po Helicella obvia (Menke, 1828) s08 smoe Helicigona faustina (Rossmíssler, 1838) k me Helicodonta obvoluta (0. E. Miller, 1774) me merid merid sme adm 7O EÚKOH: Mollusc fauna of the medium high mountain ranges of the Hungarian Holocene Table 1. (Continuation). Tabla I. (Continuación). Species 1 2 3 4 5 6 Helicopsis striata (0. E. Miller, 1774) wmoe e Helix pomatia Linné, 1758 smoe soe me soe msoe po Isognomostoma isognomostoma (Schróter, 1784) a-k me a-k a-k a-k me Laciniaria biplicata (Montagu, 1803) me-b some po Laciniaria plicata (Draparnaud, 1801) moe me me moe po Macrogastra latestriata (A. Schmidt, 1857) k Macrogastra plicatula (Draparnaud, 1801) me e e e po Macrogastra ventricosa (Draparnaud, 1801) me e me e e po Monacha cartusiana (0. E. Múller, 1774) po Nesovitrea hammonis (Siróm, 1765) p p h 0-SZ Orcula doliolum (Bruguiére, 1792) s0€ me merid soe ma Orcula dolium (Draparnaud, 1801) a-k a-wk a-k po Oxychilus depressus (Sterki, 1880) a-k me a-k po Oxychilus draparnaudi (Beck, 1837) med-we we Oxychilus glaber (Rossm(ssler, 1838) sme smoe so€ a-se po Oxychilus inopinatus (Ulicny, 1887) k Oxychilus orientalis (Clessin, 1887) k k me Perforatella incarnata (0. F. Miller, 1774) msoe mwe me msoe po Perforatella vicina (Rossm(ssler, 1842) e Punctum pygmaeum (Draparnaud, 1801) h p p h h Pupilla muscorum (Linné, 1758) h h 0-57 Pupilla triplicata (Studer, 1820) 08-0 e-a adm Pyramidula rupestris (Draparnaud, 1801) we-M oe m-a m-a ma Ruthenica filograna (Rossm(ssler, 1836) 0€ oe moe moe me Semilimax kotulai (Westerlund, 1871) a-k a-k a-k Semilimax semilimax (Férussac, 1802) a-me a-me Trichia unidentata (Draparnaud, 1805) a-k oa-wk oa-wk oa-k me Trichia hispida (Linné, 1785) e e Truncatellina claustralis (Gredler, 1856) m-sa m m hm Truncatellina cylindrica (Ferrussac, 1807) se se se se hm Vallonia costata (0. E. Múller, 1774) h h h h h h Vallonia pulchella (0. F. Múller, 1774) h h h h h h Vallonia enniensis (Gredler, 1856) mse me Vertigo alpestris Alder, 1838 na p 0-Sz Vertigo angustior Jeffreys, 1830 e e ksz Vertigo parcedentata (A. Braun, 1847) sz0 Vertigo antivertigo (Draparnaud, 1801) p ksz Vertigo pusilla (0. E. Múller, 1774) e e e-was hm Vertigo pygmaea (Draparnaud, 1801) h h h WSZ Vertigo substriata (Jeffreys, 1833) a 0-e Vitrea contracta (Westwrlund, 1871) p hm Vitrea crystallina (0. F. Miller, 1774) e e ws e e adm Vitrea diaphana (Studer, 1820) a-k a-merid a-me po Vitrina pellucida (0. E Múller, 1774) h p p p h Zebrina detrita (0. F. Múller, 1774) s08 po Zonitoides nitidus (0. E Miller, 1774) h h 71 Iberus, 15 (2), 1997 Table II. Distribution in number of 84 mollusc species in the Hungarian Holocene fauna of the medium-high mountain ranges by Faunal-centres and Biozones. Abbreviations, 1: Vallonia costata biozone; 2: Clausiliidae biozone; 3: Granaria frumentum biozone; 4: Helicigona faustina — Acicula polita biozone. Tabla 1. Distribución en número de las 84 especies de moluscos de la fauna del Holoceno de Hungría en media y alta montaña agrupadas por “centros faunísticos” y biozonas. Abreviaturas, 1: biozona de Vallonia costata; 2: biozona de Clausiliidae; 3: biozona de Granaria frumentum; 4: biozona de Helicigona faustina — Acicula polita. BIOZONES FAUNAL-CENTRES No. of species 1 2 3 4 Siberian-Asiatic 21 15 19 16 16 Middle-Asiatic 3 3 2 7 2 Caspi-Sarmatian a) 3 4 2 3 Pontomediterranean 2 18 19 18 22 Adriato-Mediterranean 5 5 4 4 5 Atlanto-Mediterranean 1 0 1 0 0 Holomediterranean o o) 5 5 5 European -Mountain 7 4 3 2 4 Middle-Eur. -Mountain 11 8 o 6 9 TOTAL 84 62 63 5 66 30 Y) o Number of species a Faunal centres Figure 3. Zoogeographical distribution in number of the mollusc fauna of the Hungarian medium high mountain ranges considering Holocene and Recent species separately. 1: Pontomediterranean; 2: Siberian-Asiatic; 3: Middle-European-Mountain; 4: Holomediterranean; 5: Adriato- Mediterranean; 6: European-Mountain; 7: Middle-Asiatic; 8: Caspi-Sarmatian; 9: Atlanto- Mediterranean. Figura 2. Distribución zoogeográfica en número de la fauna de moluscos de media y alta montaña de Hungría, considerando tanto las especies del Holoceno como las recientes. 1: pontomediterráneo; 2: sibe- riano asiático; 3: media montaña europea; 4: holomediterráneo; 5: adriático mediterráneo; 6: montaña europeo; 7: medio asiático; 8: caspiano-samartiano; 9: atlanto-mediterráneo. 2 FÚKOH: Mollusc fauna of the medium high mountain ranges of the Hungarian Holocene Table IM. Zoogeographical distribution of the Hungarian Holocene species by faunal-centres. (*) Molluscan species not cited in Table 1. Tabla 111. Distribución zoogeográfica de las especies del Holoceno de Hungría por “centros faunásticos”. (%) Especies de moluscos no citadas en la Tabla 1. 1. Siberian-Asiatic faunal-centres 21species Achantinula aculeata Euconulus fulvus Vallonia pulchella Aegopinella pura Limax maximus Linné, 1758 (*) Vertigo alpestris Bradybaena fruticum Nesovitrea hammonis Vertigo parcedentata Carychium minimum Punctum pygmaeum Vertigo pusilla Cochlicopa lubrica Pupilla muscorum Vertigo pygmaea Columella edentula Vallonia costata Vitrina pellucida Discus ruderatus Vallonia enniensis Zonitoides nitidus 2. Middle-Asiatic faunal centres 3 species Cochlicopa lubricella Orcula doliolum Pyramidula rupestris 3. Caspian-Sarmatian faunal-centres 5 species Cepaea vindobonensis Semilimax kotulai Vertigo antivertigo Evomphalio strigella Vertigo angustior 4. Ponto-Mediterranean faunal-centres 25 species Acicula polita Granaria frumentum Macrogastra ventricosa Aegopinella minor Helix pomatia Monacha cartusiana Bulgarica vetusta Helicella obvia Orcula dolium Chondrina clienta Zebrina detrita Oxychilus depressus Clausilia dubia Helicopsis striata Oxychilus glaber Clousilia pumila Laciniaria plicata Perforatella incarnata Doudebardia brevipes Laciniaria biplicata Vitrea diaphana Daudebardia ruta Macrogastra plicatula Vitrea subrimata (Reinhardt, 1871)(*) Discus perspectivus 5. Adriatic-Mediterranean faunal-centres 5 species Cecilioides acicula Helicodonta obvoluta Vitrea crystallina Discus rotundatus Pupilla triplicata 6. Atlantic-Mediterranean faunal-centres 1 species Semilimax semilimax 7. Holomediterranean faunal-centres 6 species Corychium tridentatum Ena obscura Truncatellina cylindrica Chondrula tridens Truncotellina claustralis Vitrea contracta 8. European-Mountain faunal-centres 7 species Clausilia cruciata Oxychilus inopinatus Trichia hispida Ena montana Perforatella vicina Vertigo substriata Macrogastra latestriata 9. Middle-European-Mountain faunal-centres 11 species Cochlodina orthostoma Helicigona faustina Oxychilus draparnaudi Cochlodina cerata Trichia unidentata Oxychilus orientalis Cochlodina laminata Isognomostoma isognomostoma Ruthenica filograna Daudebardia helenae Laciniaria turgida (Rossmassler, 1836) (*) 73 Iberus, 15 (2), 1997 The zoogeographical distribution of species by Faunal-centres is summarised in Table II. The formation of the characteristic zoogeographical conditions of the recent fauna began after the last cold period of the Pleistocene. This can be stated on the basis of the relative abundance analyses of the faunae (84 species) situated in nine faunal-centres (Tables II and III). Prima- rily, the abundance of Subatlantic species has increased during the last ten thou- sand years. Ponto-Mediterranean species are the most important and can be obser- ved almost in all every biostratigraphical REFERENCES ALEXANDROWICZ, S. W., 1983. Malacofauna of Holocene calcareous sediments of the Cracow Upland. Acta Geologica Polonia, 33: 117-158. ALEXANDROWICZ, S. W., 1984. Srodkowoholo- censka Malakofauna z Harcygrundu Kolo Czorsztyna (Pieninski Pas Skalkowy). Studia Geologica Polonia, 83: 96-114. BABA, K., 1982. Eine neue zoogeographische Gruppierung der Ungarische Landmollusken und die Wertung des Faunabildes. Malaco- logica, 22: 441-454. BABA, K. and FUKÓH, L., 1984. Holocén és re- cens malakológiai adatok értékelése állatfol- drajzi és 0kológiai módszerekkel a Búukkben. Malakologiai Tájékoztató (Malacological News- letter), 4: 42-53. FLASAR, I., 1971. Zur Malakofauna des nordós- tlichen Erzgebirges und des angerenzenden Gebirgsvorlands. Malakologische Abhandlun- gen, 3: 135-170. FRANK, C., 1988. Aquatische und Terrestrische Mollusken der Osterreichischen Donau-auen- gebiete und der Angrenzenden Biotope. Teil XIL. Das Osterreichische Donautal von der Os- terreichisch-Deutschen Staatsgrenze bis Linz. Linzer biologische Beitrag, 20: 413-503. FRANK, C., 1990. Pleistozáane und Holozáne Mo- lluskenfaunen aus Stillfried an der March: Ein Beitrag zur Ausgrabungs-geschichte von Stillfried und des Buhubergs nordlich von Stillfried. Wissenschaftlichen. Mitteilung Nie- derosterreichische Landesmus, 7: 7-272. FRANK, C., 1992a. Spát-und postglaciale Gas- tropoden aus dem Nixloch bei Losenstein- Ternberg (Oberósterreich). Verbindung Os- terreichisch Akademie der Wissenschaften, Mit- teilung Kommission Quartarforschung, 8: 35-69. FRANK, C., 1992b. Malakologisches aus dem Ostalpenraum. Linzer biologische Beitrag, 24: 383-662. 74 zones (Vallonía costata biozone 18; Clausi- liidae biozone 19; Granaría frumentum bio- zone 18; Helecigona faustina-Acicula polita biozone 22). Species of the Siberian-Asia- tic faunal-centre follow them in impor- tance (Vallonia costata biozone 15; Clausi- Itidae biozone 19; Granaria frumentum bio- zone 16; Helicigona faustina-Acicula polita biozone 16). The Middle-European- Mountain faunal-centres are located in third place considering the relative abundance of the species (Vallonia costata biozone 8; Clausiliidae biozone 6; Granaria frumentum biozone 6; Helicigona faustina- Acicula polita biozone 9). FUKOH, L., 1983. A búkki holocén Molluscák állatfóldrajzi csoportositása (The Zoogeo- graphical Grouppirung of the Holocene Mo- lluscs in the Búkk). Malakologiai Tájekoztató (Malacological Newsletter), 3: 37-39. FUKOH, L., 1989. A Szilvasvárad: Szalajka-vólgy (BNP) mésztufa úledekeinek malakosztrati- gráfiai vizsgálata. Folia Historico-Naturalia Musei Matraensis, 14: 39-42. FUKOH, L., 1991. Examinations on Faunal-his- tory of the Hungarian Holocene Mollusc fauna (Characterization of the Succession Phase). Folia Historico-Naturalia Musei Ma- traensis, 16: 13-28. FUKÓH, L., 1992a. Holocene Malacology in Hun- gary. In Giusti, F. and Manganelli, G. (Eds.): Abstracts of the eleventh International Malaco- logical Congress. University of Siena: 115. FuKOH, L., 1992b. The Holocene Mollusc fauna of the Búkk Mountains. Abstracta Botanica, 16(2): 101-108. FUKOH, L., 1993a. Holocene malacofaunal as- semblages in Hungary. Scripta Geologica., Spec. Issue, 2: 121-125. FUKOH, L., 1993b. Main features of the deve- lopment of the Hungarian Holocene Mollusc fauna. Malakológiai Tájékoztató (Malacological Newsletter), 12: 15-19. FUKOÓH, L., KROLOPP, E. AND SUMEGI, P., 1995. Quaternary malacostratigraphy in Hungary. Malacological Newsletter. Suppl., 1: 113-198. KERNEY, M. P., CAMERON, R. A. D. AND JUNG- BLUTH, J. H., 1983. Die Landschnecken Nord-und Mitteleuropas. P. P. Verlag Berlin: 63-284. KORNIG, G., 1983. Studie zur Gastropodenfauna der Westkarpatischen Kalk-und Karstsch- luchten. Malakologische Abbhandlungen, 8: 131- 142. O Sociedad Española de Malacología ——_—_—_—T— lIberus, 15 (2): 75-82, 1997 Aestivation responses of three populations of the giant Afri- can snail, Achatina achatina Linne (Gastropoda: Achatinidae) Respuestas a la estivación de tres poblaciones del caracol gigante afri- cano Achatina achatina Linne (Gastropoda: Achatinidae) Joseph R. COBBINAH* Recibido el 9-X-1995. Aceptado el 7-1V-1997 ABSTRACT The rates of aestivation and emergence from aestivation of three experimental populations of the giant African snails Achatina achatina (Linne, 1758) were compared. The snails were from three origins: Donyina in the Ashanti Region, Nkasem in the Brong Ahafo Region and Apedwa in the Eastern Region of Ghana. The shortest aestivation period [4 weeks) was re- corded for the Apedwa snails while the longest period (16 weeks) was recorded for the Don- yina population. Data for pre-aestivation and post-aestivation growth rates show a decrea- sing order: Apedwa > Nkasem > Donyina. The mean growth rates eight weeks before aesti- vation were 3.6 g, 14.3 g and 19.2 g for the Donyina, Nkasem and Apedwa snails respectively and differed significantly (P = 0.001). The variability in growth rates and dura- tion of aestivation reflects the optimal sizes of the natural population of the three groups. RESUMEN Se comparan las tasas de estivación y de abandono de la misma de tres poblaciones ex- perimentales del caracol gigante africano Achatina achatina (Linné, 1758). Los caracoles eran de tres localidades diferentes: Donyina, en la región de Ashanti; Nkasem, en la re- gión de Brong Ahafo, y Apedwa, al Este de Ghana. El periodo de estivación más corto (4 semanas) se registró en la población de Apedwa, mientras que el periodo más largo (16 semanas) se registró en la población de Donyina. Los datos de las tasas de crecimiento en la pre-estivación y la post-estivación muestran un orden decreciente: Apedwa > Nkasem > Donyina. Las tasas medias de crecimiento ocho semanas antes de la estivación eran 3,6 9, 14,3 g y 19,2 g para los caracoles de Donyina, Nkasem y Apedwa, respectivamente, y diferían significativamente (P = 0,001). La variabilidad en las tasas de crecimiento y la duración de la estivación reflejan los tamaños óptimos de las poblaciones naturales de los tres grupos. Key words: The giant African Snail, Achatina achatina, aestivation, Ecotypes. Palabras clave: Caracol gigante africano, Achatina achatina, estivación, ecotipos. INTRODUCTION Achatina achatina (Linne, 1758) is which is maximal in the rainy season found in the closed forest area in and minimal in the dry season. The Ghana. It shows an annual activity snail burrows into the upper 10-15 cm * Forestry Research Institute of Ghana. UST. Box 63, Kumasi, Ghana. AS Iberus, 15 (2), 1997 COTE d' IVOIRE e Nkasem _-“eKumasi _- eDonyina xx Moist Semi-deciduous GULF OF GUINEA Figure 1. Map of the closed forest area of Ghana showing the trial site (Kumasi) and the origins (Donyina, Nkasem and Apedwa) of the snails used in trials. Figura 1. Mapa del área de bosque cerrado en Ghana mostrando el lugar del experimento (Kumasz) y los orígenes de los caracoles usados en el mismo (Donyina, Nkasem y Apedwa). of soil during the dry season and remains dormant for a period ranging from three to five months (COBBINAH, 1993). This state of dormancy during the dry season is referred to as aestiva- tion. Circannual rythms are known to be induced by such factors as light, temperature, humidity and soil water deficit (OwEnN, 1966). In the period leading to the onset of aestivation, there is a progressive decline in the snail's metabolism. BARATOU (1988) asserts that in order to maintain an equilibrium between water in its tissue and the rela- tive humidity of the immediate envi- ronment, snails allow themselves to dehydrate during this period. For Achachatina marginata dehydration leads to the loss of about 42% liveweight of the non-shell tissues (STIEVENART, 1994). Later, a mucous layer is secreted to cover the shell opening; the fully formed mucous layer is impervious to both gases and water (HODASI, 1982). Snails go into a state of dormancy whe- , never conditions are too dry for their 76 liking. Whilst this behaviour is most common in the dry seasons, COBBINAH (1993) reported that even dry spells during the wet seasons may induce aestivation in A. achatina. This is, however, in contrast to the observation of HODASI (1982) that exten- sive and persistent dry conditions are necessary to induce aestivation, and that aestivation normally does not occur during the dry spells in the rainy season. Extremes of temperature and starvation is also reported to induce aestivation (KONDO, 1964). HODAsI (1982) suggested that not every individual in the population aesti- vates during the dry season. Accor- dingly in certain localities within the distribution range of A. achatina in Ghana, fresh snails can be obtained throughout the dry season. Here the results of studies undertaken to deter- mine the variability in aestivation res- ponses of populations from three dis- tinct enclaves of A. achatina in Ghana is reported. COBBINAH: Aestivation responses of the giant African snail, Achatina achatina Linne MATERIALS AND METHODS Sources of Snails: The snails for the study were obtained from (a) Donyina in the Ashanti Region, (b) Nkasem in the Brong Ahafo Region and (c) Apedwa in the Eastern Region. Donyina is 20 km from our test site on the campus of the Univer- sity of Science and Technology, Kumasi, Ghana. Nkasem to the north west and Apedwa to the south east of Donyina are 109 and 198 km respectively from our test site. Figure 1 shows the origin of the th- ree populations together with approxi- mate isohyets. Donyina (6* 45' N and 2925' W”, Nkasem (6* 15” N and 2* 20' W) and Apedwa (6” 46" N and 1? 25 E) all fall wit- hin the rainfall regime 1250 mm-1750 mm typical of the moist semi-deciduous forest type. Seasonal rainfall at the three origins is influenced by meteorological Equator (ME). Two weather systems are associa- ted with the ME, the Intertropical Front and the Intertropical Convergence Zone which cause short heavy rain storms and abundant continuous rain respectively and result in bimodal annual rainfall pattern with major rains falling between April- June and minor period of rains between September and October separated by two months of less frequent rains (LEROUxX, 1988). The mean annual rainfall for the th- ree locations are Donyina (1403 mm), Nka- sem (1395 mm) and Apedwa (1561 mm). The main dry season falls between De- cember and March. The soils at the three areas are of the forest ochrosols type. While all three areas fall within one forest vege- tation type, different levels of logging and agricultural practices have resulted in var- ying rates of deforestation. The Donyina site has completely been converted into farmland over the years. The Apedwa site falls within the mountainous Atewa range protection forest reserve where timber ex- ploitation is prohibited. However, pockets of illegal farms are not uncommon. The Nkesem site is an off reserve area with fairly dense vegetation interspersed with farmlands. Growth Rates: The snails were con- ditioned in our research plot for four we- eks on a diet consisting of pawpaw (Ca- rica papaya) leaves and fruits, cocoyam (Xanthosoma mofafa) leaves, and leaves of the fameflower plant (Talinum triangulare). Individuals that showed signs of inacti- vity during this period were not used for the trials. Twenty snails of each group (ecotype) were placed in wooden boxes measuring 0.6 x 0.6 x 0.35 m, filled to a depth of 20 cm with sieved sterile silty sandy soil obtained from an abandoned rubbish dump. The snails in the boxes were offered excess amounts of food but left over foods were removed daily and soils overturned weekly. Because of high sur- vival rates of A. achatina (COBBINAH AND OsEI-NKRUMAH, 1988) and insignificant changes in shell size over short periods of time, growth rates were measured by chan- ges in live weights of snails. Each treat- ment was replicated three times, with data recorded for eight weeks prior to onset of aestivation and eight weeks after emer- gence from aestivation. Aestivation patterns of the three populations: The aestivation patterns of individual snails used in the growth studies described above were monito- red. Snails were considered as having aestivated when they covered the shell opening with a white mucous layer. Snails were recorded as emerged from aestivation when they discarded the epiphragm and resumed feeding. A second trial was conducted to deter- mine the effects of increased humidity. Each group of snails was divided into two lots. Twenty snails from one lot were placed in a 0.6 x 0.6 x 0.3 m wooden box as described above. A second set of 20 snails of the same ecotype was placed in another box with humidity slightly incre- ased by making a platform about 15 cm above the soil level and placing a moiste- ned fibre bag on the platform. The bag was kept moist for two weeks before onset of aestivation and throughout the aesti- vation period. Similar sets were set up for the other ecotypes and each treatment was replicated three times. All snails used in the studies were labelled (paint marked) to enable obser- vation of individual activities daily. Aestivation responses (time of onset and DÍ Iberus, 15 (2), 1997 Table I. Growth rates of three populations of Achatina achatina cight weeks before and after aesti- vation period. Tabla I. Tasas de crecimiento de las tres poblaciones de Achatina achatina ocho semanas antes y después del periodo de estivación. Mean initial Mean weight gained Mean weight gained weight (g) in 8 weeks before in 8 weeks after Ecotype + s.e. aestivation + s.e. aestivation + s.e. Donyina 46.6+1.25 3.60+0.58% 10.83 + 0.172 Nkasem 49.7+0.95 14.27+0.81? 12.47+ 0.359 Apedwa SS 19.20+1.37< 19.58 + 2.87* Means within a row followed by the same letter are not significantly different (P= 0.05). emergence from aestivation) were recor- ded for all snails. The time required for 50% of each group to aestivate (TEsso) or emerge from aestivation (TEms0o) was estimated for the various treatments and populations. RESULTS AND DISCUSSION Growth rates of the three popula- tions: The pre-aestivation growth rates for the Nkasem and Apedwa snails were four and five times more than that recorded for the Donyina snails. The mean growth rates 8 weeks before aesti- vation ranged from 3.6 g for Donyina to 19.2 g for Apedwa (Table 1). Analyses of the data (ANOVA) in Table 1 indicate that the growth rates among the three populations differed sig- nificantly for the pre-aestivation (F = 83.94; df =2, 6; P< 0.001) and post-aestivation (F = 7.73, df 2, 6; P< 0.02) periods. However, Fisher's Multiple Range Test (LSD) did not show significant difference in the post- aestivation growth rates between the Donyina and Nkasem snails. The very low growth rates recorded for the Donyina group 8 weeks before aestivation suggest that, perhaps, aestivation in this group is preceded by significantly longer period of inactivity. Both the pre and post aesti- vation growth rates of the three popula- tions indicate that the Apedwa group might be the most desirable group for commercial snail farming. 78 Aestivation Patterns of the three Populations: Figure 2A shows the aesti- vation pattern of the entire snail popula- tion. It took 10 weeks for all the snails in the test to aestivate. Three peaks are found in the second, sixth and tenth week. The three peaks show the hetero- geneity in the response of the entire population to factors inducing aestiva- tion. Figure 2B shows the aestivation pat- tern of the Donyina ecotype. Aestivation commenced on 29 October 1993 and pea- ked on 6th November, 1993, one week after the beginning of aestivation. The entire Donyina group aestivated in 5 we- eks. The first observation of aestivation for the Nkasem (Fig. 2C) and Apedwa (Fig. 2D) groups was recorded on 6th November, 1993 but peak aestivation for these two groups was recorded on 4th December, 1993 and 28th December, 1993, respectively. Aestivation for these two groups spanned a period of 8 and 9 weeks. Time taken for 50% of each pro- pulation to aestivate (TEss0) were 8, 28 and 52 days for Donyina, Nkasem and Apedwa snails respectively. The relatively longer periods required for the Nkasem and Apedwa populations to complete aestivation is a reflection of the variability within these two popula- tions. The peaks observed in Figures 2B- D corresponded to the peaks in Figure 2A and suggest that the heterogeneity in the aestivation pattern of A. achatina observed in our study was due mainly to the varia- COBBINAH: Aestivation responses of the giant African snail, Achatina achatina Linne ES 3 Percent aestivating 5 0 22)r0fa3 Sin wn 3h "h2 ap Aestivation date Percent aestivating 220 Sm am 32 M2 31h2 Aestivation date Percent aestivating . 40 4, Dec —> [27] o Percent aestivating mn 3 0 2/h0 sum 14n 3/2 Aestivation date 1712 31/12 50 28 Dec.—> D uy o N o 3 0 2/0 sn. 19m 32 vh2 3112 Aestivation date Figure 2. Aestivation patterns. A: entire experimental population; B: Donyina population; C: Nkasem population; D: Apedwa population. Figura 2. Patrones de estivación. A: toda la población experimental; B: población de Donyina; C: pobla- ción de Nkasem; D: población de Apedwa. tion in the responses of the three groups to the factors inducing aestivation. The emergence period for the entire population covered a period of 7 weeks from the end of January to mid-March, 1994. Again three peaks were evident (Fig. 3A). These were in the third, fifth and seventh week and corresponded to peak emergence periods for the Apedwa (Fig. 3D), Nkasem (Fig. 3C) and Donyina (Fig. 3B) respectively. The first snail to emerge from aestivation was from the Apedwa group on 31 January 1994 (Fig. 3D). By mid-February 60% of this group had emerged from aestivation. On the other hand, not a single snail from the Donyina group had emerged by mid-February, three and half months after initiation of aestivation (Fig. 3B). Twenty percent of the Nkasem group had resumed normal metabolic activities by mid-February (Fig. 3C). Peak emergence in the Donyina group was recorded during the first week in March. Time taken for 50% of popula- tion to emerge from aestivation (T'Emso) following the outset of emergence were 9, YS Iberus, 15 (2), 1997 30 25 % Emergence from aestivation 0 6/1J94 30) az 2/2 vu 21 Emergence date 40 6Mar. —» 20 % Emergence from aestivation 0 6h 30h 13% 21 ad 7 Emergence date 40 20 Feb. E 30 u 2 E] o 5] 5 20 mn o [|] (5) o E 5 PS 0 6h 30h 132 2/2 133 27/3 Emergence date 40F 9Feb. — D 30 20 % Emergence from aestivation 0 16h 30h 19k 27h Emergence date 2/3 271 Figure 3. Emergence patterns. Á: entire experimental population; B: Donyina population; C: Nkasem population; D: Apedwa population. Figure 3. Patrones de salida de la estivación. A: toda la población experimental; B: población de Donyina; C: población de Nkasem; D: población de Apedwa. 20 and 33 days for Apedwa, Nkasem and Donyina groups respectively. Humidity is considered a major factor affecting aestivation behaviour of A acha- tina. COBBINAH (1993) reported that when the relative humidity falls during the dry season A. achatina becomes inactive, seals itself in its shell with a white calcareous layer and aestivates in order to prevent loss of water from the body. In this study the enhanced humidity (3% above the ambient condition) attained in the boxes with moistened fibre bags did not 80 influence the overall aestivation pattern of any of the three groups (see Table II). Elmslie (pers. comm.) asserts that aestivation / hibernation may be influen- ced by a programmable regulation, reset- table by environmental experience like cir- cadian rhythm, but transmitted to offs- pring in the case of parents that have changed environment in a partially reset state. It is possible that the enhanced humidity in these boxes was not adequate to destabilise the in-built mechanism which sets in motion physiological COBBINAH: Aestivation responses of the giant African snail, Achatina achatina Linne Table II. Aestivation patterns of snails in boxes with or without moistened fibre bags. Tabla II. Patrones de estivación de los caracoles en cajas con o sin bolsas de fibra humedecidas. Mean Weekly % Aestivation Mean weekly % Emergence Ecotype Dry (68-70% rh) Moist (68-74% rh) Dry (57-62% hr) Moist (56-66% rh) Donyina 22.16 SEA 26.54 25.49 Nkasem 19.42 157 15772 16.52 Apedwa 2096 23.91 13.97 14.80 All differences are not significant at (P = 0.05). changes resulting in aestivation during periods of low atmospheric humidity. Whilst aestivation has adaptive value for the snail (HODASI, 1982; STIEVENART, 1994), for the snail farmer it represents the loss of valuable growing time. The three populations show significant differences in growth rates and duration of aestiva- tion. Shorter aestivation period mean longer feeding period and ultimately larger body sizes. Based on peak aestivation and emer- gence periods for the three groups, the estimated duration of the dormant periods were 4, 10 and 16 weeks for the Apedwa, Nkasem and Donyina respectively. More- over, data on growth rates clearly show a decreasing order Apedwa > Nkasem > Donyina among the three groups during the pre and post aestivation periods. These two factors acting in concert may explain differences in adult sizes of A. achatina from various areas of the country. The Apedwa snails are usually twice the size of the Donyina snails (COBBINAH, 1993). The Nkasem snails are often intermediate in size between the two populations. Although all the three enclaves where the snails originated from are within the moist semi-deciduous forest type and are characterized by similar soil type, there are differences in mean annual rainfall and vegetation cover. Soil water regime are influenced by rainfall gradient and evapotranspiration (VAN ROMPAEY, 1993). Most studies of the soil water regime in West African tropical forest (HUTTEL, 1975; COLLINET, MONTE- NEY AND POUYAUD, 1984) suggest that seasonal soil water deficits increase with decreasing annual rainfall. Mean annual rainfall is highest at Apedwa (1561 mm) but similar at Donyina (1403 mm) and Nkasem (1359 mm). In the three enclaves the Donyina area is the most degraded due to logging and slash and burn agri- culture practices over the years. Unlike Apedwa and Nkasem where snails are mainly gathered from forest reserves and secondary forests outside reserves, the Donyina snails are mainly gathered from low vegetation farmlands. Alt- hough Donyina and Nkasem have similar mean annual rainfall, the relati- vely poor vegetation cover at Donyina would result in higher evapotranspira- tion and longer duration of seasonal drought. The snails from this area have probably adapted to this relatively long drought period through extension of dormancy period. All individuals in the three groups aestivated in these studies. Neverthe- less, a few individuals among the Apedwa group had shorter periods of aestivation than the four week group average. Further studies are, however, underway to determine whether some individuals or groups normally remain active throughout the dry season, and also to better understand the physiologi- cal, environmental and behavioural fac- tors controlling aestivation. If the varia- bility in the aestivation behaviour by the different individuals or groups has a significant genetic component, the resul- ting information would be of potential use for commercial snail farming. 81 Iberus, 15 (2), 1997 REFERENCES BARATOU, J., 1988. Raising snails for food. llu- minations Press, Calistoga, USA, 72 pp. COBBINAH, J. R. AND OSE-NKRUMAH, A., 1988. The effect of food on growth of Achatina acha- tina. Snail Farming Research, 2: 20-24. COBBINAH, J. R., 1993. Snail farming in West Africa: a practical guide. Sayce Publishing Ltd., Exeter, U. K., 49pp. COLLINET, J., MONTENEY, B. AND POUYAUD, B., 1984. Le millieu physique. Recherche de amena- gement en millieu forestier tropical humide: le pro- jet taide Cote d'Ivoire. Notes Techniques du MAB, 15. HODASL J. K. M., 1982. Some aspects of the bio- logy of Achatina (Achatina) achatina (Linne). Bulletin de l'IFAN, 44: 100-114. HUTTEL, CH., 1975. Recherches sur l'ecosys- teme de la foret subequatoriale de basse Cote d'Ivoire. IV. Estimation du Bilan hydrique. La Terre et la Vie, 29: 192-202. 82 KONDO, L., 1964. Growth rates in Achatina fu- lica Bowdich. Nautilus, 78: 6-15. LEROUX, M., 1988. La variabilite des precipita- tions en Afrique occidentale. Les compo- santes aerologigues du probleme. Veille cli- matique satellitaire, 22: 26-46. OWEN, D. F., 1966. Animal Ecology in Tropical Africa (1st Ed.). Oliver and Boyd, London, 122 bs STIEVENART, C., 1994. Artificial estivation of the giant African snail Archachatina marginata sa- turalis: Survival, weight loss and meat yield. Snail Farming Research, 5: 23-28. VAN ROMPAEY, R.S. A.R., 1993. Forest gradients in West Africa. Ph. D. Thesis, Agricultural University, Wagemningen, 98 pp. O Sociedad Española de Malacología —————— IBERUS, 15 (2): 83-93, 1997 Snail communities associated to swampy meadows and sedgy marshy meadows plant communities of the Great Hungarian Plain Comunidades de moluscos asociadas a comunidades vegetales de pra- deras pantanosas y junqueras en la Gran Llanura Húngara Károly BÁBA* and István BAGI'** Recibido el 11-X-1995. Aceptado el 29-1V-1997 ABSTRACT Simultaneous phytocoenological, malacological and pedological studies were carried out in six successional plant community types characteristic on the Great Hungarian Plain. Data were analyzed by multivariate statistical methods (PCA). Variation in the abundance of ecological, habitat type and nutritional type species groups was also followed. In the Succiso-Molinietum (swampy meadows) and Agrostio-Caricetum [sedgy marshy meadows) plant communities, the distribution of constant and differential species is mostly influenced by their range of pH tolerance. Habitat drying and salinization, and various human impacts (draining, cutting and grazing by domestic animals) influence the succession of vegetation. Changes in snail assemblages include altering proportion of living and dead individuals and decreasing diversity (H'), both reflecting habitat drying and salinization. Complementary changes in the abundance of riparian and steppe dweller species groups indicate habitat drying, while swamp dwellers become more numerous as the topsoil becomes muddy due to salt accumulation. Concerning nutritional types, the proportion of omnivores decreases with habitat drying, whereas the frequency of herbivores increases in a complementary manner. The increasing abundance of saprophagous snails reflects bio- tope eutrophization caused by cutting and grazing. RESUMEN Se han desarrollado simultaneamente estudios fitocoenológicos, malacológicos y pedoló- gicos en seis tipos de comunidades vegetales de la Gran Llanura Húngara. Los datos fue- ron analizados mediante métodos estadísticos multivariantes. También se ha estudiado la variación en los grupos de especies desde el punto de vista ecológico, de su hábitat y tipo nutricional. En las comunidades vegetales Succiso-Molinietum (praderas pantanosas) y Agrostio-Caricetum (¡junqueras), la distribución de las llamadas especies constantes y diferenciales está mayoritariamente influenciada por su rango de tolerancia de pH. La desecación del hábitat y su salinización, junto con un conjunto de alteraciones humanas (desecación, segado y ramoneado por animales domésticos) afectan la suceción vegetal. Los cambios en las comunidades de moluscos incluyen la variación en las proporciones de individuos vivos y muertos y una menor diversidad (H'); ambos cambios reflejan la desecación y salinización del hábitat. Cambios complementarios en la abundancia de los grupos de especies ribereñas y de estepa son indicadores de la desecación, mientras que las especies propias de zonas pantanosas se hacen más abundantes según la capa super- * Department of Biology, Gy. Juhász Teacher Training College, Szeged, Hungary. ** Department of Botany, A. József University, Szeged, Hungary. 83 IBERUS IS NZ)IIH 7 ficial se vuelve fangosa debido a la acumulación de sal. Por lo que se refiere a los grupos nutricionales, la proporción de omnivoros decrece con la desecación, mientras que la fre- cuencia de herbivoros crece de manera complementaria. La creciente abundancia de especies saprófagas refleja la eutrofización producida por el segado y ramoneo. KEY WORDS: Gastropoda, drainage, salinization, species groups, succession, Hungary. PALABRAS CLAVE: Gastropoda, desecación, salinización, grupos de especies, sucesión, Hungría INTRODUCTION Under the semiarid climate characte- ristic to the Eupannonicum floristic region (which the lowlands of the Car- pathian Basin belong to), habitat drying and salinization processes were studied along a successional series spanning from swampy to salt-affected meadows. The Succiso-Molinietum association represents the wet meadows on calcare- ous swampy meadows soils. The phy- siognomy of its vegetation is determi- ned by tall grasses (Molinia caerulea, M. arundinacea, Festuca pratensis, Deschamp- sia caespitosa, Agrostis stolonifera). Its stands have high species diversity and are rich in protected rare species (Dacty- lorrhiza incarnata, Orchis laxiflora palustris, Cruciata pedemontana, Veratrum album, Iris spuria, 1. sibirica, ...). The Succiso-Molinietum is in successional relation to the Agrostio-Caricetum asso- ciation; namely, moderate habitat drying and salinization involve its transforma- tion into the latter. The Agrostio-Carice- tum may occur independently, too. It is characteristic of the solontschak-solonet- zic alkaline meadows soils. Depending, on the hydro- and haloecological condi- tions the Agrostio-Caricetum forms various vegetation units (eg. subassocia- tions), that well reflect the environmen- tal impacts. These units are extremely diverse in its species composition and their physiognomy. The two associa- tions represent the meadow formation of the Great Hungarian Plain in a signi- ficant percentage. As the two associa- tions have evolved under wet condi- tions, the drought on their habitats causes drastical transformation in their vegetation structure, first of all in their species composition. The changes in the 84 structure show close relationship to the environmental conditions, therefore every distinguished vegetation unit well reflects a stage of the drought induced vegetation transformation processes (phytocoenological indication). A lower proportion of data regards to other plant associations, that are in successional relationships with Succiso-Molinietum and Agrostio-Caricetum. The water management works in the 20 century and the more and more arid climate of the last two decads endange- red the vegetation of the wet meadows. Their transformation into drier habitats would have harmful consequences for the whole ecosystem, particularly the animal assemblages, that have no enough mobility to change their habitat. The terrestrial snails belong to a little mobile group of animals, they are bound to their habitats more firmly than other ones; therefore the, however well- known, successional and zonational re- lationships of the vegetation units can be examined on the basis of their snail assemblages, too. If the transformation processes of vegetation and their animal assemblages show paralelism, the chan- ges in the composition of this animal group would have an indicative value for nature conservation. MATERIALS AND METHODS In the South-eastern part of the Great Hungarian Plain (Csongrád County) snail assemblages were sam- pled by the quadrat method. Ten plots 25 x 25 cm size were examined in para- llel with phytosociological recording of BÁBA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hungary each plant subassociation encountered (S0ó, 1964). A hight proportion of the data regards to two plant communities: Succiso-Molinietum and Agrostio-Carice- tum; only some samples originated ot- her successionally related association. Altogether 30 collection sites were visi- ted in six plant community types, while the number of subassociations studied was 22 (see Figure 3). In each quadrat, a detailed soil analysis was conducted, in- cluding measurements of relative per- centage soil moisture, total organic mat- ter content, CaCOz3 concentration, hy- groscopy and pH. The concepts applied in the coenolo- gical characterization are the following; Abundance (A) is the number of indivi- duals of a snail species found in a plant community regarding to one m? (A /m?); Dominance (D) is the ratio of indivi- duals of a species related to the total individuals of every species; Species density (SD) is the average species number of 10 quadrats in a collection site; Frequency (F) is the ratio of a species in relation to the total number of species in a collection site (consist of 10 quadrats); Constancy (K) is the ratio of a species in relation to the number of all species found in all collection sites belonging to the same plant community. When a species was found in all the quadrats, it can be considered as an absolute constant species. D, F and K are expressed as percentages. Data were analyzed by standardized Principal Components Analysis (PCA, PODANI, 1988). Shannon-diversity (H'), SD and changes in the abundance (A/m?) of living and dead individuals were followed through examination of the proportions of various species groups. Ecological species groups were defined as follows: S: sciophilous, P: swamp dweller, Ph: photophilous, R: riparian and OA: species of open areas. They were obtained by applying the block cluster method of FEOLI AND ORLÓCZI (1979). A simplified version of LoZEK'"s (1964) typology was used and the following habitat type groups were distinguished: riparian ubiquitous (RU), bush forest dweller (B), hygrophilous swamp dweller (HP) and steppe dweller (ST). Nutritional type groups (O: omni- vore, SP: saprophagous, H: herbivore) were differentiated after the system of FRÓMMING (1954). Species and their group assignments are listed in Table 1. RESULTS AND DISCUSSION Species encountered: Field studies yielded a collection of 3047 living and 3150 dead individuals belonging to 26 species (Table 1). The majority of the specimens was found in the Succiso- Molinietum (1062 + 1268) and Agrostio- Caricetum (1496 + 1445) phytocoenoses, while plant associations 1, 3, 4 and 6 altogether contained 489 + 440 alive and dead individuals, respectively. Two species new to the southern part of the Great Hungarian Plain were detected: Malacolimax tenellus (O. F. Muller, 1774) and Deroceras sturanyi (Simroth, 1894). Characteristic species and their re- quirements: On the basis of frequencies of occurrence data, the constant, sub- constant and accessorial species could be determined for the two plant associa- tions most rich in snails (Table ID). Cons- tant and subconstant species reach low levels of dominance in both communi- ties. This is probably due to unfavoura- ble changes in their environment caused by either draining, drying, salinization or grazing. Differential species are Coch- licopa lubricella (Porro, 1938) and Cary- chium minimum O. FE. Miller, 1774 in the Succisio-Molinietum plant association, and Pupilla muscorum (L., 1758) in the Agrostio-Caricetum. The occurrence of Truncatellina, Granaria and Helicopsis spe- cies in the Agrostio-Caricetum association indicates habitat drying. The distribution of constant and differential species is strongly influenced by the width of their pH tolerance range (Figs. 1, 2), as it has been shown earlier (BABA AND DOMON- KOs, 1992). According to aur data diffe- rential species of the Succisio-Molinietum association have a narrow pH tolerance range, in contrast with species occurring 85 IBERUS, 15 (2), 1997 Table I. Gastropod species found in the plant communities studied (1: Caricetum acutiformis-ripa- riae, 2: Succiso-Molinietum; 3: Bolboschoenetum maritimae, 4: Astero-Agrostietum;, 5: Agrostio- Caricetum distantis, 6: Achilleo-Festucetum pseudovinae) E: Ecological species groups (S: sciophilous; P: swamp dweller; Ph: photophilous; R: riparian; OA: species of open areas); N: Nutritional type (O: omnivore; SP: saprophagous; H: herbivore); H: Habitat type (RU: riparian ubiquitous; B: bush forest dweller; HP: hygrophilous swamp dweller; ST: steppe dweller). Tabla I. Especies de gasterópodos encontradas en las comunidades vegetales estudiadas (1: Caricetum acu- tiformis-ripariae; 2: Succiso-Molinietum, 3: Bolboschoenetum maritimae; 4: Astero-Agrostietum; 5: Agrostio-Caricetum distantis; 6: Achilleo-Festucetum pseudovinae) E: Grupos ecológicos de las especies (S: esciófilo; P: de marisma; Ph: fotófilo; R: ribereño; OA: de áreas abiertas); N: Tipo nutricio- nal (O: omnívoro, SP: saprófago, H: herbívoro); H: Tipo de hábitat (RU: ribereño ubiquo; B: zonas arbustivas; HP: marismeño higrófilo; ST: estepa). E N H 1 2 3 4 5 6 SSP OR Carychium minimum (0. F. Miller 1774) 17 5 S SP HP Carychium tridentatum (Risso 1826) 8 0A HB Cepaea vindobonensis (Ferussac 1821) l 2 6 DA SP ST Chondrula tridens (0. E. Múller 1774) 135 l 388 50 Rs 0 Bb Cochlicopa lubrica (0. F. Múller 1774) 3 4 DA 0. ST Cochlicopa lubricella (Porro 1838) + 184 R 0 RU Deroceraslueve(0.F. Miller 1774) l R 0. HP Derocerassturanyi(Simroth 1894) l 0A 0 B Euconulus fulvus (0. F. Miller 1774) 4 0A H ST Granaria frumentum (Draparnaud 1801) + 86 0A H ST Helicella obvia (Menke 1828) 3 MAS Helicopsis striata (0. E. Múller 1774) 1 Ph H B Helix pomatia (Linne 1758) 2 S 0 RU Malacolimax tenellus(0.P. Miller 1774) l PH ST Monacha carthusiana (0. F. Miller 1774) 18 99 13 AS R H RU Perforatella rubiginosa (A. Schmidt 1853) 3 0A H ST Pupilla muscorum (Linne 1758) l l 329 165 R 0 RU Succineaoblonga (Draparnaud 1801) 8 169 43 6 130 + P 0 HP Succinea elegans (Risso 1826) 1 DA SP ST Irucatellina cylindrica (Ferrusac 1807) 3 0A H ST Vallonia costata (0. E Múller 1774) | 8 6 REO p Vallonia enniensis (Gredler 1856) 69 39 244 RS Vallonia pulchella (0. E. Múller 1774) 19 40 A RES SRU Vertigo antivertigo (Draparnaud 1801) 1 A A | Vertigo pygmaea (Draparnaud 1801) 1 20 44 2 R 0 RU Zonitoides nitidus(0.F. Miller 1774) 12 1 Number of Individuals 151 1062 58 12 1496 268 Number of collection sites l 11 l l 15 l Number of Species 11 17 3 3 15 7 Dead Individuals 155 1268 OD 42 1445 243 in both Agrostio-Caricetum and Succisio- Molinietum, which tolerate a much wider range of soil pH (Fig. 2). 86 Successional changes: The ordina- tion of snail samples of collection sites (1-30) clearly indicates (Fig. 3) the suc- BÁBA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hungary Table I1. The constant (above), subconstant (middle) and accesorial (below) species of the two plant communities (Succiso-Molinietum, Agrostio-Caricetum) studied in more detail. K: constancy; D: dominance. Tabla 11. Las especies constantes (arriba), subconstantes (medio) y accesorias (abajo) en las dos comuni- dades vegetales estudiadas (Succiso-Molinietum, Agrostio-Caricetum). K: constancia; D: dominancia. Succiso - Molinietum K D Monacha carthusiana 100 9.32 Succinea oblonga 90.9 15.91 Cochlicopa lubricella 81.81 17.32 Vallonia enniensis 1272 36.72 Chondrula tridens IDEA Vertigo pygmaea 45.45 1.88 Carychium minimum 21.17 0.47 Vallonia pulchella 18.18 3.76 Cochlicopa lubrica 18.18 0.28 Carychium tridentatum 9.09 0.75 Perforatella rubiginosa 9.09 0.28 Cepaea vindobonensis 9.09 0.09 Deroceras laeve 9.09 0.09 Deroceras sturanyi 9.09 0.09 Malacolimax tenellus 9.09 0.09 Pupilla muscorum 9.09 0.09 Vertigo antivertigo 9.09 0.09 Agrostio - Caricetum K D Chondrula tridens 100 25.93 Monacha carthusiana 100 13.03 Succinea oblonga 80 8.68 Pupilla muscorum 73.33 21.99 Vertigo pygmaea 33.33 2.94 Vallonia pulchella 26.66 3.67 Cepaea vindobonensis 20 0.4 Helix pomatia 13.13 0.13 Granaria frumentum 6.66 5.74 Vallonia costat 6.66 0.53 Cochlicopa lubricella 6.66 0.26 Iruncatellina cylindrica 6.66 0.2 Helicopsis striata 6.66 0.06 Zonitoides nitidus 6.66 0.06 cessional and zonational relationships of the plant associations studied, and a gradual habitat drying (Baci, 1988). The main lines of the drying processes can be outlined as follows: (A) Non-salinic, wet line Caricetum acutiformis-ripariae (L, 1), Succiso-Molinietum typicum (IL, 2-10), S. -M. poetosum (X, 11), S. -M. agrostieto- sum (XI, 12). The latter is a connection to a different, saline line: (B) Bolboschoene- tum maritimae (IV, 14), Agrostio-Caricetum bolboschoenetum (IV, 21), A. -C. fac. Juncus (UL 16), Agrostio-Caricetum plantagineto- sum maritimae (V, 17-20). Later, while the drying procces continues, the two lines originated a common line (C), whose re- presentative associations are A-C. festu- cetosum arundinacene (VI, 28), A. -C. poe- tosum (Vlla, 22-24), A. -C. festucetosum pseudovinae (VII, 15) and finally the Achilleo-Festucetum pseudovinae (IX, 29). Roman numbers indicates groups of co- llection sites with similar features obtai- ned from the analysis. The successio- nally closely connected plant communi- ties often form zonation systems in the field. The drying processes have been particularly accelerated since the sixties, due to the draining of the region. The three successionally related lines could be distinguished in the process of drai- ning-generated habitat drying by inves- tigation of snails, too. The declining density of dead shells and living indivi- duals, the fall of individual density (ID) and diversity (H”) indicate habitat dr- ying (X, V, Vlla, b, IX and VIII) and sometimes salinization (Fig. 4). Habitat drying accelerates with draining, which then leads to higher snail abundance again at the end of the successional xero-series at dry localities (Achilleo-Fes- tucetum association). Chondrula and Pu- pilla can become especially numerous. Snail species groups were used to evaluate these processes, for which the 87 IBERUS, 15 (2), 1997 2. axis 1.907 3 8 1 4 2 6 9 7 -2.043 5 Í | 1. axis -2.038 2.23 Figure 1. Distribution of constant and subconstant species according to soil pH (standardized PCA). 1: Carychium minimum, 2: Succinea oblonga; 3: Cochlicopa lubricella; 4: Vertigo pygmaea; 5: Pupilla muscorum; 6: Vallonia costata;, 7: Vallonia pulchella; 8: Vallonia enniensis, 9: Chondrula tridens. Figura 1. Distribucion de las especies constantes y subconstantes de acuerdo con el pH del suelo (Análisis de componentes principales estandarizado). 1: Carychium minimum, 2: Succinea oblonga; 3: Cochlicopa lubricella; 4: Vertigo pygmaea; 5: Pupilla muscorum,; 6: Vallonia costata; 7: Vallonia pulchella; 8: Vallonia enniensis; 9: Chondrula tridens. 9.2 No) 8.8 8.6 8.4 pH interval (0,2) 159) 8 9 2 3) 4 3 7 6 l Figure 2. Distribution of the pH tolerance ranges of constant and subconstant species. Numbers refer to species as in Figure 1. Figura 2. Distribución de los rangos de tolerancia al pH de las especies constantes y subconstantes. Los números de las especies son idénticos a los de la Figura 1. 88 BÁBA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hungar; 2.axis 1.744 - 0.896 -1.329 1.258 Figure 3. Ordination of collection sites by using data of snails samples (standardized PCA). Roman numbers indicate sample groups with similar phytological and malacological features; groups obtai- nes by ordination of data. Arabic numerals in brackets indicate collection sites. Caricetum acutifor- mis-ripariae 1 (1); Succiso-Molinietum a. typicum facies Veratrum album 1 (9), b. typicum facies Phragmites 11 (10), c. typicum normale 1 (2-8), d. poetosum angustifoliae X (11), e. agrostietosum XI (12); Bolboschoenetum maritimae YV (14); Astero - Agrostietum IV (30); Agrostio - Caricetum distan- tis, a. agrostietosum IV (13), b. agrostietosum facies Juncus compressus YI (16), c. plantaginetosum maritimae Va, b (17-20), d. poetosum angustifoliae Vlla (22-24), e. festucetosum arundinaceae VÍ (28), £. festucetusum pseudovinae VII (15), g. bolboschoenetosum IV (20); Achilleo - Festucetum pseu- dovinae IX (29). See the text for further details. Figura 3. Ordenación de las estaciones de muestreo utilizando datos de muestras de moluscos (Análisis de componentes principales estandarizado). Los números romanos indican grupos de estaciones con similares características fitológicas y malacológicas; los grupos se obtuvieron por ordenación de los datos. Los núme- ros arábigos entre paréntesis indican las localidades de muestreo. Caricetum acutiformis-ripariae / (1); Succiso-Molinietum 4. typicum facies Veratrum album /7 (9), b. typicum facies Phragmites // (10), c. typicum normale /7 (2-8), d. poetosum angustifoliae X (11), e. agrostietosum AX7 (12); Bolboschoenetum maritimae /V (14); Astero - NIV (30); Agrostio - Caricetum distantis, 4. agros- tietosum /V (13), b. agrostietosum facies Juncus N1I] (16), c. plantaginetosum maritimae Va, b (17- 20), d. poetosum angustifoliae V//a (22-24), e. festucetosum arundinaceae VI (28), f. festucetusum pseudovinae VIT (15), g. bolboschoenetosum 7V (20); Achilleo - Festucetum pseudovinae /X (29). Véase el texto para más detalles. abundantial changes are shown (Figs. 5, quists (R, RU) monotonously decreases 6, 7). Habitat drying and salinization (collection sites 11 and 12 represent sta- have different consequences in the two ges of ramification in the successional se- plant associations. In the Succiso-Molinie- ries). As the wet terrain dries down gra- tum one, the abundance of riparian ubi- dually, the abundance of species typical 89 IBERUS, 15 (2), 1997 1000 900 800 700 600 DA /m? 500 5 la) /m? VI Vila Vilb 1X vi A B C Figure 4. Variation in species density (1D), species diversity (H”), pH and density of living (A/m?) and dead individuals (DA/m?) in groups of phytocoenologically similar samples (Roman numerals) influenced by the dominant process of the habitat changes. A, B and C refer to the three succession lines of the vegetation. A includes groups 1, II, X and XI; the main processes are drying (L, ll and X) and salinization (XI). B includes groups IV, II! and V; the main processes are drying (IV and 111) and an additional salinization with cutting (Va, 18-20) and degradation (Vb, 17). C includes groups VI, VIL VII and IX; the main processes are drying (VI, Vlla 22-24 and IX) with cutting (VIIb 25- 27) and drainage (VII. Figura 4. Variación en la densidad de las especies (ID), diversidad (H), pH y densidad de individuos vivos (A/m*) y muertos (DA/m?) en los grupos de muestras con características fitocoenológicas similares (números romanos) influenciadas por los procesos dominantes en los cambios de hábitat. A, B y C se refieren a las tres líneas de sucesión de la vegetación. A incluye los grupos l, Il, X y XI; los principales pro- cesos son la desecación (L, 1I y X) y la salinización (X1). B incluye los grupos IV; 11 y V: los principales procesos son la desecación (IV y 11) y una salinización adicional con el segado (Va, 18-20) y degrada- ción (Vb, 17). C incluye los grupos VI, VI, VII y LX; los procesos principales son la desecación (VI, Vlla 22-24 y IX) junto con el segado (VIIb 25-27) y el drenaje (VII). in open areas (OA, ST) decreases, para- lleled by a similar decline in the number of species and individuals (Figs. 5, 6). Concerning ecological species groups, the abundance of swamp dwellers (P, HP, Monacha) becomes higher with bio- tope salinization (12). Na* accumulation in the topsoil is responsible for its muddy character. 90 Sciophilous (S) and bush forest dwe- ller (B) snail species may also appear in the tall and dense stands of Caricetum acutiformis-ripariae community. The abundance of OA and ST species groups increases at the end of the successional series in Agrostio-Caricetum and Achilleo- Festucetum associations. Groups P and HP are also more abundant there, due to BÁBA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hung; 80 s F 600 20 A / il L 500 - Ph j / 60 le ==-==-=R 11 ¡lp 400 ===. OA 07 — ¡pl S j! e 2 pa300 “ lle SE J £ l. < Ni l N l F 200 iS 4 y e Ñ US y o S z Ni L 100 SUN N E IN Ñ A 7% IE == NEÁ E 0 T 1 T T e A pO I u Xx XI Iv TI Va Vb VI Vila Vilb IX VI Figure 5. Variation in the abundance (A/m?) of ecological snail species groups among plant asso- ciations and subassociations. Abbreviations, S: sciophilous, P: swamp dweller, Ph: photophilous, R: riparian and OA: species of open areas. Figura 5. Variación en la abundancia (A/m?) de los grupos ecológicos de especies de moluscos en el con- junto de las asociaciones y subasociaciones vegetales. Abreviaturas, S: esciófilo; P: de marisma; Ph: fotófi- lo; R: ribereño; OA: de áreas abiertas. 120 RU L 700 le perio 0 coria 100 4 Í: ee E p E V : ' Pa En eS DON IV IM Va Vb VI Vila Vb IX VIn Figure 6. Variation in the abundance (A/m?) of snail habitat type groups among plant associations and subassociations. Abbreviations, RU: riparian ubiquist, B: bush forest dweller, HP: hygrophilous swamp dweller and ST: steppe dweller. Figura 6. Variación en la abundandia (A/m”) de los tipos de hábitat en las especies de moluscos en el con- junto de las asociaciones y subasociaciones vegetales. Abreviaturas, RU: ribereño ubiquo; B: zonas arbus- tivas; HP: marismeño higrófilo; ST: estepa. 91 IBERUS, 15 (2), 1997 250 (0) 2 Sp 200 OSA A 150 o 2) E < 100 50 y IU X Xi iv IM : E 400 :l al 350 Sl al E 300 20) ; 250 po 25 200 au E < 150 100 50 Va Vb VI Vila Vilb Ix MIU Figure 7. Variation in the abundance (A/m?) of snail nutritional type groups among plant associa- tions and subassociations. Abbreviations, O: omnivore, SP: saprophagous, H: herbivore. Figura 7. Variación en la abundancia (A/m?) de los grupos nutricionales de moluscos en el conjunto de las asociaciones y subasociaciones vegetales. Abreviaturas, O: omnívoro, SP: saprófago, H: herbívoro. the muddy topsoil formed under Na* accumulation (Figs. 5, 6). In both collection site groups of the Agrostio-Caricetum association, habitat salinization and drying, grazing and cutting result in the decline of the pro- portion of riparian ubiquists (R, RU: Succinea, Vallonia pulchella), and a com- plementary increase in the abundance of steppe dwellers (OA, ST: Chondrula, Pupilla). Stands of XL, Va and Vlla are regularly cut, while site of VIII is both cut and drained (Figs. 5, 6). In terms of nutritional types, omni- vorous snail species dominate in each collection site group, as it was also found elsewhere in willow-poplar fo- rests (Bába, 1993). With habitat drying the vegetation becomes denser, resulting in a higher abundance of herbivores. 2 Subsequent cutting increases the pro- portion of saprophagous species (Vallo- nia, Chondrula, Vertigo). The omnivore and herbivore-saprophagous groups were found to change in a complemen- tary manner (Fig. 7). According to our data, the connec- tion between the vegetation units and the species groups of snails seems to be very close. The composition of snail as- semblages indicates the most important environmental changes, such as drying and salinization, and the human im- pacts, eg. cutting, mowing and some ot- her disturbances. The structural transfor- mation of snail assemblages can be follo- wed at a level of species groups and also within these groups. The snail assembla- ges indicate not only the differences bet- ween plant communities but the diffe- BABA AND BAGI: Snail communities in swampy and sedgy marshy meadows in Hungary rent impacts of natural and anthropoge- nic factors within a plant community as well. The consequences of the habitat dr- ying are the decrease in species number and increase in abundance; the saliniza- tion causes the decrease of species num- ber and change of species groups in a particular habitat. Mowing leads to the decrease of species number and increase of the ratio of saprophagous and steppe dweller species groups. The changes can be traced back to pedological reasons, eg. habitat drying, increase in pH value, REFERENCES BABA, K. AND DOMOKOS, T., 1992. The occu- rrence and ecology of Chilostoma banatica (Rossmassler, 1838) in Hungary. Abstract of the Eleventh International Malacological Con- gress, Siena: 383-385. BABA, K., 1993. Effect of the regions of the Tisza Valley on the malacofauna. Tiscia, 18: 97-102. Bacií, L, 1988. The role of water management in the degradation processes of halophytic ve- getation in Hungary. Environmental Conser- vation, 15: 359-362. FEOLI, E. AND ORLÓCZI L., 1979. Analysis of con- centration and detection of underlying fac- tors in structured tables. Vegetatio, 40: 49-54. and accumulation of organic matter. The investigations of the structural and com- positional changes of snail assemblages of the studied six plant communities may provide a way to detect the conse- quences of the salinization as a characte- ristic successional process of Hungarian Great Plain. The studies on the changes of snail assemblages in meadow plant communities can indicate the main pro- cesses in this vegetatation type similar to the investigations carried in forest ecosystems (BABA, 1993). FRÓMMING, E, 1954. Biologie der mitteleuropais- chen Landgastropoden. Duncker-Huniblot, Ber- lin 1-404. LoZEk, V., 1964. Quartermollusken der Tschec- hoslowakei. Tschechoslowakischen Akademie der Wissenschaften. Praha, 374 pp. PODANI, J., 1988. Syn-Tax III. Users Manual. Abstracta Botanica, 12: 1-179. S06, R., 1964. Synopsis systematico-geobotanica flo- rae vegetationisque Hungariae I. Akadémiai Kiadó, Budapest, 589 pp. 93 dYas am sinezio do m SEO ui YY Apu rá sd egsldrnseas Un la de Aga esti ta pS de ond de Prosa dolia pil oia A 0 soi N 35% Ni ¡Je Ji wo da ya na leds De Ñ Fe 5 Caloabezá nt HE VAT lee 20 0 a lie 40 neta) 4 ] pS de 1 o $ dio 7 1999 ) M0 Ae. DUDA. al dhe E e oso sabe e - siniles 4 ET! De barba | rd sd ma 5 . bl Esos AS ebRol gré 939 bns eto o Y DA wtT Sr lo mb: pc a, | Subs DD cut ue moi mh E Ca ¿hepital sembla vic 13 AA TRI id . O Sociedad Española de Malacología Iberus, 15 (2)495-121, 1997 Morphological and biometrical researches on Austrian Clausiliids. Shell morphology and variability in Clausilia dubia Draparnaud, 1805 Investigaciones biométricas y morfológicas en Clausílidos de Austria. Morfología y variabilidad de la concha de Clausilia dubia Drapar- naud, 1805 Karl EDLINGER* Recibido el 8-[-1996. Aceptado el 28-V-1997 ABSTRACT Morphological and biometrical studies on shells of some Austrian populations of Clausilia dubia Draparnaud, 1805, show a great variability in size and in the other morphological features within the species as a whole and also within single populations. Investigations of the variability of characters by metrical and statistical methods in some populations dis- close impressive metrical divergences and morphological differences. These results give rise to the question whether the characterization of Clausilia dubia as a polytypic species, as suggested by Kemm (1960, 1973], is justified. RESUMEN Estudios morfológicos y biométricos en conchas de algunas poblaciones austriacas de Clausilia dubia Draparnaud, 1805, muestran una gran variabilidad en el tamaño y en otros caracteres morfológicos tanto en el conjunto de la especie como en poblaciones ais- ladas. Investigaciones sobre la variabilidad de caracteres por medio de métodos métricos y estadísticos de algunas poblaciones mestran importantes divergencias tanto métricas como morfológicas. Estos resultados originan la pregunta de si está ¡justificada la caracterización de Clausilia dubia como una especie politípica, como sugiere KiemM (1960, 1973). KEY WORDS: Clausilia dubía, subspecies, measures, distribution, morphological continuum. PALABRAS CLAVE: Clausilia dubía, subespecies, medidas, distribución, continuidad morfológica. INTRODUCTION As in great parts of Europe also in Austria, specially in the eastern parts of the Alps and in the adjacent areas, Clau- silia dubiía Draparnaud 1805 is a widely diffused and in some localities a com- mon species that is believed to be polyty- pic (KLeEmMM, 1960, 1973; FECHTER AND FALKNER, 1990; KEARNEY, CAMERON AND JUNGBLUTH, 1983; NORDSIECK, 1990). KLEMM (1960, 1973) gave a survey of di- * Naturhistorisches Museum Wien, 3. Zoologische Abteilung, Burgring 7, A-1014 Wien, Austria. YO Iberus, 15 (2), 1997 Y Y S ZP $9 ] ] [TD STD IB [ID Y ON C. d. dubia C. d. speciosa C. d. huettneri C. d. schlechti 9] 7 io 57) [TD 27 [> S [NIP PIIRAL La 2_/ [TD |) Dm A 7] [PI — E TD eZ |] E [15 EI] lA E C. d. gracilior C. d. tettelbachiana C. d. floningiana C. d. bucculenta — C. d. runensis C. d. kaeufeli Figure 1. Distribution and vertical succession of Clausilia dubia subspecies as is suggested by KLEMM (1960). Figura 1. Distribución y sucesión vertical de las subespecies de Clausilia dubia, tal y como sugiere KLEMM (1960). verse “subspecies” of Clausilia dubia ac- cording to the characters suggested by several authors. Traditional classification was based on a subjective selection of characters belie- ved to be important. This classification depended on the preferences of the single authors. In some cases, subspecies are even described according to the presence of typical features in the shell of few or sin- gle specimens. HOLOYAK AND SEDDON (1988) and NORDSIECK (1990) examined some of these descriptions and offered re- asonable revisions. Some of these Clausilia dubia “subspe- cies” are considered to be widely distri- buted, others to be localized in small areas or subdivided into isolated populations that live in separated sites (KLEMM, 1960, 96 1973). Furthermore, KLeMM (1960) sug- gested a vertical pattern of distribution and an altitude dependent succession of diverse subspecies at the eastern ranges of the Alps (Fig. 1). He tried to explain the ob- served distribution pattern by probable re-immigration events in the alpine re- gion after the Pleistocene and adaptation, enforcing the role of the environment (al- titude). Examination and possible revision of these interpretations have to critically con- sider the definition and the meaning of the term subspecies as it is used under va- rious perspectives by several authors. For the practical requirement of the collector and for the taxonomic ordering, of collec- tions the technical term “subspecies” as ba- sed on peculiar morphological features EDLINGER: Shell morphology and variability in Clausilia dubiía Draparnaud, 1805 Rax Schneeberg Hohe Wand Wienerwald Figure 2. A profil of the estern edge of the Alps and the Wienerwald with the sample localities. Figura 2. Perfil de la cara este de los Alpes y del Wienerwald con las localidades de muestreo. may certainly be useful. However, it is scientifically more important to consider the conceptual background of the termi- nology. In the given context the concept of race or subspecies refers to biological units that are groups of related populations which are genetically characterized and, in the case of very long isolation, may be the forerunners of “valid species” (MaYr, 1967: 387; SUDHAUS AND REHFELD, 1992). This view is highly important for tradi- tional evolutionary biology, especially in the frame of Darwinian models for evo- lutionary change. In respect to the biological relevance of the subspecies concept, Sudhaus and REH- FELD (1992) are suggesting that “geo- graphic races” (subspecies) are allopatric populations of a species, which can be distinguished taxonomically. The diag- nostic characters of races should be pre- sent in 90 or more percent of individuals of the population. “ OSCHE (1994) claims that 75 percent or more members of a population must be morphologically distinguishable from the members of another population. Popula- tions only in this case should be accepted as valid subspecies. In this paper Osche's definition is assumed as a very tolerant and useful concept. According to the presuppositions just given, studies referring to the subspecies problem have to treat large numbers of in- dividuals and to apply methods of grea- ter exactness than the usual descriptions and discriminations of characters based on a subjective and rather arbitrary approach. It is therefore necessary to study as great a number as possible of morphological characters and to take measurements of the greatest possible exactness to establish a solid basis for statistical evaluations (NE- MESCHKAL AND KOTHBAUER, 1988; KoTH- BAUER, NEMESCHKAL, SATTMANN AND WAWwRa, 1991; NEMESCHKAL, 1990, 1991, 1993; MYLONAS, KRIMBAS, TSIAKAS AND AYOUNTANTI, 1990). Such investigations may clear up, whether the populations studied by Klemm meet all requirements of a reliable identification as subspecies (EDLINGER AND FISCHER, 1997). It is also worth while to attempt a critical revision of some sam- ples of the Museum of Natural History in Vienna (NHMW). This paper is first at- tempt at elaborating a new basis for the discussion of the Clausilia dubia problem by morphological measurements. In future the anatomy of the soft parts and geneti- cal analyses might be also considered. MATERIAL AND METHODS 606 specimens of Clausilia dubia from different lots collected in various areas of the “Wienerwald” (Lower Austria in the southwest of Vienna) the massif of the “Hohe Wand”, the “Schneeberg” and the “Rax” were investigated. The localities from which the samples came were si- tuated at altitudes between 270 and 1850 m (Fig. 2). 7% Iberus, 15 (2), 1997 RA (in mm/10, x factor 20) Angle A BR (1-6) GW 15) B (in mm/10) MH (in mm/10) EA Ñ MB Gan mm/10)< N LE (1-5) Figure 3. Measures taken from the shells: shell-height (H), shell-width (B), height (MH) and width (MB) of the aperture, distance of ribs (RA), number of whorls (WZ), angle between the spindle axis and the upper palatal (left side) (A). Figura 3. Medidas tomadas en las conchas: altura (H), anchura (B), altura (MH) y anchura(MB) de la apertura, distancia de las estrías (RA), número de vueltas (WZ), ángulo entre el eje del huso y palatal superior (lado izquierdo) (A). Samples (Number of Sample (Sample localities, altitude/ numbers of specimens. R = Rax: R4 (Reichenau, 700 m/1 spec.), R5 (Aufstieg z. Knappenhof, 730 m/5 spec.), R7 (Knappenhof, 800 m/24 spec.), R10 (Thórlweg, 850 m/5 spec.), R11 (Thorl- weg, 960 m/1 spec.), R12 (Thórlweg, 1120 m/29 spec.), R13 (Thórlweg, 1260 m/20 spec.), R14 (Thórlweg, 1320 m/12 spec.), R15 (Jakobskogel, 1685 m/8 spec.); S = Schneéberg; S1 (Puchberg, 560 m/15 spec.), S2 (Schneebergbahn, 750 m/57 spec.), S3 (Schneebergbahn, 790 m / 5 spec.), S4 (Sch- neebergbahn, 945 m/30 spec.), S5 (Sch- neebergbahn, 1165 m/9 spec.), S6 (Schne- ebergbaln, 1370 m/6 spec.), S8 (Waxriegl 11, 1820 m/4 spec.), S9 (Waxriegl 11, 1850 m/26 spec.), S10 (Schneebergbahn, 1650 m/11 spec.), H= Hohe Wand: H1 (Dreistet- ten, 530 m/60 spec.), H2 (Einhornhóhle, 600 m/12 spec.), H3 (Drobilsteig, 700 m/24 spec.), H4 (Drobilsteig, 760 m/69 spec.), H5 (Auffahrt z. Plateau, 830 m/74 spec.), H6 (Plateau 1020 m/6 spec.), H7 (Plateau 1020 98 m /34 spec.), H8 (Plateau 1020 m/4 spec.); W = Wienerwald: W1 (Anninger, 400 m/11 spec.), W2 (Anninger, 450 m/4 spec.), W3 (Moódling-Klause, 260 m/8 spec.), W4 (Hu- sarentempel, 480 m/14 spec.), W5 (Auf- gang Aminger, 270 m/11 spec.), W6 (Peils- tein, 400 m/8 spec.), Several individuals were dissected. Dissections did not reveal significant dif- ferences in the genital apparatus. 18 shells, syntypes of Austrian and South Tyrolean (Italy) localities, given in loan by the Natur-Museum Senckenberg as typical representatives of various subspe- cies were used for comparisons (Fig. 1). They are: 1. Clausilia dubia dubiía Draparnaud, 1805 (SMF 163024a) 2. Clausilia dubia speciosa A. Schmidt, 1857 (SMF 163025a) 3. Clausilia dubia speciosa A. Schmidt, 1857 (SMF 163026a) 4. Clausilia dubia obsoleta A. Schmidt, 1857(SMF 163027a) EDLINGER: Shell morphology and variability in Clausilia dubía Draparnaud, 1805 Figure 4. Shell forms (7 stages from club-shaped -left- to spindle-shaped -right-). Figura 4. Formas de la concha (siete estadios desde forma de maza -izquierda- hasta abusada -derecha-). 5. Clausilia dubia huettneri Klemm, 1960 (SMF 1630248) 6. Clausilia dubia schlechti A. Schmidt, 1857 (SMF 163030a) 7. Clausilia dubia gracilior Clessin, 1887 (SMF 163031a) 8. Clausilia dubia tettelbachiana Ross- massler, 1838 (SMF 163032a) 9. Clausilia dubia otvinensis H. Gallens- tein, 1895 (SMF 163033a) 10. Clausilia dubia grimmeri L. Pfeiffer, 1848 (SMF 163034a) 11. Clausilia dubia floningiana Tscha- pek, 1886 (SMF 163035) 12. Clausilia dubia floningiana/gracilior (SMF 163036) 13. Clausilia dubia bucculenta Klemm, 1960 (Holotypus, SMF 163037) 14. Clausilia dubia runensis Tschapek, 1883 (SMF 163039a) 15. Clausilia dubia moldanubica Klemm, 1960 (Holotypus, SMF 163040) 16. Clausilia dubia kaeufeli Klemm, 1960 (Holotypus, SMF 163042) 17. Clausilia dubia alpicola Clessin, 1878 (SMF 31969) 18. Clausilia dubia reticulata Pini, 1883 (SMF 31936) The shells were measured under a bi- nocular microscope; the measurements were repeated three times. In the case of different results a special check was made. The height (H, Fig. 3) and the width (B, Fig. 3) of the shell as a whole, the height (MH, Fig. 3) and the width (MB, Fig. 3) of the aperture, the form of the aperture (MF, a series of 9 stages from pear-shaped to deltoid form, the angle between the spindle axis and the edge of the upper palatal (0.5 degree exactness), the mean of 5 rib distances (RA, Fig. 3) on the last whorl, and the number of whorls per shell (WZ, exact- ness: 0.25) were recorded. By compari- son with stencils the morphological cha- racters of the form of the shells (GH, a series of 7 stages from club-shaped, to extremely spindle-shaped specimens, Fig 4), the depth, and the thickness of the basal groove (BR, 6 stages, Fig. 5), the la- teral internal bulge (GW, on the left side of the aperture, 5 stages of thickness (Fig. 5), and the incision in the columellar la- mella (LF, 5 stages, Fig. 5) were recorded. The measured values of the follo- wing features were processed by a WIN- DOWS-EXCEL 5.0 and a WINDOWS SPSS 6.0 program (BROSIUS AND BROSIUS, 1995): - Mean of shell heights in each spot check - Standard deviation of shell heights in each spot check - Mean of shell heights in each spot check - Standard deviation shell heights in each spot check - Correlation (Pearson s) Coefficient of all 11 values: 99 Iberus, 15 (2), 1997 Figure 5. Basal groove (6 stages, upper row); lateral bulge (5 stages, middle row); incision in the columellar lamella (5 stages, lower row). Figura 5. Surco basal (6 estadios, arriba); protuberancia lateral (5 estadios, centro); incisión en la lame- la columelar (5 estadios, abajo). (R= Pearson“s Coefficient; N= number of cases; X, Y= variables; Sx, Sy= standard deviation of the varia- bles). By means of the WINDOWS SPSS 6.0 programs a factor extraction and a principle component analysis were exe- cuted. “Community” delivers informa- tion about the quota of spreading of one value that can be traced back to all other values. “Eigenvalue” is a value of the regression factors. It represents the quota of spreading of all values as interpreted by special regression factors. A reduction process restricts the numbers of factors in the final statistics to that exceeding 1.0. The factor matrix shows the influence of the regression factors on every variable as a percen- tage of 1. The measured variables of the samples in conjunction with the values of the specimens described by KLEmMM (1960) and the values of specimens of Clausilia dubia alpicola and C. d. reticulata were utilized for computing hierarchical clusters as dendrograms. For purpose of cluster analysis the measured values were transformed to “z-values”, values with a mean of 0 and a standard devia- tion of 1. Hierarchical clusters result 100 from dissimilarities computed on the basis of the sums of squared values of distances of each character. Thereby the spectrum of similarities and differences between all individuals of a spot check could be elaborated. The dendrograms contain specimens of various clusters according to their graduated similarity (BROSIUS AND BROSIUS, 1995). The formula of the general distances: D*= xvi) (D= distance; v= number of varia- bles; X, Y= cases) RESULTS Means of shell height and shell width: The means of the shell height and shell width differ in all sampling areas. The lowest value was found at the “Hohe Wand” region, the highest in the “Wienerwald” area. Comparisons of the means at different altitudes reveal that KLEMM's (1960) suggestion of a succes- sion of different shell heights (according to a succession of “races” resp. subspe- cies, high values at low altitudes, low values at high altitudes) is not generally convincing (Fig. 7). EDLINGER: Shell morphology and variability in Clausilia dubia Draparnaud, 18 Figure 6. Two spot-checks of the collection of the NHAMW (Naturhistorisches Museum, Wien). In the upper row “Clausilia dubia schlechti”, Inv. Nr. 11. 229 NHMW. The specimen in the upper row at the left belongs to Neostyriaca corynoides (Held, 1836). In the row below C. d. Eschlecht?”, Inv. Nr. 62. 348 NHMW. These spot-checks show us a high variability in the “subspecies”. Scale bar 1 cm. Figura 6. Dos muestras de la colección del NAMW (Naturbistorisches Museum, Wien). En la fila supe- rior Clausilia dubia schlechti ”, /nv. Nr. 11. 229 NAMW El especimen de la izquierda de la fila supe- rior pertenece a Niostyriaca corynoides (Held, 1836). En la fila inferior C. d. “schlechti”, lnv. Nr. 62. 348 NAMW. Las fotografías muetran una alta variabilidad en las subespecies. Escala 1 cm. Correlation coefficients: A very tude and the shell height (Table 1) but a remarkable outcome of the study was a high correlation between altitude and low positive correlation between the alti- the incicion in the columellar lamella. 101 Iberus, 15 (2), 1997 2000 — AA A se e. ——_—_—_—. == AS AA _ AAA 1000 El TAE dl A Na —.— ===> =0 == RRRAXáÁ —— —A— —e— 5 - a —r— —A— r ai” SYMB 1500m “+. <1000m GR e 1000-1500m vo >1500m . <1000m Figure 13. Two-dimensional scatter plots with factors 1/2 and 3/4 of a spot check of the Rax sam- ples, arranged in three groups in accordance with the altitude of the sample areas (less 1000 m, 1000-1500 m, more than 1500 m). Figura 13. Diagramas de puntos de dos dimensiones con factores 1/2 y 3/4 de las muestras de Rax , orde- nadas en tres grupos de acuerdo con la altitud de las áreas de muestreo (menos de 1000 m, 100-1500 m, más de 1500 m). (less than 1000m, 100-1500m, more than 1500m) results no altitude dependence of the factors. Cluster analysis: For the first cluster analysis the measured values were ta- ken from a group of specimens which consisted of selected samples from diffe- rent localities and altitudes; data from the SMF specimens were also included (Fig. 14). The hierarchical cluster which is presented as a dendrogram contains groups of different size which were co- llected at different localities. 111 Iberus, 15 (2), 1997 Notable is the isolated position of the SMF specimens of Clausilia dubia spe- ciosa and C. d. dubia which constitute a cluster of their own together with two specimens of the samples. Almost the same phenomena occur in dendrograms of samples collected in the areas “Hohe Wand”, “Schneeberg” and “Rax”. In these cases specimens of the SMF were also taken into consideration. In general the clusters show remar- kable segregations. Similarities of the individuals coming out from the same locality and appearing together in clus- ters may be seen as indication of close relationship. At the other side the spot checks from various areas are overlapping at a high degree. Relevant portions of indivi- duals, which present all the characters of several “subspecies” (subspecies seen in the traditional way) do not occur. It is remarkable that most specimens from the SMF which were considered to be typical for specific regions, appear also in the different branches of the dendro- grams and in various clusters. Only the SMF specimens of Clausilia dubia speciosa, and in a most astonishing way, the SMF specimen of C. d. dubia, an individual belonging to the nominotypi- cal subspecies” are seen at own separate branches of the dendrograms. This finding is in full agreement with the above mentioned scatter plots of the analysis of the main components. All three above mentioned specimens are characterized by strongly deviating measures and stand in isolated posi- tions. It is evident that this finding disa- grees with the geographical distribution shown in the literature (KLeMM, 1960). Wienerwald: The samples from the Wienerwald area come from altitudes between 270 and 400 m and sites similar in climate and ecological conditions (fir, pine, and mixed forests). The shells are rather similar and club shaped, but differ considerably in shell height and width, height and width of aperture, and distance of ribs. The same is true for all the other measures taken. In the hierarchical cluster analysis of samples from the Wienerwald area (Fig. 15) which was considered to be the type locality of Clausilia dubia dubia, the posi- tion of C. d. dubia, C. d. kaeufeli, and C. d. speciosa is found to be extremely isolated in a cluster of their own (Fig. 16). Within the second cluster also other SMF speci- mens appear in entirely isolated bran- ches. Only C. d. runensis, C. d. tettelba- chiana, C. d. grimmeri, C. d. schlechti and C. d. moldanubica appear in a branch together with the specimens of the samples from the Wienerwald area. Hohe Wand: Analyses of samples taken from the Hohe Wand (altitude between 560 m and 1080 m; Fig. 16) also lead to results which are not in accor- dance with generally held views. Clau- silia dubia speciosa and C. d. runensis are in an isolated position. They are in a cluster of their own together with one specimen of the local sample from H8. Also C. d. bucculenta, C. d. floningia- na/lgracilior, C. d. floningiana, C. d. reticu- lata, C. d. obsoleta, C. d. gracilior, C. d. (Right page). Figure 14. Hierarchical cluster of a spot-check of all samples and the SMEF specimens (Wi-y= specimens of the Wienerwald area; i= sample; y= number of the specimen; Hi-y= specimens of the Hohe Wand area; Si-y= specimens of the Schneeberg area; Ri-y= specimens of the Rax area; Cd-= specimens of the SME: du= dubia; sp= speciosa; ob= obsoleta; hn= huettneri, sc= schlechti, gr= gracilior, te= tettelbachiana, ot= otvinensis, gi= grimmeri, l= floningiana; flgr= floningianal gracilior, bu= bucculenta; ru= runensis, mo= moldanubica; ki= kaeufeli, al= alpicola; re= reticulata). (Página derecha). Figura 14. Cluster de todas las muestras y los especimenes SMF (Wi-y= especímenes del área de Wienerwald; ¡= muestra; y= número del especimen; Hi-y= especímenes del área de Wand; Si-y= especímenes del área de Schneeberg; Ri-y= especímenes del área de Rax; Cd-= especímenes SMF: du= dubia; sp= speciosa; 0b= obsoleta; /n= huettneri; sc= schlechti; gr= gracilior; te= tetrelbachiana; 01= otvinensis; g/= grimmeri, fl= floningiana, flgr= floningiana/gracilior; b4= bucculenta; 7u= runensis; mo= moldanubica; Rá= kaeufeli; 4/= alpicola; re= reticulata). 112 EDLINGER: Shell morphology and variability in Clausilia dubía Draparnaud, 180 Cdob Cdflgi Cdbu cafl Cdsp Cdsp W6 Cddu R4 32 Rescaled Distance Cluster Combine IE En - 1 EJE 113 Iberus, 15 (2), 1997 Rescaled Distance Cluster Combine CASE 0 5 10 15 20 Label Num —+-=-=--=-=-=--- Moscas MiS E O o == w1 6 wl 7 ia] wl1 3 w1 4 z W4 33 w1 5 E] wW2 12 W4 27 y] W3 23 w1 2 W4 30 poa W4 29 —— W3 24 Cdmo 73 Al wW3 18 op po W3 20 — W2 14 == wW4 3/9) — W3 119 — wW4 38 => W2 16 = w2 157) 4 — W3 22 == W4 31 W4 32 7 w4 28 — W4 34 a) W3 21 A Wa 36 —— WS 50 Á W3 25 == wW4 37 — > W4 35 — wW2 13 —— WS 42 WS 40 => W5 41 — HZ W2 15 ———— 7 w1 de WS 44 0 ¡FR WS 45 W5 48 — W6 52 WS 47 — 0) W6 a) == W6 56 — W6 53 W6 54 -=) WS 43 W4 26 === WS 46 W5 49 — HH W6 Sl A [EU wl1 8 w1 11 iS Wl 10 w Sparta Cdsc 64 =——, Cdgi 68 EIA ===) Cdte 66 Cdru 72 SIS y) W6 57 W6 58 A ¡E Cdflgi 70 => Cdbu NS EA E Cdga 65 Cdre 76 A Cdob 62 _—A Cdhn cp AS Cdot 67 Cdal IS pe Cdká 74 E EE cdf1l 69 Cddu 59 Cdsp 6l ] ' Cdsp 60 Ea Figure 15. Hierarchical cluster of a spot check of all samples of the Wienerwald area and the SMF specimens. Abbreviations as in Figure 14. | Figura 15. Cluster de todas las muestras del área de Wienerwald y los especímenes SME Abreviaturas como en la Figura 14. 114 EDLINGER: Shell morphology and variability in Clausilia dubía Draparnaud, 1805 Rescaled Distance Cluster Combine CASE 0 5 10 15 20 25 Label Num +--------- OS ASFALTO ERARIO a E H5 39 HS 43 HS 41 H5 42 H6-2 46 H6-4 48 H5 40 H2 9 H6- 3 47 H5 35 — Cdká 76 H4 24 Cdsc 66 H1 1 [asta H8-2 58 HS 34 H8-4 60 Cdhn 65 HS 44 H6-6 50 H7-25 53 H8-3 s9 H7-24 52 H7-23 51 —y H1 3 Cdmo 75 Tf) H3 14 H3 22 H1 5 H2 8 H7-27 55 H7-26 54 —— H4 26 ES do ¿E A H4 23 H4 25 H1 6 H2 10 H1 7 | H4 29 H3 19 H7-28 56 J H3 15 H3 18 H3 21 H5 38 H6-1 45 H2 12 H4 32 pitos H3 20 HS 36 ——— H4 27 H4 30 H1 2 H3 16 H3 17 H4 28 H2 11 H3 13 H5 37 F Cdgi 70 Cdot 69 Cdal 27 HS 33 ae Cdga 67 1] - Cdob 64 PR Cdre 78 Ccdfl 71 Cdflgi 72 Cdbu 73 E Cdsp 62 Cdsp 63 En _——— H8-1 57 Cddu 61 Ertasto de a] Figure 16. Hierarchical cluster of a spot check of all samples of the Hohe Wand area and the SMF specimens. Abbreviations as in Figure 14. Figura 16. Cluster de todas las muestras del área de Hohe Wand y los especímenes SME Abreviaturas como en la Figura 14. 115 Iberus, 15 (2), 1997 alpicola, C. d. otvinensis and C. d. grim- meri occur in separated branches and only C. d. tettelbachiana, C. d. moldanu- bica, C. d. huettneri and C. d. kaeufeli are integrated in branches together with the major part of the specimens from the Hohe Wand area. There are no indi- cations of the succession postulated by former workers (KLeMM, 1960) when one takes into consideration altitudes of the localities from which they came and the similarities that appear in the den- drogram. Schneeberg: The studied material was collected at altitudes between 700 to 1850 m. The cluster analysis (Fig. 17) reveals morphologically isolated posi- tions for Clausilia dubia speciosa, C. d. dubia, C. d. kaeufeli, C. d. floningiana, C. d. reticulata, C. d. obsoleta, C. d. bucculenta, C. d. floningiana / gracilior, C. d. alpicola, C. d. otvinensis, C. d. tettelbachiana, C. d. runensis and C. d. moldanubica. Only C. d. grimmeri, C. d. gracilior and C. d. schlechti occur in branches together with most of the specimens of the Schneeberg spot checks. Remarkable is that specimens with close morphological relations to C. d. kaeufeli, as expected for the peak of the mountain, don't occur in the clus- ters. The SMF specimen of C. d. kaeufeli is isolated in the dendrogram. Rax: The studied material was collected at altitudes between 700 to 1685 m. As in the Clusters analyses of the other areas Clausilia dubia dubia, and C. d. speciosa are isolated in a Cluster of ist own (Fig. 18). Also C. d. kaeufeli, C. d. floningiana, C. d. bucculenta, C. d. flonin- giana / gracilior, C. d. runensis, C. d. alpi- cola, C. d. otvinensis, C. d. otvinensis, C. d. obsoleta, C. d. reticulata and C. d. gracilior occur very isolated in branches together with only few specimens of the local samples. Only C. d. grimmeri, C. d. mol- danubica, C. d. tettelbachiana, C. d. sch- lechti and C. d. huettneri can be seen as well integrated in clusters with the major part of the local samples. No remarkable position of C. d. kaeufeli or a succession as suggested by Klemm was visible. 116 DISCUSSION AND CONCLUSION The conclusions presented here must be seen as being valid only for samples of Clausilia dubia collected in restricted areas. Only further research might lead to conclusive results on possible geo- graphic variation in C. dubia. The results of the present research as well as a critical evaluation of those of earlier researchers show us the limits inherent the intuitive, subjective method used in the analysis of characters. Earlier studies by EDLINGER AND MILDNER (1979) on Clausilia dubia in Carinthia, using traditional morphologi- cal methods based on a number of shell characters, also showed a high variabi- lity of most of these characters within each population. This may be a common phenomenon when analyses are based on a large number of characters. In this Case, techniques of measurement, as well as applied statistical methods might present potential sources of error. Nevertheless, all observations are indicative of high variability within the species. Similar observations can be made in many of the collection samples (Fig. 6). This can be clearly discerned, in spite of the fact that many of these samples have been classified as belon- ging to various subspecies. To gain a better understanding of shell variability, we must also consider the influence of ecological factors, and the life history of the specimens. Morp- hological features may be influenced by non hereditary factors too (GOODFRIEND, 1986). Against the background of low correlation coefficients between altitude and most characters, except columellar lamella, altitude in itself cannot be con- ceived as an substantial ecological factor, because the measures of the shells except the columellar lamella don't vary significantly in correlation with altitude. A reason for the remarka- ble correlation of altitude with the colu- mellar lamella might be that the Clausi- lia dubia shells of the Wienerwald area commonly have very pregnant incisions of the columellar lamella. EDLINGER: Shell morphology and variability in C/ausilia dubia Draparnaud, 1805 Rescaled Distance Cluster Combine s8 Cdmo Cdru Cdte s10 Cdot Cdal Cdflgi Cdbu Cdob Cdre Ccdf1 Cdká Cddu Cdsp Cdsp Num 26 29 25 33 2 34 21 4 5 12 17 22 35 41 39 6 36 1 37 14 42 19 38 10 18 27 30 8 11 2 13 15 32 40 9 20 66 16 28 65 23 24 67 70 50 51 53 52 31 55 57 56 58 60 54 47 48 46 3 45 49 43 44 75 74 68 59 69 77 72 73 64 78 7/3 76 61 63 62 0 5 10 15 20 25 00 3 a] B| +] al gol ll ejes ==. UTE J Ip uu Figure 17. Hierarchical cluster of a spot check of all samples of the Schneeberg area and the SMF specimens. Abbreviations as in Figure 14. Figura 17. Cluster de todas las muestras del área de Schneeberg y los especímenes SME Abreviaturas como en la Figura 14. 117 Iberus, 15 (2), 1997 Rescaled Distance Cluster Combine DIU SBPBRRA An e VwWWWVWVWA NN POUOWWVWWWW ny >bBIAOIWADIOBONIW R7 15 R10 20 R13 31 R14 44 R14 46 R14 47 = R14 41 R14 42 R14 43 R15 50 R15 Sp R7 Y aa R7 13 R10 21 R14 38 R12 23 JA R13 29 R12 24 NA R12 25 EE cal E RS 6 Cdhn 60 a) R7 el R7 12 R7 9 a R14 40 Cdsc 61 R7 19 Cdte 63 R14 45 R14 49 R14 48 — Cdmo 70 Cdgi 65 RS 2 R7 17 R11 22 Cdga 62 Cdre ve ) Cdob 59 Cdot 64 Cdal 72 AN Cdru 69 R4 1 Cdflgi 67 AS pa cdf1 66 Cdká 71 Cdsp 57 Cdsp 58 Cddu 56 Figure 18. Hierarchical cluster of a spot check of all samples of the Rax area and the SMF speci- mens. Abbreviations as in Figure 14. Figura 18. Cluster de todas las muestras del área de Rax y los especímenes SME Abreviaturas como en la Figura 14. 118 EDLINGER: Shell morphology and variability in Clausilia dubiía Draparnaud, 1805 Other ecological factors may be of relevance, but we must consider them as arranged in ensembles and affecting upon animals collectively. Certainly, individual living condi- tions, as well as hereditary dispositions, result in specific modifications of cha- racters. nevertheless we can reject the hypothesis of EDLINGER AND MILDNER (1982) that the characteristics of Clausilia dubia runensis might be a result of hel- minth parasitism. A representation of variability of species or populations by character analysis also has to take into considera- tion the fact that many characters act as correlated variables. In these cases one can speak of character complexes. Arrangements in complexes restrict variability. The question arises, whether there are other factors influencing the variables mentioned above, which are not subjects of the researches presented above. Shells, for example, function as hard skeletons and as mechanical abut- ments for musculature. Hard skeleton and musculature must be suited to each other. So it is evident, that morphologi- cal relations between various measures of the shell are indirectly caused by constructional needs of the animal as a whole and also of the soft body in parti- cular (GUTMANN, 1989; EDLINGER, 1991). Considering the local patterns of dis- tribution of values and primary compo- nents inside the species and its popula- tions, these patterns must be seen as varying locally and gradually. The variation of patterns must be seen as the result of local variation, following from step by step changes of frequencies of characters. When different frequencies are developed under similar ecological conditions, we may assume that these changes are a result of genetic drift. Certain local accumulation of special characters resp. values of variables may result from genetic drift too. Typical characters of the so called “Clausilia dubia runensis”, C. d. speciosa (shell form) or “C. d. otvinensis” (distances between shell ribs), for example, occur in sepa- rate areas with large distances between them. Other characters of the same animals very often don't differ from that of surrounding populations in adjacent areas (EDLINGER AND FISCHER, 1997). This leads us to the conclusion that frequencies of special characters cannot automatically be taken as a reason for a common origin of separated popula- tions resp. of a heterogeneous origin of contiguous populations. Above all this is true, when other characters are identi- cal with those of contiguous popula- tions. So it may be that genetic drift or special environmental factors have an influence on single characters. With regard to these characters we may con- ceive of some populations as homoge- nous and “pure bred”. At the other side there cannot arise populations being homogeneous and “pure bred” in all or most characters and containing such a high number of “typical specimens” homogenous in most or in all characters by simple environmental influences. In any case, “typical” or “pure bred” specimens were recognized mostly by earlier researchers, and are extremely arbitrary. So, the natural populations of Clausilia dubia which were investigated do not match the preconceived expecta- tions of well established races or subs- pecies occurring in well delimited areas with rare interbreeding occurrences. Therefore, we must question, if a morp- hologically uniform population of Clau- silia dubia which can be defined as a subspecies or race, might ever have existed in the eastern Alpine region. KLEMM (1960) who was of the same opinion, believed that the transitional stages between the races and subspecies might be the consequence of (post glacial) re-immigration of pure bred populations, and their subsequent mixing with other local forms. Shell characters, and their distribu- tion patterns do not support this hypot- hesis. Additionally, cluster analysis shows that very variable samples come from regions where the forerunners of present populations must definitely have lived and survived during the Pleistocene. Contrary to KLemMM (1960), we must state that a reconstruction of 119 Iberus, 15 (2), 1997 events after the Pleistocene gives us no evidence to support this view. Why should areas, glaciated during the Pleis- tocene be resettled by the descendants of single, pure bred populations after the disappearance of the ice? A new interpretation of character distribution seems to be more adequate for the case presented, and discussed here. This interpretation requires the (for the case discussed here) use of theo- rems which are accepted by most anth- ropologists (CAVALLI-SFORZA, MENOZZI AND PIAZZA, 1994; KLEIN, TAKAHATA AND AYALA, 1994). According to their theoretical guide- lines, the definition of subspecies and race depends on subjective argumenta- tions, and does not mirror objective, REFERENCES BROSIUS, G. AND BROSIUS, F., 1995. SPSS Base Sys- tem und Professional Statistics. International Thomson Publishing, Bonn-Albany, 1002 pp. CAVALLI-SFORZA, L. L., MENOZZI, P. AND PIAZZA, A., 1994. The History and Geography of Human Genes. Princeton, NJ. EDLINGER, K., 1991. The mechanical constraint's in mollusc constructions. The function of the shell, the musculature, and the connective tis- sue. In Schmidt-Kittler, N, and Vogel, K. (Eds.): Constructional morphology and evolution. Springer, Berlin, Heidelberg, New York. EDLINGER, K. AND MILDNER, P., 1979. Mono- graphie der in Kárnten lebenden Clausilia dubia-Rassen. Carinthia, 11/171: 251-266. EDLINGER, K. AND MILDNER, P., 1982. Trema- todenbefall bei Clausilia dubia runensis Ts- chapek, 1883 aus Kárnten. Carinthia, 11/172: 319-324. EDLINGER, K. AND FISCHER, W., 1997. Clausilia dubia bucculenta Klemm, 1960. Zur statistis- chen Realitát einer Unterart. Club Conchylia Informationen, 29 (1/2): 11-25. FECHTER, R. AND FALKNER, G., 1990. Weichtiere. Europaische Meeres- und Binnenmollusken. Mo- saik-Verlag, Múnchen, 287 pp. GOODFRIEND, G. A., 1986. Variation in land- snail shell form and size and its causes: a re- view. Systematic Zoology, 35(2): 204-223. GUTMANN, W. F., 1989. Die Evolution hydraulis- cher Konstruktionen. Organismische Wandlung statt altdarwinistischer Anpassung. W. Kra- mer, Frankfurt /M, 200 pp. 120 concrete facts. They also accept diversity within given populations. Therefore, distribution and frequency of characters are subject to constant change. Thus, in any investigated snail population, only the distributions of characters, and the frequency of their occurrence can be reasonably recorded. This entirely altered interpretation also corresponds better with the results of genetic analysis in various species of Albinaría (MYLONAS, KRIMBAS, TSIAKAS AND AYOUNTANTI, 1990; SCHILTHUIZEN, 1994). The results of these studies show that there is an overlap of genetic featu- res in some of the traditionally recogni- zed subspecies, and even of species. Similar results may be expected for Clausilia dubia. HOLOYAK, D. T. AND SEDDON, M. B., 1988. Sys- tematics and biogeography of some western European Clausilia (Gastropoda: Clausilii- dae). Journal of Conchyology, 33: 55-80. KEARNEY, M. P., CAMERON, R. AND JUNGBLUTH, J., 1983. Die Landschnecken Nord und Mitte- leuropas. P. Parey-Verlag, Hamburg-Berlin, 387 pp. KLEIN, J]., TAKAHATA, N. AND AYALA, F. J., 1994. MHC-Polymorphismus und Ursprung des menschen. Spektrum der Wissenschaft, 2: 56-62. KLEmMM, W., 1960. Clausilia dubia und ihre for- men in Osterreich. Archiv fúr Molluskenkunde, 89: 81-109. KLeMM, W., 1973. Die Verbreitung der rezen- ten Land-Geháuse-Schnecken in Osterreich. - Denkschriften der Osterreichischen. Aka- demie der Wissenschaften, mathematisch-natur- wissenschaftliche Klasse, 117: 1-503. KOTHBAUER, H., NEMESCHKAL, H. L., SATT- MANN, H. AND WAWRA, E., 1991. Uber den Aussagewert von Typen und qualitativen Aufsammlungen: Eine kritische Sicht am Beispiel von Artanta arbustorum styriaca (Frauenfeld, 1868) (Pulmonata: Helicidae). Annalen des Naturhistorischen, 92 B: 229-240. MAayr, E., 1967. Artbegriff und evolution. P. Pa- rey, Hamburg, 617 pp. MYLONAS, M., KRIMBAS, C., TSIAKAS, S. AND AYOUNTANTI, A., 1990. The genus Albinaria Vest. (Clausiliidae, Gastropoda). Is there any true species?. Biología Gallo-hellenica, 13: 161- 164. EDLINGER: Shell morphology and variability in Clausilia dubiía Draparnaud, 1805 NEMESCHKAL, H. L., 1990. Uber die form der sch- neckenschale: morphometrische grundlagen und vorbereitungen fúr ein statistisches ta- xon-Modell. Zoologisches Jahrbuch Systema- tik, 117: 491-534. NEMESCHKAL, H. L., 1991. Character coupling for taxa discrimination: a critical appraisal of quadratic assignment procedures (QAP). Zeitschrift fur zoologische Systematik und Evo- lutionsforschung, 29: 87-96. NEMESCHKAL, H. L., 1993. Character coupling in land snails (Arianta, Helicidae). Does Man- telís test detect equality or similarity?. Zoo- logischer Anzeiger, 230(1/2): 1-17. NEMESCHKAL, H. L. AND KOTHBAUER, H., 1988. Arianta arbustorum alpicola (Ferussac, 1819) (Pulmonata, Helicidae). Uber Interpretation und Realitát eines Taxon. Zoologischer An- zeiger, 221(5/6): 343-354. NORDSIECK, H., 1990. Revision der Gattung Clausilia Draparnaud, besonders der Arten in SW-Europa (das Clausilia rugosa-Problem). Archiv fúr Molluskenkunde, 119 (416): 133-179. OSCHE, G., 1994. Rasse. In Becker, U., Ganter, S., Jost, C. and Sauermost, R. (Eds.): Lexikon der Biologie 7. Spektrum Verlag, Heidelberg- Berlin-Oxford: 82-83. SCHILTHUIZEN, M., 1994. Differentiation and hybridization in a polytypic snail. Den Haag. 178 pp. SUDHAUS, W. AND REHFELD, K., 1992. Einfúhrung in die Phylogenetik und Systematik. G. Fischer, Stuttgart-Jena-New York, 241 pp. THORPE, R. S., 1987. Geographic variation. a synthesis of cause, data, pattern and con- gruence in relation to subspecies, multivariate analysis and ohylogenesis. Bollettino Zoologico, 54: 3-11. 121 . lo Pad cn nbiegas 0 0 M po De mb pese Ne se E Ds Lo Qu NE 1 ganes | dl RUBIA pee 2092 e pe RBinanseral vola a E AE NORMAS DE PUBLICACIÓN e La revista IBERUS publica artículos de fondo, notas y monografías que versen sobre cualquiera de los aspectos rela- cionados con la Malacología. Se entiende por artículo un trabajo de investigación de más de 5 páginas de texto, mecanografiadas, incluidas láminas, gráficos y tablas. 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Se recomienda reducir el número y extensión de ilustraciones, láminas o tablas al mínimo necesario. * Los artículos que no se ajusten a las normas de publicación serán devueltos al autor con las indicaciones de los cam- bios necesarios. » El Comité Editorial comunicará al autor responsable del trabajo la fecha de recepción del trabajo y la fecha de envío a revisión. Cada original recibido será sometido a revisión por al menos dos investigadores. El Comité Editorial, a la vista de los informes de los revisores decidirá sobre la aceptación o no de cada manuscrito. El autor recibirá en cada caso copia de los comentarios de los revisores sobre su artículo. En caso de aceptación, el mismo Comité Editorial, si lo considera conveniente, podrá solicitar a los autores otras modificaciones que considere oportunas. Si el trabajo es aceptado, el autor deberá enviar una copia impresa del mismo corregida, acompañada por una versión en disco flexi- ble (diskette), utilizando procesadores de texto en sus versiones de DOS o Macintosh. La fecha de aceptación figura- rá en el artículo publicado. + Las pruebas de imprenta serán enviadas al autor responsable, EXCLUSIVAMENTE para la corrección de erratas, y deberán ser devueltas en un plazo máximo de 15 días. Se recomienda prestar especial atención en la corrección de las pruebas. » De cada trabajo se entregarán gratuitamente 50 separatas. Aquellos autores que deseen un número mayor, deberán hacerlo constar al devolver las pruebas de imprenta, y NUNCA POSTERIORMENTE. El coste de las separatas adi- cionales será cargado al autor. INSTRUCTIONS TO AUTHORS » 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: Dr. Ángel Guerra Sierra, Editor de Publi- caciones, Instituto de Investigaciones Marinas (CSIC), C/Eduardo Cabello 6, 36208 Vigo, 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. When a paper has joint authorship, one author must accept responsability for all correspondence. * Papers should conform the following layout: First page. This must include a concise but informative title, with mention of family of higher taxon when appropriatte, and its Spanish translation. It will be followed by all authors' names and surnames, their full adress(es), an abstract (and its Spanish translation) not exceeding 200 words which summarizes not only contents but results and conclusions, and a list of Key Words (and their Spanish translation) under which the article should be indexed. Following pages. These should content the rest of the paper, divided into sections under short headings. Whenever pos- sible the text should be arranged as follows: Introduction, Material and methods, Results, Discussion, Conclusions, Acknowledgements and References. Unusual abbreviations used in the text must be grouped in one alphabetic sequence after the Material and methods section. * Notes should follow the same layout, without the abstract. e Footnotes and cross-references must be avoided. The International Codes of Zoological and Botanical Nomencla- ture must be strictly followed. The first mention in the text of any taxon must be followed by its authority including the year. In systematic papers, when synonyms of a taxon are given, they must be cited IN FULL, including the perio- dical, in an abbreviate form, where they were described, and the type localities in square brackets when known. Follow this example (please note the punctuation): Dendrodoris limbata (Cuvier, 1804) Synonyms Doris limbata Cuvier, 1804, Ann. Mus. H. N. Paris, 4 (24): 468-469 [Type locality: Marseille). Doris nigricans Otto, 1823, Nov. Act. Ac. Caes. Leop. Car., 10: 275. These references must not be included in the Bibliography list, except if referred to elsewhere in the text. I£a full list of references of the taxon is to be given immediately below it, the same layout should be followed (also excluding those nowhere else cited from the Bibliography list). Only Latin words and names of genera and species should be underlined once or be given in ¿talics, No word must be written in UPPER CASE LETTERS. SI units are to be used, together with their appropriate symbols. In Spanish manuscripts, decimal numbers must be separated with a comma (,), NEVER with a point (.) or upper comma ('). » References in the text should be written in small letters or SMALL CAPITALS: Fretter 82 Graham (1962) or FRETTER 82 GRAHAM (1962). The first mention in the text of a paper with more than two authors must include all of them [Smith, Jones 82 Brown (1970)], thereafter use et al. [Smith et al. (1970)]. Ifan author has published more than one paper per year, refer to them with letters: (Davis, 1989a; Davis, 1989b). Avoid op. cit. The references in the reference list should be in alphabetical order and include all the publications cited in the text but only these. ALL the authors of a paper must be included. These should be written in small letters or SMALL CAPITALS. The references need not be cited when the author and date are given only as authority for a taxonomic name. Titles of periodicals must be given IN FULL, not abbreviated. For books, give the title, name of publisher, place of publication, indication of edition if not the first and total number of pages. Keep references to doctoral theses or any other unpu- blished documents to an absolute minimum. See the following examples (please note the punctuation): Fretter, V. and Graham, A., 1962. British Prosobranch Molluscs. Ray Society, London, 765 pp. Ponder, W. F., 1988. The Truncatelloidean (= Rissoacean) radiation - a preliminary phylogeny. In Ponder, W. F. (Ed.): Prosobranch Phylogeny, Malacological Review, suppl. 4: 129-166. Ros, J., 1976. Catálogo provisional de los Opistobranquios (Gastropoda: Euthyneura) de las costas ibéricas. Miscelá- nea Zoológica, 3 (5): 21-51. + Figures must be original, in Indian ink on draughtsman's tracing paper. Keep in mind page format and column size when designing figures. These should be one column (57 mm) or two columns (120 mm) wide and up 194 mm high, or be proportional to these sizes. Two columns format is recomended. It is desirable to print figures with their legend below, so authors are asked to take this into account when preparing full page figures. If computer generated graphics are to be included, they must be printed on high quality white paper with a laser printer. Photographs must be of good contrast, and should be submitted in the final size. When mounting photographs in a block, ensure spacers are of uni- form width. Remember that grouping photographs of varied contrast results in poor reproduction. Take account of necessary reduction in lettering drawings; final lettering must be at least 2 mm high. In composite drawings, each figure should be given a capital letter; additional lettering should be in lower-case letters. A scale line is recomended to indi- cate size, magnification ratio must be avoided as it may be changed during printing. UTM maps are to be used if neces- sary. Figures must be submitted on separate sheets, and numbered with consecutive Arabic numbers (1, 2, 3,...), without separating Plates' and “Figures”. Legends for Figures must be typed in numerical order on a separate sheet, and an English 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, 300 US$ per page). They should be submitted in the same way that black and white prints. + Tables must be numbered with Roman numbers (1, II, IL...) and each typed on a separate sheet. Headings should be typed on a separate sheet, together with their English translation. Complex tables should be avoided. As a general rule, keep the number and extension of illustrations and tables as reduced as possible. + Manuscripts that do not conform to these instructions will be returned for correction before reviewing. + Authors submitting manuscripts will receive an acknowledgement of receipt, including receipt date, and the date the manuscript was sent for reviewing. Each manuscript will be critically evaluated by at least two referees. Based of these evaluations, the Editorial Board will decide on acceptance or rejection. Anyway, authors will receive a copy of the refe- rees” comments. If a manuscript is accepted, the Editorial Board may indicate additional changes if desirable. Accep- table manuscripts will be returned to the author for consideration of comments and criticism; a finalized manuscript must then be returned to the Editor, together with a floppy disk containing the article written with a DOS or Macin- tosh word processor. Dates of reception and acceptance of the manuscript will appear in all published articles. » Proofs will be sent to the author for correcting errors. At this stage no stylistic changes will be accepted. Pay special attention to references and their dates in the text and the Bibliography section, and also to numbers of Figures and Tables appearing in the text. e Fifty reprints per article will be supplied free of charge. Additional reprints must be ordered when the page proofs are returned, and will be charged at cost. NO LATER orders will be accepted. 2 S y " e J 1. y d ¡ 1 se - o y ” “ ol W Ñ My m0) ha A » CIA E ) EI é e MESE: EEN A: dei DE DA AE IS Mail Ab Mide 7 ELN PM a y ie de IGAC Ad Un is e > LA SociEeDAD ESPAÑOLA DE MALACOLOGÍA Junta directiva desde el 18 de octubre de 1996 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 Ángel Guerra Sierra Instituto de Investigaciones Marinas, c/ Eduardo Cabello 6, 36208 Vigo, España Bibliotecario Rafael Araujo Armero Museo Nacional de Ciencias Naturales, CSIC, c/ José Gutierrez Abascal 2, 28006 Madrid, España Vocales Eugenia María Martínez Cueto-Felgueroso María de los Ángeles Ramos Sánchez Francisco Javier Rocha Valdés Gonzalo Rodríguez Casero 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 1997: Socio numerario (en España): 5.000 ptas. (= 50 U.S. $) (en extranjero): 7.000 ptas (= 70 U.S. $) Socio estudiante: 2.000 ptas. (= 20 U.S. $) Socio Familiar: 500 ptas. (= 5 U.S. $) Socio Protector: 6.000 ptas. (= 60 U.S. $) (mínimo) Socio Corporativo 6.000 ptas. (= 60 U.S. $) INSCRIPCIÓN: 1.000 ptas. (= 10 U.S. $) 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. Members living in foreing countries can deduce 10 U.S. $ 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 15 (2) 1997 MALHAM, S. K., RUNHAM, N. W, AND SECOMBES, C. J. Phagocytosis by haemocytes from the Lesser Octopus Eledone cirrhosa Fagocitosis en hemocitos del pulpo blanco Eledone cirrhosa........ooooocooccooccooconocooo: 1-11 PÉREZ, A. M. AND LÓPEZ, A. New data on the morphology and the distribution of Bulimulus corneus Sowerby, 1833 (Gastropoda: Pulmonata: Orthalicidae) in Nicaragua Nuevos datos sobre la morfología y la distribución de Bulimulus corneus Sowerby 1833 (Gastro- pora; Dulmonatas Orthalicidae) en INTIAgUa a a on ade 13-24 LAZARIDOU-DIMITRIADOU, M. AND SGARDELIS, S. Phenological patterns and life history tactics of Helicoidea (Gastropoda, Pulmonata) snails from Northern Greece Patrones fenológicos y estrategias de vida en Helicoidea (Gastropoda, Pulmonata) del Norte de A O O A a A UE ad 25-34 SALVINI-PLAWEN, L. VON. Fragmented knowledge on West-European and Iberian Caudofoveata and Solenogastres Conocimiento fragmentado de los Solenogastros y Caudofoveados de Europa occidental y Península IA RA AN OD nd E 35-50 EDLINGER, K. AND GUTMANN, W. E Molluscs as evolving constructions: necessary aspects for a dis- cussion of their phylogeny Los moluscos como construcciones en evolución: aspectos necesarios para una discusión sobre su VILA A E A ORO OOO do 51-66 FÚKOH, L. Mollusc fauna of the medium high mountain ranges of the Hungarian Holocene: a zoo- geographical research Fauna de moluscos de media y alta montaña del Holoceno de Hungría: una investigación zo0ge- ORVAES ao E SUS da ERE Coalo AUN ERA AS 67-74 COBBINAH, J. R. Aestivation responses of three populations of the giant African snail, Achatina acha- tina Linne (Gastropoda: Achatinidae) Respuestas a la estivación de tres poblaciones del caracol gigante africano Achatina achatina Linne UCA OPOda: AC Uaae) sl ira Ed an aee e e o DD SOON 75-82 BABA, K. AND BAGL I. Snail communities associated to swampy meadows and sedgy marshy meadows plant communities of the Great Hungarian Plain Comunidades de moluscos asociadas a comunidades vegetales de praderas pantanosas y junqueras ena Gran Llanura AUR Cara o dinos So Ue de DUNN rai Se CSIC EE eE 83-93 EDLINGER, K. Morphological and biometrical researches on Austrian Clausiliids. Shell morphology and variability in Clausilia dubia Draparnaud, 1805 Investigaciones biométricas y morfológicas en Clausilidos de Austria. Morfología y variabilidad de la concha de Clausilia dubia Draparnaud, 1805... 95-121 ISSN 0212-3010 a , AS 1 < K y Y E F a Es E > y y . E 1d ERNST MAYR LIBRARY NR 3 2044 110 337 PEA A PRO A ASNO AI o e e ia ia S E . z S AS ren ro dl A - Sy a eo rr A ps A OS : SN z a IS ETS PIS : CAN . : : ; aaron 2 O er raras - TO eo CNEA EN SENS a Nota SER OS ES o a E : : A OS Ñ g Pa 4 Ed z EEES aa : E dez 4 O paro PAE, > 7 E E E z E CIA ] 29. E, EA ENTE Edel E IN E . roo Daria o a AA