6X-, ¡,. HÚ¡ ¡ Í¿1'~> m^evus Vol. 26 (1) REVISTA DE LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA Oviedo, junio 2008 Iberas Revista de la Sociedad Española de Malacología Comité de Redacción (Board of Editors) Editor di Publicaciones (Editor-in-Chief) Serge Gofas Universidad de Málaga, España Director de Redacción (Exkutive Editor ) Gonzalo Rodríguez Casero Apdo. 1 56, Mieres del Camino, Asturias, España Editora Ejecutiva (Managimg Editor) Eugenia Mg Martínez Cueto-Felgueroso Apdo. 1 56, Mieres del Camino, Asturias, España Editores Adjuntos (Associate editors) Francisco Javier Conde de Saro Embajada de España, Japón Benjamín Gómez Moliner Universidad del País Vasco, Vitoria, España Ángel Antonio Luque del Villar Universidad Autónoma de Madrid, Madrid, España Emilio Rolán Mosquera Universidad de Vigo, Vigo, España José Templado González Museo Nacional de Ciencias Naturales, CSIC, Madrid, España Jesús S. Troncoso Universidad de Vigo, Vigo, España Comité Editorial (Board of Reviewers) Kepa Altonaga Sustacha Universidad del País Vasco, Bilbao, España Eduardo Angulo Pinedo Universidad del País Vasco, Bilbao, España Rafael Araujo Armero Museo Nacional de Ciencias Naturales, Madrid, España Thierry Backeljau Instituí Royal des Sciences Naturelles de Belgique, Bruselas, Bélgica Rüdiger Bieler The Field Museum, Chicago, Estados Unidos Sigurd v. Boletzky Laboratoire Arago, Banyuls-sur-Mer, Francia José Castillejo Murillo Universidad de Santiago de Compostela, Santiago de Compostela, España Karl Edlinger Naturhistorisches Museum Wien, Viena, Austria Antonio M. de Frías Martins Universidade dos Acores, Acores, Portugal José Carlos García Gómez Universidad de Sevilla, Sevilla, España Gonzalo Giribet de Sebastián Harvard University, EE.UU. Edmund Gittenberger National Natuurhistorisch Museum, Leiden, Holanda Ángel Guerra Sierra Instituto de Investigaciones Marinas, CSIC, Vigo, España Gerhard Haszprunar Zoologische Staatssammlung München, München, Alemania Yuri 1. Kantor A.N. Severtzov Institute of Ecology and Evolution, Moscú, Rusia María Yolanda Manga González Estación Agrícola Experimental, CSIC, León, España Jordi Martinell Callico Universidad de Barcelona, Barcelona, España Ron K. O'Dor Dalhousie University, Halifax, Cañada Takashi Okutani Nihon University, Fujisawa City, Japón Marco Oliverio Universitá di Roma "La Sapienza", Roma, Italia Pablo E. Penchaszadeh Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Buenos Aires, Argentina Winston F. Ponder Australian Museum, Sydney, Australia Carlos Enrique Prieto Sierra Universidad del País Vasco, Bilbao, España M^ de los Ángeles Ramos Sánchez Museo Nacional de Ciencias Naturales, CSIC, Madrid, España Francisco Javier Rocha Valdés Instituto de Investigaciones Marinas, CSIC, Vigo, España Paul G. Rodhouse British Antarctic Survey, Cambridge, Reino Unido Joandoménec Ros i Aragonés Universidad de Barcelona, Barcelona, España María Carmen Salas Casanovas Universidad de Málaga, Málaga, España Gerhard Steiner Instituí für Zoologie der Universitát Wien, Viena, Austria Victoriano Urgorri Carrasco Universidad de Santiago de Compostela, Santiago de Compostela, España Anders Warén Swedish Museum of Natural History, Estocolmo, Suecia Portada de Iberus Iberus gualtieranus (Linnaeus, 1758), una especie emblemática de la península Ibérica, que da nombre a la revista. Dibujo realizado por José Luis González Rebollar “Toza”. Iberus REVISTA DE LA SOCIEDAD ESPAÑOLA DE MALACOLOGÍA Vol. 26 (1) Oviedo, junio 2008 Iberus Revista de la Sociedad Española de Malacología Iberus publica trabajos que traten sobre cualquier aspecto relacionado con la Malacología. Se admiten también notas breves. Iberus edita un volumen anual que se compone de dos o más números. Instrucciones para los autores Los manuscritos deben remitirse a: Serge Gofas, Editor de Publicaciones, Departamento de Bio- logía Animal, Universidad de Málaga, Campus de Teatinos, s/n, 29071, Málaga, España. Los trabajos se entregarán por triplicado (original y dos copias). Se recomienda a los autores leer cuidadosamente las normas de publicación que se incluyen en cada número de la revista. SUBCRIPCIONES Iberus puede recibirse siendo socio de la Sociedad Española de Malacología, en cualquiera de sus formas, o mediante intercambio. Aquellos socios que deseen adquirir números atrasados deberán diri- girse al bibliotecario. Los no socios deberán ponerse en contacto con BACKHUYS PUBLISHERS, P.O. Box 321, 2300 AH Leiden, The Netherlands. Tel.: +31-71-31 70 208, Fax: +31-71-31 71 856, Correo Elec- trónico: backhuys@euronet.nl Los resúmenes de los artículos editados en esta revista se publican en Aquatic Science and Fisheries Abstracts (ASFA) y en el Zoological Records, BIOSIS. Contents list published in Aquatic Science and Fisheries Abstracts and Zoological Records , BIOSIS. Dep. Leg. B-43072-81 ISSN 0212-3010 Diseño y maquetación: Gonzalo Rodríguez Impresión: LOREDO, S. L. - Gijón © Sociedad Española de Malacología Iberas, 26 (1); 1-4, 2008 "(w Re-discovery of living specimens of Heliacus ( Gyriscus ) jef- freysianus (Tiberi, 1867) (Gastropoda: Architectonicidae) Redescubrimiento de ejemplares vivos de Heliacus ( Gyriscus ) jef- Jreysian i/s (Ti be ri , 1867) (Gastropoda: Architectonicidae) Constantine MIFSUD* and Panayotis OVALIS** Recibido el 17-VII-2007. Aceptado 9-X-2007 ABSTRACT Heliacus (Gyriscus) ¡effreysianus (Tiberi, 1 867), a rare architectonicid species associated with Corallium rubrum (L., 1758), has been rediscovered alive off Crete, eastern Mediter- ranean. RESUMEN Heliacus ( Gyriscus ) ¡effreysianus (Tiberi, 1 867), una especia rara de architectonicido aso- ciado a Corallium rubrum (Linnaeus, 1 758), ha sido redescubierta con ejemplares vivos, frente a la isla de Creta en el Mediterráneo oriental. KEY WORDS: Mollusca, Gastropoda, Architectonicidae, Heliacus j effreysianus, Crete. PALABRAS CLAVE: Mollusca, Gastropoda, Architectonicidae, Heliacus jeffreysianus, Creta. INTRODUCTION Heliacus ( Gyriscus ) jeffreysianus was described by Tiberi (1867) from three specimens collected from a red coral substratum off Sardinia. He described the species a year after its discovery, as stated by Coen (1932). SYSTEMATICS Original description: Cochlea turbinata, elato-conica, turrita, modice umbilicata, inteo- fulvescens; apex obtusiusculus, laevigatus, vértice intorto, subperf orato, spiraliter invo- luto; anfr. 7 convexi, sutura profunda divisim transverse cingulati, cingulis confortis, alter- natim majoribus (num.fere 15 totidemque minaribus in ultima anfractu; num. 6 in penúltimo), eleganter granulosis, submonili- formibus; anfr. Ultimus rotundatus, subin- flatus, basi paululum depressus; umbilicus mediocris, pervius, superne circolariter crenu- latus; apert. Subcircularis, effusa, intus haud margaritacea, marginibus acutis, callo pari- etali junctis; margine columellari sinuato, reflexo, umbilici partem occultante. - Diam. Maj. 9, min. 8, alt 10? mili. - Operculum corneum, superne núcleo centrali depresso lamellaque erecto-crenulata multispiratum, inferné processu centrali styliformi paucispi- rato praeditum, limbo peripherico incrassa- tum- Animal hucusque incognitum. Hab. Infundís coralligenis maris Sar- diniam meridionalem ambientis. * 5, Triq ir-Rghajja, Rabat RBT 2486, Malta ** Agisilaou 37-39, Tzitzifies/Kallithea, 17674 Athens, Greece 1 Iberus , 26 (1), 2008 Figure 1. Heliacus (Gyriscus) jeffreysianus (Tiberi, 1867), off Crete Island on Corallium rubrum (Linnaeus, 1758) in 120m. Size: 9x8 mm. Photo P. Ovalis. Figura 1. Heliacus (Gyriscus) jeffreysianus (Tiberi, 1867), frente a la isla de Creta sobre Corallium rubrum (Linnaeus, 1758) en 120m. Size: 9x8 mm. Foto P. Ovalis. Tiberi dedicated his species to the illustrious malacologist John Gwyn Jef- freys (1809-1885). Two of the specimens are presently in the Coen collection at The Hebrew University, (Department of Zoology) Jerusalem, Israel and the other specimen is in the Jeffreys collection at the Smithsonian Institution, Washington (USA) (Melone and Taviani 1984). New material: Two live specimens of Heliacus ( Gyriscus ) jeffreysianus were recently recovered from a living colony of Corallium rubrum (Linnaeus, 1758) from off the island of Crete at a depth of 120 m. Figure 1 shows one of the two similar specimens, measuring h= 9 mm x w= 8 mm which is about the same size as those found by Tiberi. The specimens were hand picked from a coral colony by a professional scuba diver from Crete using specialized diving equipment. It is not known whether the molluscs were actually feeding on the coral polyps. However, the species seems to have a cióse connection with C. rubrum. Several species from the family Architectonici- dae are known to be parasitic on coráis (Bieler 1993). Judging from the two presently known records, the species has always been found with these red coráis, and as it is well known, it was never found in any other Mediterranean habitat notwithstanding the large amount of research carried out in the last hundred years. Since its original description, the species had never been found again until recently, when living specimens of Heliacus jeffreysianus were collected in the Adriatic Sea (Stanic and Schiapar- elli, 2007). It is very curious, but not surprising, that both this and our dis- coveries were recorded at the same time and at distantly separated places. This is certainly due to the more recent interest being undertaken both by amateur and professional researchers in the depths of the red and white coral substratum of the Mediterranean Sea. The authors are also aware of other unrecorded species living in this part of the Mediterranean, and originally described from deep water in the Atlantic. Otherwise, Heliacus jeffreysianus has only been cited in the literature in a few species lists and catalogues pertaining to the Mediterranean malacofauna and it was even thought to be extinct. More- over, no fossil specimens of the species have ever come to light (Melone and Taviani 1984). Monteros ato (1880) had com- mented about the species of Tiberi "II G. jeffreysianus, é una delle nostre piú rare gemme, la cui scoperta, come di tante altre raritá coralligene, si deve al Dr. Tiberi. Soltanto tre esemplari ne sono conosciuti sinora. Beati possidentes! lo voglio ammettere che nella mente di un naturalista il Gyriscus abbia piú valore 2 Mifsud AND OvaliS: Re-discovery of living specimens of Heliacus (Gyriscus) jeffreysianus Figure 2. Map of the Mediterranean showing the three locadons cited for H. jeffreysianus. 1: Sardi- nia Island; 2: Adriatic Sea; 3: Crete Island. Figura 2. Mapa del Mediterráneo mostrando las tres localidades citadas para H. jeffreysianus. 1: isla de Cerdeña; 2: mar Adriático; 3: isla de Creta. del piú grosso diamente della corona d'Inghilterra, ma francamente il prezzo al quale lo vende il Dr. Tiberi, per una conchiglia cosí piccola, sará riguardato tanto dai naturalisti quanto dai mercanti come assai esagerato" In transía tion: "The G. jeffreysianus is one of our most rare gems, its discovery, like many other coralligeneous rarities, is owed to Dr. Tiberi. Only thee speci- mens are known up to now. Blessed are the possessors\ I must admit that in the minds of the naturalist the Gyriscus has more valué than the largest diamond from the British crown, but frankly the price which Dr. Tiberi has asked, for such a small shell is regarded by natu- ralists and also by merchants as being exaggerated." H. jeffreysianus is one of the most beautiful species from the family Archi- tectonicidae. Its sculpture of small- beaded spiral chords and its trochoid shape make it outstanding and very dis- tinguished among the other species of the family in the Mediterranean. In fact, Tiberi (1867) created a genus for it, Gyriscus, which is nowadays a subgenus of Heliacus. A very similar Pacific species is H. ( Gyriscus ) asteleformis (Powell, 1965) from New Zealand. Coen (1932) figured the operculum from one of the two specimens in his collection (also figured in Melone and Taviani, 1984). Melone and Taviani (1984) described and figured a syntype from the Coen collection. They also figured the radula (as redrawn after Merril, 1970). Other descriptions and diagnoses of the genus and the species can also be found in Coen (1932), Melone and Taviani (1984) and in Bieler (1993). CONCLUSIONS This very long elapsed period of o ver 140 years for the re-discovery for H. jeffreysianus is in all probability due to its rarity and the very particularly restricted and difficult to sample habitat. Red coral was traditionally har- vested by fishermen utilizing a particu- lar gear called the Cross, which consists of two large beams tied together in the form of a cross and armed with dan- gling ropes and pieces of netting to enable the coral to entangle during dredging. This heavy equipment, although it seemed adequate for the purpose, besides being very destructive to the substratum, constantly shakes the entangled coráis and the molluscs seem to always fall off before the gear is brought aboard the vessel. The modern method of manual harvesting through SCUBA is more selective, and therefore much less destructive. However, it is much more risky and dangerous. Curiously, this species had not been discovered on red coral before, although 3 Iberus , 26 (1), 2008 the method of manual harvesting of coral by specialized deep diving equipment (using a mixture of gases) has now been in practice for many years, especially by Spanish, Italian and Greek coral divers. More probably, it may take the keen eyes of a biologist, a naturalist or a shell col- lector to notice and pick out the small mollusc shells in situ. Therefore any spec- imens brought up by chance by the fish- ermen or the divers are probably either thrown back into the sea with the other rubbish as in the case of the fishermen or in the case of the diver, they fall off the coral unnoticed during the long decom- pression process. Moreover, the diver would be even more concerned for his "treasure" rather than a few "worthless" shells. Although Mediterranean Corallium rubrum is also found at great depths in the Mediterranean, the two existing records of H. jeffreysianus are both from shallow BIBLIOGRAPHY Bieler, R., 1993. Architectonicidae ofthe Indo-Pa- cific. Gustav Fischer Verlag, pp. 377. Stuttgart. Coen, G. S., 1932. Sul genere Gyriscus, Tiberi, 1867. Bollettino della Societd Veneziana di Scienze Naturali, 1 (1): 9-13. Melone, G. and Taviani, M., 1984. Revisione delle Architectonicidae del Mediterráneo. Atti del Simposio: Sistemática dei Prosobranchi del Mediterráneo, Bologna 24-26 Setiembre 1992, (B. Sabelli, ed.) Lavori S.I.M., 21: 149-192. Merrill, A. S., 1970. The family Architectonici- dae (Gastropoda: Mollusca) in the western and eastern Atlantic. Unpubl. Ph.D. Thesis, Univ. Delaware (Univ. Microfilms Int. Inc., Ann Arbor, Michigan, U.S.A., Nr. 7144444). water living colonies. Stanic and Schia- parelli (2007) did not mention the species on which they found their specimens. Finally, this record extends the species dis- tribution to quite a larger area of the Mediterranean (Fig. 2) and it is expected that more specimens which could con- tribute to the study of the biology of the species are likely to turn up. ACKNOWLEDGEMENTS We would like to thank Agios Nico- laos, a specialized deep sea coral diver for donating one of the two specimens which he collected to one of the authors (P. O.) for this study. Thanks are also due to Sophie Valtat (Belgium) for pro- viding important literature and to S. Gofas (Spain) for revising and enhanc- ing the manuscript. Monterosato, T. A., 1880, Nota sopra alcune conchiglie coralligene del Mediterráneo. Bol- lettino della Societd Malacologica Italiana, 6: 243-259. Stanic, R. and Schiaparelli, S., 2007. New finding of living specimens of the rare ar- chitectonicid Heliacus (Gyriscus) jeffreysianus (Tiberi, 1867). Bollettino Malacologico, 43 (9-12): 143-146. Tiberi, N., 1867. Diagnose du nouveau genre méditerranéen Gyriscus. Journal de Conchy- liology, 15: 303. Paris. 4 © Sociedad Española de Malacología Notes on West African Perrona (Gastropoda: Clavatulidae), with the description of a new species Notas sobre Perrona (Gastropoda: Clavatulidae) en Africa Occidental, con la descripción de una nueva especie Emilio ROLÁN*, Peter RYALL** and Juan HORRO*** Recibido el 31 -VI 1-2007. Aceptado el 19-X-2007 ABSTRACT The type species of the genus Perrona is poorly known and given an erroneous type local- ity. One of the purposes of this paper is to clarify its status and correct spelling as well as illustrating the species and defining its distribution area. A new species from Angola belonging to this genus is described. The shell, protoconch, operculum and radula are illustrated, and the reasons for assigning this new species to this genus are discussed. All known species of Perrona are figured. RESUMEN La especie tipo del género Perrona es poco conocida y la localidad tipo dada es errónea. El objetivo de este trabajo es clarificar su situación, su correcto nombre, además de repre- sentar la especie y clarificar su área de distribución. Se describe una nueva especie de este género de Angola. Se muestran concha, protoconcha, opérculo y rádula de la nueva especie y se discuten las razones para asignarla a este género y no a otros con similar morfología. Se ilustran todas las especies conocidas de este género. KEY WORDS: Clavatulidae, Perrona, West Africa, Angola, new species. PALABRAS CLAVE: Clavatulidae, Perrona, África occidental, Angola, nueva especie. INTRODUCTION Classical malacological works like Bruguiére (1792), Lamarck (1801, 1816 and 1822), Kiener (1840) and Reeve (1843, 1845) and publications dealing with Turridae (s.l.) such as Récluz (1851), Petit de la Saussaye (1851), von Maltzan (1883, 1884), Sykes (1905), Dautzenberg (1912), Strebel (1912, 1914) have described numerous West African species assigned to the family Clavatulidae Gray, 1853. Their generic placements and syn- onymies are still not clear. Some modem works like Knudsen (1952, 1956), Ardovini (2004), Boyer and Hernán- dez (2004), Nolf (2006), Nolf and Ver- straeten (2006) and Boyer and Ryall (2006) have described new species or revised others, placing most of them in the genus Clavatula. Among our material from Angola, dredged over many years by Xico Fer- * Museo de Historia Natural, Campus Universitario Sur, E 15782 Santiago de Compostela, Spain ** St. Ulrich 16, A-9161 Maria Rain, Austria *** Montero Ríos, 30-3° 36201 Vigo, Spain 5 Iberas, 26 (1), 2008 nandes, a new small unidentified species was found. Its description is one of the subjects of this work. The generic attribution to the genus Perrona (Schu- macher, 1817) requires a review of the type species because it is a rather poorly known species. This paper therefore also comments on the type species. The identity of Perrona perron Martini and Chemnitz (1788: 278, pl. 164, figs. 1573, 1574) illustrated a shell they named "Der Perron ". Later Gmelin (1791), referring to Chemnitz's figures, described it as Murex perron and indicated "in Océano australi" as its geographic origin. This type locality is obviously wrong. These figures, or the shell they represent, can be considered to be the lectotype. Its present where- abouts, as noted below, are unknown. When Lamarck (1816) described Pleurotoma spirata he also made refer- ence to the same illustrations, but sepa- rated his species from them by stating "La figure citée de Chemnitz offre sur la base du dernier tour ; des sillons dont notre coquille est absolutement dépourvue". Schumacher (1817) also referred to the same illustrations of Chemnitz when he described the genus Perrona and nomi- nated Perrona tritonium as the type of this genus. P. tritonium is therefore a júnior objective synonym of Perrona perron (Gmelin). Again no locality was given. The species was subsequently described for a third time by Reeve (1843) who referred once more to Chemnitz's illustrations. He gave it the ñame Pleuro- toma perronii, drawing his own figure from his own shell. He likewise did not State a locality or habitat. Reeve cited Lamarck 's spirata as a different species, writing "1 have a shell before me ofvery dif- ferent character (Pleurotoma Perronii, nobis), answering exactly to the figure in question". Reeve (1843) did not cite Gmelin and therefore appeared to con- sider his ñame as a valid new species. However it is clearly a júnior synonym of Gmelin's species and it is concluded that this was an omission in Reeve's work. Marrat (1877) was the first to give an indication of the true range of the species when he included it in a list of North-west African shells which he stated carne "from Madeira to the Gulf of Guinea". Tryon (1884), although stating his Opinión that this species was an inter- medíate form between lineata and spirata , mentioned it as Clavatula (sub- genus Perrona ) perron Chemnitz and provided a detailed description. Strebel (1912), in his revisión of genus Perrona, treated as clearly differ- ent species both spirata and perron. He was able to examine Chemnitz's speci- men from which the famous drawings were made and reillustrated it. He referred immature specimens of the species to Reeve's description, calling them perron var. reevei. He also correctly indicated part of the correct habitat range of the species, citing specimens he had examined from Pointe Noire, Congo to Ambriz, northem Angola. Knudsen (1952) employed the ñame "Clavatula perronii (Reeve)" noting that the "Atlantide expedition" could not obtain any additional material and that Chemnitz's type could no longer be traced at the ZMUC. Powell (1966), who referred cor- rectly to the type species of Perrona, did not figure Perrona perron but illustrated Perrona spirata (Lamarck, 1816). More recently, Cernohorsky (1974) figured a syntype from Chemnitz (in ZMUC), commenting that "the species most probably Uves in West Africa but this locality has been not substantiated by either literature records or authentic specimens. Another possibility is that Murex perron is conspecific with Pleurotoma spirata Lamarck, 1816, as suggested by Nordsieck (1968: 154) although the two species appear quite distinct". Finally, Bernard (1984, fig. 182) figured under the erroneous ñame "Clavatula kraepelini" three specimens of this species. He unknowingly extended Strebel's habitat range northwards, quoting the localities of both Cape Este- rias and Mayumba in Gabon, and indi- cating a depth of -20 to -50 meters. After examining Reeve's type speci- men, the available literature and some ó RoláN ET AL.: Notes on West African Perrona with the description of a new species shells in the collection of one of the authors (PR), it was decided to record some additional information about this poorly known group. Abbreviations: AMNH American Museum of Natural History, New York ANSP Academy of Natural Sciences, Philadelphia BMNH The Natural History Museum, London MCZ Museum of Comparative Zoology, Harvard university MHNS Museo de Historia Natural "Luis Iglesias" Universidad, Santiago de Compostela MNCN Museo Nacional de Ciencias Naturales, Madrid MNHN Muséum national d'Histoire naturelle, París ZMUC Zoologisk Museum, Copenhagen USNM National Museum of Natural History, Washington ZSM Zoological Staatmuenchen Museum, Muenchen CJH collection of Juan Horro, Vigo CPR collection of Peter Ryall, Maria Rain SYSTEM ATIC PART Family Clavatulidae Gray, 1853 Genus Perrona Schumacher, 1817 Type species: Perrona tritonium Schumacher, 1817 = Murex perron Gmelin, 1791 = Pleurotoma perronii Reeve, 1843. Perrona perron (Gmelin, 1791) (Figs. 1-13) Der Perron Martini and Chemnitz, 1788: 278, pl. 164. figs. 1573, 1574 (Fig. 1). Murex perron Gmelin, 1971: 3559. Perrona tritonium Schumacher, 1817: 218. Pleurotoma perronii Reeve, 1843: pl. 11, sp. 94 (Fig. 2). Type material: Figured syntype of Der Perron, whereabouts presently unknown; this specimen is reported as examined by Strebel (1912) but could not be traced by Knudsen (1952) who never- theless mentions another shell labelled in the handwriting of Chemnitz. Pleurotoma perronii: holo- type BMNH 1900.2.8.26. Other material examined: 8 shells (Figs. 4-10), Pointe Noire, Congo (CPR). Type locality: Erroneous for Der Perron (in Océano australi). Not mentioned in the original descrip- tion of any of the other taxa. Here designated at Pointe Noire, Congo. Description: Shell (Figs. 3-12) see the original descriptions as well as Tryon (1884) and Strebel (1912). The follow- ing data can be added: the protoconch (Fig. 13) consists of about 2 1 ¡i shiny, smooth whorls, honey brown in colour; the first whorl slightly darker and of 750 pm diameter. From the protoconch whorls a series of opisthocline axial ribs start to form, first as thin riblets extend- ing from the top to the bottom of the whorl, becoming stronger and shaped as an inverted C in the second teleo- conch whorl where their bases become more globose. Between them small striae can be seen. In the subsequent whorl they detach themselves entirely from the upper subsutural cord which also increases in prominence. In the fourth teleoconch whorl these vestiges of the axial ribs become closer and closer to the lower suture, declining in prominence until they entirely disap- pear. The following whorls are smooth with an irregular, raised subsutural collar and the whorls are traversed by s- shaped striae. At the same time as the axial riblets form in the third whorl. 7 Iberus , 26 (1), 2008 about 9 radial striae also start to emerge becoming slightly stronger in the fol- lowing whorls but then decreasing in intensity in the fourth or fifth whorls and thereafter are hardly noticeable. The shell is a light honey-brown throughout. On the fourth whorl the subsutural colar starts to become lighter, being entirely white in the fifth whorl. In subsequent whorls it can be interrup- ted by patches of light brown; in the final whorl the two major cords are white interspaced with small brown patches the same colour as the whorl itself and the cords on the columnella are also white. Size : the shell can reach up to 37.2 mm (Bernard, 1984). Remarks : The comparison of juvenile specimens of P. perron with species of the genus Clavatula (type species: Clavat- ula coronata Lamarck, 1801) show much similarity in the aspect of the opistho- cline axial ribs, which tend to disappear with maturity in the subsequent whorls. For this reason it is concluded that there is a stronger relationship between these two genera, than with other genera within this family. Whilst some authors synonymized P. perron with P spirata Lamarck, 1816 it must be noted that not only are they morphologically different, but the latter inhabits an area further to the south, being restricted to the Luanda area where it has been found at -5/50 meters. Distribution : This species inhabits the West African infralittoral coast from Cap Esterias (Gabon) to Ambriz (northern Angola). Special remarks: The type locality is always related with a holotype or a lec- totype. In the present case, the original figures are sufficient to determine the identity of the species and it is not nec- essary to designate a neotype; Pleuro- toma perronii was described lacking type locality as confirmed by the label of this holotype in BMNH. If the type locality of the first description is erroneous, and no other is mentioned in the description of the synonymized taxa, it is necessary to give more detailed information. Pointe Noire (Congo) is designated as the type locality being in the center of the known distribution area, as well as a locality mentioned by Strebel (1912) and the source of our specimens. Our material agrees very well with the figures and the known holotype. Concerning the correct spelling for the ñame of this species, it must be perron, which is the first ñame employed by Gmelin (1791) and which is appa- rently a ñame not dedicated to a person. Tryon (1884) stated that Chemnitz employed the common Dutch ñame "perron" (= flight of stairs, in reference to the profile) and that Reeve (1843), misinterpriting Chemnitz's ñame to be a personal ñame, Latinized it to read "perronii”. Perrona micro spec. nov. (Figs. 14-19, 21, 23, 24-28) Material examined: Holotype (Figs. 14, 15), in MNCN (15.05/47050). Paratypes in the following collections: MNHN (1, Fig. 16); BMNH (1, Fig. 17); AMNH (1); USNM (1); ZSM (1); ANSP (1); MCZ (1); CPR (11); CJH (6); MHNS (16, Fig. 18). Type locality: Palmeirinhas, south of Luanda, Angola, between 3 and 20 meters depth. Etymology: The specific ñame refers to the small size of the species. Description: Shell (Figs. 14-18) small for the genus, very solid, lanceolate. Protoconch (Figs. 21, 23, 24-26) with a little more than one whorl, of 530 pm in diameter and white in colour. The teleo- conch begins with three grooves that sepárate 4 cords, later increasing to four and then five spiral grooves; there are six in the subsequent whorl; the two cords immediately below the suture as well as the lowest cord are more promi- nent. This gives the middle part of the whorl a concave appearance whilst the upper and lower parts protrude before their separation at the suture. There is no axial sculpture except for growth 8 RoláN ET AL.: Notes on West African Perrona with the description of a new species Figures 1-13. Perrona perron. 1: figure from MARTINI AND CHEMNITZ (1788); 2: figure of the holotype of Pleurotoma perronii from Reeve (1843); 3: holotype (see below Figs. 11, 12) of Pleuro- toma perronii (BMNH) at the same comparative size; 4-6: shell (27.6 mm), Pointe Noire, Congo (CPR); 7: shell, 28.6 mm, Pointe Noire (CPR); 8, 9: juveniles, 18.6 and 14.2 mm, Pointe Noire (CPR); 10: shell, 27.2 mm, Pointe Noire (CPR); 11, 12: holotype of P. perronii , 25 mm (BMNH); 13: spire and protoconch, Pointe Noire (CPR). Figuras 1-13. Perrona perron. 1: figura de MARTINI Y CHEMNITZ ( 1 788); 2: figura del holotipo de Pleurotoma perronii, de Reeve ( 1 843); 3: holotype ( véase abajo Figs. 1 1, 12) de Pleurotoma perronii (BMNH) con el mismo tamaño relativo; 4-6: concha, 27,6 mm, Pointe Noire, Congo (CPR); 7: concha, 28,6 mm, Pointe Noire (CPR); 8, 9: juveniles, 18,6 y 14,2 mm, Pointe Noire (CPR); 10: concha, 27,2 mm, Pointe Noire (CPR); 11, 12: holotipo de P. perronii, 25 mm (BMNH); 13: spire and protoconch, Pointe Noire (CPR). 9 Iberus, 26 (1), 2008 Figures 14-19. Perrona micro spec. nov. 14, 15: holotype, 10.3 mm (MNCN); 16: paratype, 10.6 mm (MNHN); 17: paratype, 8.5 mm (BMNH); 18: paratype, 7.6 mm (scanning electrón micro- scope) (MHNS); 19: operculum (SEM). Figures 20-23. Comparison of the spire and protoconch of Clavatula quinteni (Figs. 20, 22) and Perrona micro (Figs. 21, 23). Figuras 14-19. Perrona micro spec. nov. 14, 15: holotipo, 10,3 mm (MNCN); 16: paratipo, 10,6 mm (MNHN); 17: paratipo, 8,5 mm (BMNH); 18: paratipo, 7,6 mm (microscopio electrónico de barrido) (MHNS); 19: opérenlo ( MEB). Figuras 20-23. Comparación de la espira y protoconcha de Clavatula quinteni (Figs. 20, 22) y Perrona micro (Figs. 21, 23). 10 ROLÁN ETAL.: Notes on West African Perrona with the description of a new species Figures 24-27. Perrona micro. 24: detail of the spire; 25, 26: detail of the protoconch; 27: radular tooth; 28: radula. Figures 29-33. Perrona obesa. 29: holotype, 34.0 mm (BMNH); 30: shell, 31.2 mm, 10 m, Namibe, Angola (CER); 31: shell, 35.7 mm, Namibe, Angola (CPR); 32, 33: shells, 40.2, 41.6 mm, Sacomar, Namibe, Angola (CPR). Figuras 24-27. Perrona micro. 24: detalle de la espira; 25, 26: detalle de la protoconcha; 27: dientes de la rádula; 28: rádula. Figuras 29-33. Perrona obesa. 29: holotipo, 34. 0 mm (BMNH); 30: concha, 31,2 mm, 10 m, Namibe, Angola (CER); 31: concha, 35,7 mm, Namibe, Angola (CPR); 32, 33: conchas, 40.2, 41.6 mm, Sacomar, Namibe, Angola (CPR). Iberus, 26 (1), 2008 lines which form an U-shape in the concave central portion; this is reversed on both the upper and lower protruding parts forming an overall S-shape on the whorl. There are about 6 whorls on the teleoconch, increasing gradually in width and height. The last whorl becomes elongated towards the base, the siphonal canal is widely open and almost similar in length to the lowest part of outer lip. The aperture is ovoid, with an U-shape sinus in the upper part, a sharp external lip, and a small callus on the columella. There are 20 spiral grooves from the periphery to the base. The colour is typically light brown or orangish, with white and dark blotches alternating on the subsutural cords, but in a few specimens the colour is darker and the white pattern is absent. Soft parts: Study of a retracted speci- men preserved in alcohol reveáis a light coloured animal; the operculum (Fig. 19) is ovoid with the nucleus centrally placed on the interior. The radula (Fig. 28) is formed by two rows of about 70 elongate and sharply pointed marginal teeth (Fig. 27) at the center of which is a very small central tooth reduced to a cusp. Dimensions : Flolotype is 10.3 mm. The largest paratype is 12 mm. Distribution: Only known from the type locality, where it is probably endemic. Discussion: There is no known species from West Africa with which it can be confused. Only juvenile forms of Clavatula quinteni Nolf and Verstraeten, 2006, which are much larger when adult, are similar in profile and coloura- tion; they both lack axial sculpture (see comparison in Figs. 20 and 21, 22 and 23) but C. quinteni has a much more elongated siphonal canal, a larger proto- conch (more than double the diameter) and lacks spiral grooves. There was some hesitation about the generic assignment of this new species. Comparison with morphologically similar species in some genera was made, in particular Microdrillia Casey, 1903; Suavodrillia Dalí, 1918; Parecuneus Laseron, 1954, and Maoritomella Powell, 1942, Tomopleura Casey, 1904, and Viridi- turris Powell, 1964. In all cases opercu- lum and radula do not agree. Perrona micro spec. nov., in spite of its small size, belongs to Clavatulidae. This is confirmed by the operculum with a medium-lateral nucleus and the wishbone-like marginal teeth with a vestigial rachidian one in the middle. Powell (1966) reviewed the Turri- dae (s. 1.) and defined the subfamily Clavatulinae H. and A. Adams, 1858 with four recent and two fossil genera; the recent genera being Clionella, Clavat- ula, Perrona (with subgenus Tomellana) and Pusionella. Kilburn (1985), who dealt only with South African species, noted 4 genera; Toxiclionella, Bentho- clionella , Clavatula and Clionella. Tucker (2004) considers 492 fossil and 97 recent species in this group. Later Bouchet and Rocroi (2005) elevated Clavatuli- dae to family status. The genus Clavatula Lamarck, 1801 is relatively abundant along the West Africa coast. Powell (1966) mentions 11 species. Most of them, including the type species ( Clavatula coronata Lamarck, 1801) are of large size. Other species recently illustrated and/or described, as detailed in our introduction, are also rel- atively large. Although Boyer and Ryall (2006) noted that Knudsen (1952, 1956) classified other genera of Clavatul- idae under the genus Clavatula, subse- quent authors seem to ignore these genera and their paramaters as estab- lished by Kilburn (1985). A revisión could conclude that some of them belong to other genera. The genus Clavatula is described by Powell (1966): "shell modera tely large to large sized, 15 - 55 mm, very solid, buccinoid, coarsely axially and spirally sculptured; with a tall, often coronated spire, and a trúncate body-whorl, termi- nated in a relatively short reflected and deeply notched anterior canal.... Oper- culum ovate-lunate with a medio-lateral nucleus. Radula consisting of a pair of stout narrowly pointed marginal teeth of modified wishbone type, the proxi- mal extremity sepárate and superim- posed upon the larger element. A small 12 RoláN ET AL. : Notes on West African Perrona with the description of a new species Figures 34-37. Perrona spirata. 34-36: shell covered by organic material, 31.5 mm, Luanda, Angola (CPR); 37: operculum. Figures 38-41. Perrona subspirata. 38: shell, 31.5 mm; 39-41: shell, 32.0 mm, Namibe, Angola (CPR); 42: operculum. Figuras 34-37. Perrona spirata. 34-36: concha recubierta por material orgánico, 31,5 mm, Luanda, Angola (CPR); 37: opérculo. Figuras 38-41. Perrona subspirata. 38: concha, 31,5 mm; 39-41: concha, 32,0 mm, Namibe, Angola (CPR); 42: opérculo. 13 Iberus, 26 (1), 2008 but we 11 formed unicuspid central tooth is present also." According to Kilburn (1985) the main features of Clavatula are: deep anal sinus, realtively long siphonal canal, distict parietal tubercle and proto- conch that is bluntly domed with about 2 1/2 whorls. The species P. micro spec. nov. has some shell characters of Clavatula but has no axial sculpture and is smaller than all of them. Clavatula cossignanii Ardovini, 2004, the smallest Clavatuli- dae known in West Africa, is different in most features including the fewer number of protoconch whorls and is endemic to Senegal. The holotype is 13.2 mm but it can reach a size of 15.9 mm (CPR). According to Powell (1966) the genus Clionella Gray, 1847 differs from Pusionella Lamarck, 1801 by its clavi- form shape with tall, flat-sided spire whorls; simple long flexuous axials ribs without either subsutural or peripheral processes and a truncated body-whorl. Kilburn (1985) defines Clionella with an anal sinus; a slight notch; siphonal canal short with no distinct parietal tubercle; protoconch somewhat conical, of about 2 whorls, the first one rounded and tilted. The genus is usually restricted to South Africa. P. micro spec. nov. lacks axial sculpture, its sinus is deep and does not have a truncated body-whorl: therefore it does not agree with Clionella. The new species was also ruled out of the genus Pusionella Gray 1847 because many specific characters are absent in P. micro ; the flat-sided whorls; the outer lip bordered by a broad, sub- sutural, very slight sinus and the smooth and polished surface. Finally, the genus Perrona Schu- macher, 1817 is described by Powell (1966) as: "shell moderately large, 25 - 40 mm, rather narrowly fusiform, with a tall spire of rapidly increasing whorls ACKNOWLEDGEMENTS The authors wish to thank the Euro- pean Synthesys Program which allowed one of the authors (ER) to examine the and a narrow body-whorl... surface smooth, or nearly so... with a conspicu- ous narrowly carinate subsutural collar.... operculum as in Clavatula... radula with a pair of elongate margináis and a narrow-based unicusp central tooth." The known West African species of Perrona, all figured in the plates in the present work, are the following: P. perron (Gmelin, 1791) (Figs. 1-13), P. spirata (Lamarck, 1816) (Figs. 34-37), both already mentioned, P. obesa (Reeve, 1843) (Figs. 29-33) and P subspi- rata (von Martens, 1903) (Figs. 38-41), both endemic to South Angola. Other authors have placed other species in this genus, for example Perrona nifat (Bruguiére, 1789) by Abbott and Dance (1986), but this generic place- ment is erroneous in our opinión. Powell (1966) also mentions another West African species belonging to this genus but placed in the subgenus Tomellana Wenz, 1843: Perrona ( Tomel - lana) lineata Lamarck, 1816; but this shell has a short, concave-outlined spire and a deep slit-like anal sinus and does not resemble our species. Strebel (1912) referred other species to Tomel- lana; they also do not correspond with the species currently reviewed. Perrona micro spec. nov. is more similar to Perrona perron than to any other type species and the new species is placed in this genus as being the closest. Also noticed is a similarity in the colouration of the subsutural cord as well as the S-shaped axial striae. We find this at present the better solution rather than to create a new genus. Whilst the similarity noted with Clavatula quinteni should not create con- fusión, a result of this paper must also be to assign this species to the genus Perrona although this has not been the reason for this work. type material of Pleurotoma perronii and other types; Jesús Méndez of the Centro de Apoyo Científico y Tecnológico a la 14 ROLÁN ET AL. : Notes on West African Perrona with the description of a new species Investigación (CACTI) of the University of Vigo for the SEM photographs; Jesús S. Troncoso for allowing us to use the Photographic apparatus of the Departa- BIBLIOGRAPHY Abbott, R. T. and Dance, S. P., 1982. Com- pendium ofSea Shells. E. P. Dutton, New York, ix + 411 pp. Ardovini, R., 2004. Clavatula cossignanii sp. n. (Gastropoda; Turridae) dal Senegal, West Africa. Malacologia, Mostra Mundiale, 43: 5-6. Bernard, P. A., 1984. Seashells ofGabon. Pierre A. Bernard, Libreville, 140 pp. Bouchet, P. and Rocroi, J.-P., 2005. 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Spécies général Muséum d'His- toire naturelle de París, collection Lamarck, celle du Prince Masséna et les découverts récente des voyageurs. París: Rousseau. 84 pp, 27 pls. Kilburn, R. N., 1985. Turridae (Mollusca: Gas- tropoda) of Southern Africa and Mozam- bique. Part 2. Subfamily Clavatulinae. Annals ofthe Natal Museum, 26 (2): 417-470. Knudsen, R. N., 1952. Marine Prosobranches of Tropical West Africa collected by the "Atlantidae" Expedition 1945-46. Part 1. Videnskabele Meddeleser fra Dansk naturhis- torisk Forening I Kjobenhavn, Bd. 114: 129- 185, pl. 13. Knudsen, R. N., 1956. Marine Prosobranchs of Tropical West Africa (Stenoglossa). Atlantide Report n° 4. Danish Science Press, Ltd. Co- penhaguen, 1956 pp. 93-105 Lamarck, J. B. P. A. de, 1801. 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Zootaxa, 682: 1-1295. ló Iberus, 26 (1): 17-28, 2008 © Sociedad Española de Malacología Pattern of reserve storage of the two mussel species Perna perna and Mytilus galloprovincialis living on Moroccan coasts: annual variation and effect of pollution Patrones de almacenamiento de reservas en dos especies de mejillón Perna perna y Mytilus galloprovincialis de las costas de Marruecos: variación anual y efecto de la contaminación Abdellatif MOUKRIM*1, Mohamed ID HALLA*, Abderazak KAAYA*, Abdellah BOUHAIMI*, Soumaya BENOMAR* et Michel MATHIEU** Recibido el 23-XI-2005. Aceptado el 30-X-2007 ABSTRACT The pattern of reserve storage and ¡ts ¡mportance ¡n the reproductive cycle of the two mus- sel species living in Moroccan coasts, Perna perna and Mytilus galloprovincialis, were studied comparatively, during two annual cycles. Study was conducted at a polluted and an unpolluted site in Agadir bay. Stereological analysis shows that P. perna presents only one type of storage cells (vesicular cells) storing glycogen. In M. galloprovincialis, two types of storage cells are present: vesicular cells and adipogranular cells (containing glycogen, lipids and proteins). In both species, seasonal variations of reserve tissue vol- ume are conversely proportional to those of the germinal tissues. Reserve tissue appeared in spring (May) and increased in summer. It decreased quickly from August onwards, and disappeared between January and April. In M. galloprovincialis, adipogranular cells dis- appeared before vesicular cells and reappeared first. In the polluted site, seasonal varia- tion of reserve and germinal tissues is comparable to that in unpolluted site but the máxi- mum valué of reserve tissue percentage is less ¡mportant. Moreover, during the year, this tissue did not disappear. Biochemical analysis shows that mean valúes of glycogen and proteins quantities are significantly higher in P. perna than in M. galloprovincialis. How- ever, total lipids quantity mean is higher in M. galloprovincialis. Seasonal variations of the three biochemical parameters present a similar profile in the two species. Compared to unpolluted site animáis, in polluted one, molluscs possess low glycogen and high lipids lev- éis. Seasonal variations of these parameters show a perturbed profile. RESUMEN Se ha estudiado y comparado el patrón de almacenamiento de reservas y su importancia en el ciclo reproductor en dos especies de mejillón de las costas de Marruecos, Perna perna y Mytilus galloprovincialis, durante dos ciclos anuales. Este estudio se llevó a cabo en un sitio contaminado y otro no contaminado de la bahía de Agadir. El análisis estere- ológico demostró que P. perna presenta un solo tipo de células de reserva (células vesicu- lares) almacenando glucógeno. En M. galloprovincialis, hay dos tipos de células de * Laboratory Aquatic Ecosystems: Marine and Continental Field, Biology Department, Sciences Faculty, Ibn Zohr University, BP 8106, 80000, Agadir, Morocco. ** Laboratory Marine Biology and Biotechnology, I. B. B. A., Caen University, France. 1 Corresponding author. 17 Iberus , 26 (1), 2008 reserva: células vesiculares y células adipogranulares (conteniendo glucógeno, lípidos y proteínas). En ambas especies, las variaciones estacionales del volumen de tejido de reserva son inversamente proporcionales a las del tejido germinal. El tejido de reservas apareció en la primavera (mayo) y se incrementó en verano. A partir de agosto, dismin- uyó rápidamente y desapareció entre enero y abril. En M. galloprovincialis, las células adipogranulares desaparecieron antes que las células vesiculares y son las primeras en aparecer en el siguiente ciclo. En el sitio contaminado, la variación estacional de tejidos de reserva y germinales es similar a la del sitio no contaminado, salvo que el porcentaje máximo de tejido de reserva es menor. Además, este tejido no desapareció a lo largo del año. Los análisis bioquímicos muestran que las cantidades promedias de glucógeno y de proteínas son significativamente más altas en P. perno que en M. galloprovincialis. Sin embargo, el promedio de lipidos totales fue mayor en M. galloprovincialis. Las varia- ciones estacionales de los tres parámetros presentan un perfil similar en las dos especies. Comparados con los del sitio no contaminado, los animales del sitio contaminado poseen niveles bajos en glucógeno y altos en lípidos. Las variaciones estacionales de estos parámetros muestran un perfil perturbado. KEY WORDS: Agadir, Morocco, mussels, Mytilus galloprovincialis, Perna perna, pollution effects, reproductive cycle, reserves strategy. PALABRAS CLAVE: Agadir, Marruecos, mejillones, Mytilus galloprovincialis, Perna perna , efectos de la conta- minación, ciclo reproductor, estrategias de reserva. INTRODUCTION Two sympatric mussel species are found along the Moroccan coast: the African mussel Perna perna and the Mediterranean mussel Mytilus gallo- provincialis. The first species is located in the north of its geographical distribu- tion; the second one in its Southern limit. In Agadir bay, these local popula- tions of mussels have important com- mercial valúes (Id Halla, 1997) and environmental interest (Najimi, Bouhaimi, Daubéze, Zekhnini, Pel- LERIN, NARBONNE, AND MOUKRIM, 1997; Kaaya, Najimi, Ribera, Narbonne, and Moukrim, 1999; Moukrim, Kaaya, Najimi, Roméo, Gnassia-Barelli, and Narbonne, 2000) as sentinel species. The few studies carried out on mussel biology in this area, concern only some aspects of the reproduction cycle (Shafee, 1989; Id Halla, Bouhaimi, Zekhnini, Narbonne, Mathieu, and Moukrim, 1997). They showed that P. perna and M. galloprovin- cialis present synchronous reproductive cycles, and breed throughout the year with a principal spawning period in spring. The only differences observed by Id Halla et al. (1997), regard the duration of the principal spawning period (more important for P. perna ) and the genital activity rhythm in summer (reduced in this species). In view of the importance of reserves in the reproductive cycle (Bayne, Bubel, Gabbott, Livingstone, Lowe and Moore, 1982; Lowe, Moore and Bayne, 1982; Pipe, 1987) and of the particularities of reserve strategy in Mytilidae (strategy essentially based on the glycogen, presence of specific storage cells and existence of a direct relationship between reserve metabo- lism and reproductive cycle), it was interesting to conduct a comparative study of P. perna and M. galloprovincialis in order to describe the reserves strat- egy for these two species living in the same environmental conditions. The two molluscs present an important dif- ference regarding the cells involved in the reserve storage. As indicated by Lunetta (1969) P. perna presents only one type of storage cells (vesicular cells or glycogen cells), storing especially the glycogen ; however, in M. galloprovin- 18 Moukrim ET AL.: Reserves strategy of mussels in Moroccan coasts cialis (Herlin-Houtteville, 1974; Dan- ton, Kiymoto, Komaru, Wada, Awaji and Mathieu, 1996), two types of storage cells are present, vesicular cells (storing glycogen) and adipogranular cells (containing glycogen, lipids and proteins). Furthermore, beyond the determina- tion of pattern of reserve storage and its importance in the reproductive cycle of the two mussels living in Moroccan coasts, this work studies the impact of pollution on this pattern. Therefore, a comparative study of two sites (unpol- luted and polluted) was conducted. MATERIAL AND METHODS Sampling This study was conducted during two annual reproductive cycles (October 1994 to August 1996). Two types of sites, representative of the Agadir marine bay were considered: i) a reference site (unpolluted). Cap Ghir, located 50 km north of Agadir City and far from any human activity, and ii) a polluted site (Anza) located 5 km north of Agadir and receiving the industrial and domestic untreated waste waters of Anza zone. Many studies, conducted during the same period of the present study, showed that this site is contami- nated by heavy metáis and PAHs either accumulated by mussels or in sediment (Id Halla, 1997; Najimi, 1997; Moukrim, Kaaya, Najimi, Roméo, Gnassia-Barelli and Narbonne 2000; Kaaya, 2002), with some perturbations in the physical and Chemical parameters of seawater (Id Halla, 1997). During this study, for each month, ninety individuáis of 35 mm for each species are collected at random in each of the two sites. 30 individuáis/ species/sites were subject to stereologi- cal analysis and 60 individuáis/ species/sites were dedicated to the bio- chemical analysis. Stereological analysis In order to follow the seasonal vari- ation of the glycogen according the reproduction cycle in Mytilus gallo- provincialis and Perna perna, an histo- chemical study (stereological analysis) was conducted in the mantle. Thirty adult mussels were collected and directly fixed in Gendre liquid for 3 days. Animáis were then removed from shells and a central portion of mantle tissue was correctly excised and embed- ded in paraffin. Sections of 6 pm were cut and stained by the periodic acid of Schiff (PAS). Stereological analysis was applied according the method of Weibel, Kistler, and Scherle (1966) which quantified the volume occupied by the reserve and germinal tissues. The cell types were determined accord- ing the definition of Lubet (1959): i) adipogranular cells (20-25 pm for length, 4.5-5 pm for length of nucleus, green coloration, contain lipids, pro- teins and glycogen), and ii) vesicular cells (50-80 pm for length, 3-5 pm for length of nucleus, pink coloration, contain only glycogen) Biochemical analysis Biochemical analysis (glycogen, lipids and proteins) was carried out on mussel mantle which in Mytilidae, con- stitutes an important organ in storage of reserves. Monthly, sixty adults were ran- domly sampled, quickly transferred to the laboratory in isotherm conditions and frozen at -30°C. Glycogen, total lipids and proteins were respectively measured according the methods of Duchateau and Florkin (1959), Folch, Lees and Sloane-Stanley (1957) and Lowry (1951). Rates of these compounds were expressed as mg/g fresh weight (mg/ g FW). Statistical analysis Stereological analysis was expressed as a percentage of the total volume of mantle tissue ; biochemical results as means ± standard deviations. The statis- tical significance of difference between samples was evaluated by the "t" test using the Statistica software (Release 4.5 A StatSoft Inc. Ed. 1993). A "P" valué of less than 0.05 was considered as sta- tistically significant. 19 Iberus , 26 (1), 2008 Figure 1. Cells implicated in the reserve storage in Perna perna (A) (only one type of cells: ve: vesicular cells) and Mytilus galloprovincialis (B) (two types of cells, ve: vesicular cells; adge: adi- pogranular cells). Scale bars 10 pm. Figura 1. Células implicadas en la acumulación de reservas en Perna perna (A) (solo un tipo de células: ve: células vesiculares) y Mytilus galloprovincialis (B) (dos tipos de células, ve: células vesiculares; adge: células adipogranulares). Escalas 10 pm. RESULTS Stereological analysis Figure 1A shows that the mantle of Perna perna presents only one type of storage cells: vesicular cells (VC). In Mytilus galloprovincialis (Fig. IB), two types of storage cells are presentí adi- pogranular cells (ADGC) and vesicular cells (VC). For P. perna, the respective volumes of germinal and reserve tissues are con- versely proportional (Fig. 2A). The sea- sonal profile is similar during the two studied annual eyeles. The volume of germinal tissue increases from October to February, when the máximum is reached (respectively 72% and 77% in the first and second year). From March onwards, the germinal tissue volume decreases to reach the minimum in August (respectively 8% and 6% in the first and second year). The reserve tissues appeared in spring (May) and increased until reaching maximal valúes in summer (August). They decreased quickly and disappeared between January and April. In the polluted site (Fig. 3A), the seasonal variation of reserve and germinal tissues is similar to that in the reference site but the máximum valué of reserve tissues per- centage is lower (57% and 62% respec- tively in the first and second year). Moreover, during the year, this tissue did not disappear. The minimum valué recorded was 7-10% in February. For Mytilus galloprovincialis (Fig. 2B), the volume of germinal tissue increases in autumn and in the beginning of winter to reach a máximum in February (75%). Thereafter, we observe a reduc- tion of the surface occupied by this tissue until October where the minimum (12%) is reached. During the second eyele a similar evolution is recorded. Otherwise, as with P. perna, the volume of germinal tissue is inversely proportional to reserve tissue. Nevertheless, in this case, the máximum is reached in October (64%), whereas the disappearance takes place from January to March for the two studied yearly eyeles. In the polluted site (Fig. 3B), the reserve tissue presents a seasonal evolu- tion similar to the reference site. Besides, this tissue never disappears as in P. perna. The occupied minimal volume is 5% reached in February. Moreover, for M. galloprovincialis, in the reference site, the adipogranular cells disappeared before the vesicular cells and then reappeared first. In the polluted site the two cellular categories persist 20 Moukrim ET AL. : Reserves strategy of mussels in Moroccan coasts A Reserve tissue ^ Germinal tissue O Others B 100% 90% OND JFMAM J JASOND JFMAM J JA ADGC VC Germinal tissue I | Others Figure 2. Seasonal variation of germinal and reserve tissues in Perna perna (A) and Mytilus gallo- provincialis (B) in the unpolluted site. (VC: vesicular cells storing glycogen; ADGC: adipogranular cells, containing glycogen, lipids and proteins). Figura 2. Variación estacional de los tejidos germinal y de reservas en Perna perna (A) y Mytilus gallo- provincialis (B) en el sitio no contaminado. (VC: células vesiculares almacenando glucógeno; ADGC: células adipogranulares, conteniendo glucógeno, lípidos y proteínas ). during all the year. However, an oscilla- tion can be observed in the volumes occupied by the two categories of cells. Biochemical analysis Glycogen: The mean amount of glycogen is significantly (F= 11.19 and P= 0.027) higher in Perna perna than in Mytilus galloprovincialis (respectively 59.6 ± 7.36 and 48.7 ± 4.78 mg/g FW). For the two species, the seasonal varia- tions are significant (F= 4.95 ; P= 10"6 for Perna perna and F= 2.65 ; P= 4.7xl0'6 for Mytilus galloprovincialis) and exhibit a similar profile (Fig. 4A). The glycogen increases in spring (from April), reaches a maximal valué in August (219.4 and 191.9 mg/g FW in Perna perna and Mytilus galloprovincialis respectively), then decreases during autumn and winter. The minimal valúes are noted in February (0.39 and 0.34 mg/g FW in Perna perna and Mytilus galloprovincialis respectively). 21 Iberus, 26 (1), 2008 ioo%- 90% - 80% - 70% ' 60% - 50% - 40% ' 30% ' 20% ' 10%' o%- i ¡i i I li I l i i ¡ I p II lli Wm i i 11. li ,1 H 1* " L i i ■I i» 1 OND JFMAM J JA SOND JFMAM J JA Reserve tissue ^ Germinal tissue Q Others 100% 90% 80% 70% 60% 50% 40% 30% 20% 10% 0% B Figure 3. Seasonal variation of germinal and reserve tissues in Perna perna (A) and Mytilus gallo- provincialis (B) in the polluted site. (VC: vesicular cells storing glycogen; ADGC: adipogranular cells, containing glycogen, lipids and proteins). Figura 3. Variación estacional de los tejidos germinal y de reservas en Perna perna (A) y Mytilus gallo- provincialis (B) en el sitio contaminado. (VC: células vesiculares almacenando glucógeno; ADG: células adip ogranulares, conteniendo glucógeno, lípidos y proteínas). ONDJFMAMJ JASONDJFMAMJ JA ¡H ADGC H VC ^ Germinal tissue Q Others In the polluted site (Fig. 4B), the dif- ference between the mean amount of glycogen in the two species (39.75 and 23.44 mg/g FW in P. perna and Mytilus galloprovincialis respectively) is not sig- nificant (F= 6.257 and P= 0.130). Other- wise, their seasonal variation are signifi- cantly different (F= 4.03 and P= 10'6 in P. perna and F= 9.78 and P= 10~7 in M. galloprovincialis). The seasonal profile is completely different for the two mol- luscs: For P. perna , it presents three peaks, the first in June (96.05 mg/ g FW), the second in November (118.87 mg/g FW) and the third in May of second year (159.02 mg/g FW). For M. galloprovin- cialis, only two peaks are recovered 65.03 and 153.64 mg/g FW respectively in May and November of the first year. The comparison of valúes for refer- ence and polluted site animáis shows a significant difference. The valúes in the polluted site are less important (39.8 ± 13.36 instead of 59.6 ± 7.36 mg/g FW in 22 Moukrim ET AL Reserves strategy of mussels in Moroccan coasts P perna ▲ M. galloprovincialis Month Figure 4. Seasonal variation of glycogen in the mantle of Perna perna and Mytilus galloprovincialis in the reference site (A) and the polluted site (B) Figura 4. Variación estacional del glucógeno el el manto de Perna perna y Mytilus galloprovincialis en el sitio de referencia (A) y en el sitio contaminado (B) the reference site, for Perna perna). More- over, the seasonal profile of glycogen contení (Fig. 4B) presents some pertur- bations compared to the reference site. Lipids : A significant difference was noted between the mean amounts of total lipids in the two mussels species (92.86 and 125.04 mg/g FW in P. perna and Mytilus galloprovincialis respectively). However, the difference of their seasonal variation are significant with F= 4.42 and P= 10'6 in P. perna and F= 3.6 and P= 4.5xl0'5 in M. galloprovincialis. In the first year, the seasonal profile (Fig. 5A) is similar for the two molluscs. The lipid contení increased in autumn, with a maximal valué in December (respectively 354.3 and 316.9 mg/ g FW in Perna perna and Mytilus gal- loprovincialis), and decreased in winter and spring. However, in the second year, a slight increase was noted in summer (June - July) only in P. perna. In the polluted site, the two species present a significant difference between total lipids mean valúes (respectively 231.4 ± 21.47 and 135.5 ± 16.93 mg/g FW in Perna perna and Mytilus gallo- provincialis). Besides, the difference in their seasonal variation (Fig. 5B) is sig- nificant with F= 2.59 and P= 0.002 in P. perna and F= 3.60 and P= 4.5xl0"4 in M. galloprovincialis. Ótherwise, the valúes are generally higher for the polluted site than the ref- erence site for the two species. More- over, compared to the unpolluted site, the seasonal variation (Fig. 5B) shows a perturbed profile with several peaks during the annual cycle and a máximum valúes is reached rather in August (777.03 mg/ g FW in P. perna and 659.95 mg/g FW in M. galloprovincialis). Proteins : The mean contení of protein is significantly higher in Perna perna (28.9 23 Iberus, 26 (1), 2008 P. perna a M. galloprovincialis Month Figure 5. Seasonal variation of lipids in the mantle of Perna perna and Mytilus galloprovincialis in the reference site (A) and the polluted site (B). Figura 5. Variación estacional de los lípidos en el manto de Perna perna y Mytilus galloprovincialis en el sitio de referencia (A) y en el sitio contaminado (B). ± 3.51 mg/g FW) compared to Mytilus galloprovincialis (20.2 ± 3.17 mg/g FW). However, the seasonal profile (Fig. 6A) is similar for the two species. The maximal valúes were generally recorded in summer and autumn in the first year and in winter and beginning spring in the second year, whereas the minimal valúes were reached at the end of autumn and the beginning of winter. In the polluted site (Fig. 6B), the mean amount of protein is higher in P. perna (30.11 ± 7.742 mg/g FW) than in M. galloprovincialis (19.74 mg/g FW). However, the mean valúes and the sea- sonal profile do not show any signifi- cant difference compared with those observed from the reference site (F= 1.08 and P= 0.30 for Perna perna ; F= 0.31 and P= 0.57 for Mytilus galloprovincialis ). For the two species, the máximum valúes were recorded in summer and autumn. DISCUSSION The comparative study of the reserve pattern of the two mussels species living in the Moroccan coasts, Perna perna and Mytilus galloprovincialis, sampled in the same site (Cap Ghir, Agadir bay), shows that, in spite of the difference in their reserve tissues (only one cell type, vesicular cells, in Perna perna ; two cell types, adipogranular cells and vesicular cells, in Mytilus gallo- provincialis), the two molluscs have a similar fluctuation. This strategy presents cycles (of re- serve compounds and cells involved in the storage) which, compared to the cy- cle of reproduction (determined in these same animáis by Id Halla et al., 1997) are inversely proportional to it. The ac- cumulation of reserves in these cells is related to the period of reduced sexual 24 Moukrim ET AL. : Reserves strategy of mussels in Moroccan coasts P. perna a M. galloprovincialis Figure 6. Seasonal variation of proteins in the mantle of Perna perna and Mytilus galloprovincialis in the reference site (A) and the polluted site (B). Figura 6. Variación estacional de las proteínas el manto de Perna perna y Mytilus galloprovincialis en el sitio de referencia (A) y en el sitio contaminado (B). activity (in summer). The low levels of reserve are obtained in autumn and par- ticularly in winter (gametogenesis and spawning periods). Similar results have been reported in Perna perna from Brazilian coasts (Lunetta, 1969) and in Mytilus galloprovincialis from the French coasts (Herlin-Houtteville, 1974; Danton, Kiymoto, Komaru, Wada, Awaji and Mathieu, 1996). Also, in the Ria de Vigo (Spain), the profile of the variations of the gonadic Índex and so- matic Índex in cultured mussels show clearly their inverse and gradual fluctu- ations (Cáceres-Martínez and Figueras, 1998). This association be- tween gonad and storage tissue cycles in mussels is well known. Decline of the ADG cells occurs during gametogenesis by a lysosomal autophagic mechanism (Bayne et al., 1982; Lowe, Moore and Bayne; 1982; Pipe, 1987). The energy used in gonad restoration following spawning during spring and summer probably derived directly from feeding since ADG cells disappeared from the mantle in early spring (Villalba, 1995). Furthermore, the reserve accumula- tion in the two mollusc species is related to the proliferation of the phytoplanc- tonic biomass linked to upwelling cur- rents which take place, between Febru- ary and August, in the Agadir bay and more precisely in Cap Ghir (Belveze, 1984 ; Agoumi and Orbi, 1992). Cáceres-Martínez and Figueras (1998) reported that this increase in food availability for mussels in the area favours the accumulation of reserves during this period. These results also confirm the cióse dependence, described by Mathieu (1987), between the reserve storage and environmental conditions in marine ecosystems. As suggested by 25 Iberus, 26 (1), 2008 this author, the disappearance of reserve tissue in mussels seems to be under endocrine control via cerebroid ganglia which provoke disappearance of adi- pogranular cells and vesicular cells in the mantle after liberation of their reserves which are indispensable for gametogenesis and spawning process. The synchronic character of this disap- pearance, in Perna yerna and Mytilus gal- loyrovincialis, could be explained by the existence of the same mechanism for the control of reserves in both species. The comparison of the reserve com- pounds in the two molluscs shows that the glycogen, total lipids and proteins contents are higher in Perna yerna (in spite of the presence of only one type of reserve storage cells, vesicular cells) compared to Mytilus galloyrovincialis. These results could explain the differ- ences observed by Id Halla et al. (1997) between the reproductive cycles of these bivalves. According to these authors, in Perna yerna, compared to Mytilus galloyrovincialis, the main- spawning period in spring is longer and the sexual activity in summer is reduced. Otherwise, the important reserve levels in this species could be attributed to the spatial distribution of each species: Perna yerna lives essen- tially at the infra-littoral level and is more immersed than Mytilus galloyrovin- cialis (Id Halla et al., 1997) and, conse- quently, has access to more nutrients, thus allowing the synthesis and storage of more reserves. Seed (1976) showed that the gonadal development was faster in mussels from the low intertidal zone, than in those from the upper zone and related these results to food avail- ability. Other studies associated local variations in gonadal cycle with envi- ronmental conditions (Ferrán, 1991; Villalba, 1995). According to Cáceres- Martínez and Figueras (1998), there is no influence of locality and depth in the gonadal development of cultured mussels. The seasonal profile of the biochemi- cal reserves studied shows that the first peak of glycogen and lipids, recorded in summer, is related to the occurrence of upwelling currents which provide food availability, ensuring an abundant planktonic food supply for mussels. According to Cáceres-Martínez and Figueras (1998), massive spawns occur in spring coinciding with an increase in temperature and chlorophyll-a concen- traron in the area providing favorable conditions for larval growth. In winter, the second peak of lipids coincides with the maturity period of gametes. This peak takes place just after the disap- pearance of the glycogen peak. It is probably a result of the glycogen trans- formaron. The metabolic conversión of glycogen to lipids has been reported by Zaba and Davies (1980), using 14C- glucose. According to Gosling (1992), the mantle is considered as the organ of many and extensive metabolic transfor- mations during the sexual cycle. The reserves, particularly the glycogen, accumulated during summer, are used in autumn and winter for the gametoge- nesis. Similar results were reported by Shafee (1989) in Perna yicta of Temara (North of Morocco). In the polluted site (Anza), many perturbations in the metabolism of reserves were noted comparatively to the reference site (Cap Ghir). The reserve tissues, which disappeared in winter and at the beginning of spring, in mussels of Cap Ghir, persist throughout the sexual cycle in Anza mussels. This could probably be a result of the pollu- tion effect on the cerebroid ganglions neurosecretions which are, according to Lubet, Herlin, Mathieu, and Collin (1976) and Mathieu (1987) involved in the control of reserve cells. The analysis of the seasonal profile of the reserve levels in mussels sampled in the polluted site shows some pertur- bations as compared to the reference site. For example, the glycogen content is low in summer (August), in spite of the availability of phytoplanctonic biomass in this period. This is probably linked to the stress caused by pollution of industrial and domestic waste waters discharged directly in this site, without any treatment. This fact was indicated by Deslous-Paoli, Wolowicz, and 26 Moukrim ET AL.: Reserves strategy of mussels in Moroccan coasts Boromthanarat (1991) who reported that, in Mytilus edulis, reserves could be used both in reproductive process and to overeóme the hard environmental conditions. According to Thompson (1972) the reserves are used in order to reach the basal level of energy necessary for stressed animáis. Contrary to the glycogen, the total lipids are more important in Perna yerna and Mytilus galloprovincialis living in Anza (polluted site). This could be explained by an eventual direct assimi- lation of lipids from the organic matter of waste waters and/or a change of the reserve storage process. According to Gosling (1992) the lipid storage process in molluscs is considerably linked to the environmental conditions, particularly to the presence of pollutants. As a general conclusión, in the two species of mussel P. yerna (with only vesicular cells) and M. galloprovincialis (with adipogranular cells and vesicular cells), the respective germinal and reserve tissues clearly show their inverse and gradual profile. Their sea- sonal fluctuations are similar in the two molluscs. In the polluted site, many per- turbations of the reserve metabolism were noted comparatively to the refer- ence site. Then, contrary to animáis of this latter site, which presented a glyco- BIBLIOGRAPHY Agoumi, A. and Orbi, A., 1992. Evolution météorologique et upwelling le long de la cote atlantique marocaine. Hydroécologie Appliquée, 2 (4): 149-158. Bayne, B. L., Bubel, A., Gabbott, P. A., Liv- INGSTONE, D. R., LOWE, D. M. AND MOORE, M. N., 1982. 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Marine Biology Eetters, 1: 235-243. 28 Iberus, 26 (1): 29-42, 2008 © Sociedad Española de Malacología Reproduction of the cockle Cerastoderma edule (Linné, 1758) in the estuary of Oued Souss (southwestern Morocco) Reproducción del berberecho Cerastoderma edule (Linné, 1758) en el estuario del Oued Souss (suroeste de Marruecos) Hafida BERGAYOU**, Abdellatif MOUKRIM*1, Michel MATHIEU** and Jean-Pierre GIMAZANE** Recibido el 14-11-2006. Aceptado el 3 l-X-2007 ABSTRACT Field and laboratory investigations were carried out from 2001 to 2003 on a population of Cerastoderma edule living ¡n the estuary of Oued Souss, ¡n order to determine the impact of domestic wastewater discharges on the biology of this species. During waste- water discharges, several episodes of partial spawning occurred from November to March, followed by another in April-May. The main spawning occurred nevertheless in August. The gonadic Índex was always higher than 1, so that the period of sexual rest was short. The period of recruitment (from June to October) was mainly correlated with spawning peaks. The renewal of the population was ensured by the recruitment of June- July, whereas young cockles recruited from August to October only represented 9.2% of the population at the end of autumn. Most 1-year-old cockles disappeared during the sum- mer of the second year following their recruitment. After wastewater pollution stopped, an important recruitment of young cockles occurred. The spatial distribution of this species progressed upstream and the biomass of cockles had strongly increased. Because of its reproduction which takes place throughout the year, C. edule , as an opportunistic species, succeeded in colonizing this site. RESUMEN Se realizaron, entre 2001 y 2003, estudios de campo y de laboratorio sobre una pobla- ción de Cerastoderma edule del estuario del Oued Souss, con el objetivo de determinar el impacto de los vertidos de aguas residuales sobre la biología de esta especie. Coinci- diendo con los vertidos, se produjeron varios episodios de puesta parcial entre Noviem- bre y Marzo, seguidos de otro en Abril-Mayo. La puesta principal, sin embargo, se pro- dujo en Agosto. El índice gonádico fue siempre superior a 1 , indicando que el periodo de reposo sexual fue corto. El periodo de reclutamiento (de Junio a Octubre) estuvo esencial- mente relacionado con picos de puesta. La reposición de la población fue asegurada por el reclutamiento de Junio-Julio, mientras berberechos jóvenes reclutados entre Agosto y Noviembre representaban tan solo el 9,2% de la población a finales de otoño. La mayor parte de los berberechos con un año de edad desaparecieron durante el verano del segundo año después de su reclutamiento. Al finalizar los vertidos se produjo un impor- tante reclutamiento de berberechos jóvenes. La especie se extendió río arriba y la bio- * Laboratory Aquatic Ecosystems : Marine and Continental Field, Biology Department, Sciences Faculty, Ibn Zohr University, BP 8106, 80000, Agadir, Morocco. ** Laboratory of Marine Biology and Biotechnology, University of Caen, F- 14000, Caen, France. 1 Corresponding author 29 Iberus , 26 (1), 2008 masa de berberechos aumento fuertemente. Por su reproducción que se extiende sobre todo el año, C. edule, como especie oportunista, fue exitosa en la colonización de este lugar. KEY WORDS: Cerastoderma edule, estuary, Oued Souss river, pollütion, recruitment, reproductive cycle, waste- water discharges. PALABRAS CLAVE: Cerastoderma edule, estuario, Oued Souss, contaminación, reclutamiento, ciclo reproductor, INTRODUCTION In marine environment, many biotic and abiotic factors may influence the life cycle of bivalves. If temperature, salin- ity, food supply, and tidal exposure are the most important causes known to modulate development in mytilids (Seed, 1975), other elements, such as intraspecific competition or environ- mental contaminants, can result in great variations in growth rate of bivalves (Seed and Suchanek, 1992). These factors also have an effect on the repro- duction, settlement, recruitment, and production of these molluscs so that their variations directly affect the devel- opment of beds for each species of bivalve (Widdows and Donkin, 1992). Contrary to numerous ecological reports on the effects of contaminants on different marine bivalves, the recovery of mollusc populations after the disap- pearance of pollütion has been less investigated. The implantation of a wastewater pu- rification plant since November 2002 along the estuary of Oued Souss, at 4 km from the mouth of the estuary, had re- sulted in the fact that the fresh water still running in the river and the decanted wastewater were no longer discharged in the estuary but diverted to another Coastal site: M'Zar, located at 3 km south. For this reason, the estuary, upstream to the purification plant, was only swept by sea tide at the present time. As there existed a population of Cerastoderma edule living in the estuary of the Oued Souss river, it was interest- ing to determine the physiological State of these cockles during the pollütion period by wastewater and the changes which have occurred after this contami- nation stopped. In view of these objec- tives, the following two questions aróse: Had the discharge of wastewater before November 2002 caused repercussions on the reproductive cycle of C. edule and on the dynamics of this population? What consequences did the presence of sea water in the estuary after November 2002 have on the spatial distribution and biomass of C. edule? To answer the first question, monthly investigations from January 2001 to December 2002 were carried out in the habitat of C. edule i) to follow the different stages of sexual maturity in males and females, and ii) to specify the development of the different generations and cohorts of cockles. To tackle the second question, two surveys in 2002 and 2003 during summer months were performed in the estuary of Oued Souss river. This study complements other reports made by our research team in the same ecosystem (Mouneyrac, Pel- LERIN, MOUKRIM, AlT ALLA, DUROU AND Viault, 2005; Ait Allá, Mouneyrac, Durou, Moukrim and Pellerin, 2005; Ait Allá, Gillet, Deutsch, Moukrim and Bergayou, 2005; Bergayou and Moukrim, 2003 and Gillet, Gorman, Tallec, Moukrim, Mouloud, Anajjar, Ait Allá, Bergayou and Kaaya, 2003). MATERIALS AND METHODS The estuary of Oued Souss is located on the Atlantic coast, in southwestern Morocco and is subjected to an arid climate. The mouth is swept by an intense marine hydrodynamism, responsible of the presence of great 30 Bergayou ET AL.: Reproduction of Cerastoderma edule in Southwestern Morocco Figure 1 . Location of sampling sites along the estuary of Oued Souss in the Bay of Agadir, south- western Morocco. Figura 1. Situación de las localidades de maestreo a lo largo del estuario del Oued Souss, en la bahía de Agadir, suroeste de Marruecos. sandbanks in the estuary, with currents linked to high or low tides, and with high salinity (from 31.2 to 34.2 %o). Reproductive cycle The study of reproductive cycle and population dynamics for C. edule were carried out at station 2 (30° 21.97'N north, 9o 35.98'W west), (Fig. 1). Selected because of its high biomass of the cockle population, this station was character- ized by fine silty sand 2-5% of organic matter in the substratum and 340-440 mg/1 of suspended matter in the water. Two methods for studying the repro- ductive cycle of C. edule were used. The first was a classical histological study of gonads and was performed from January 2001 to March 2002. The second was a complementary study of condi- tion Índex and microscopic examination of gonad smears (Guillou, Bachelet, and Glémarec, 1991) and was per- formed from August 2001 to August 2002. For the first and second study. respectively, 30 and 100 cockles (length, 20 to 30 mm) were collected by hand at low tide, at monthly intervals (respec- tively, a total of about 450 and 1300 accumulated cockles was examined). For the first study (histological study of gonads), the shells of C. edule were opened and soft masses were prefixed in the Gendre's fixative for 24 h. In the laboratory, the shell of each bivalve was removed and small pieces of soft masses were post-fixed in a new solution of Gendre's fixative for 48 h before being dehydrated through a graded series of ethanol and butanol, and finally embed- ded in cytoparaffin (56°-58°C). Serial sections (thickness, 5 pm) were made before being stained with Gabe's trichrome, hemalun-eosin, or Mann- Dominici's method (Gabe, 1968). The maturity of gonads was determined using the scale proposed by Lubet (1959) for Mytilus edulis (Table I). The gonadic Índex (Seed, 1975) indi- cates the State of gonad maturity for 31 Iberus, 26 (1), 2008 Table I. Terminology used by LUBET (1959) and LUCAS (1965) to study morphologically and his- tologically the gonadic developmental stages in bivalves. Tabla I. Terminología empleada por Lubet (1959) y Lucas (1965) para el estudio morfológico y his- tológico de estadios de desarrollo gonadal en bivalvos. Lubft's scale (1959) Lucas's scale (1965) Staqe 0: sexual rest. Staae 1: earlv qametoaenesis with numerous qoniae. Staqe III D: spent, completelv emptv lumina. Staqe A: sex undetectable. Ihis stage can correspond to sexual rest, previtellogenesis, or gamete resorption. Staqe II: activelv developinq qonads but mature qametes were not observed. Staqe B: sex detectable with difficully to the naked eye. Staqe IIIA: near ripe follides with mature qametes. Staqe IIIB: spawninq, follicles distended. Staqe C: identifiable gonad. Ihe foot is salmon-stained in males and pearly white in females. Gonad maturation and spawning occurred during this stage. Staqe IIIC: partial spawninq, partiallv emptv lumina. Staqe D: gonadic reconstitution with co-existence of empty follicles and of tubules showing numerous goniae. each population and is evaluated from histological slides. It was determined by giving a number to each of Lubet's gametogenic stages: stage 0 (number 1), stages I and II (2), stage IIIA (3), stages IIIB and IIIC (2), and stage IIID (1). For each sample of cockles, the number of gonads showing a gametogenic stage is multiplied by the corresponding number; the figures obtained were then added and the sum was then divided by the total number of cockles studied. This gonad Índex varied from 1 (all gonads were spent, with completely empty lumina) to 3 (all gonads were ripe). The stereological analysis was made on the whole bivalves used for the histo- logical examinations of gonads. For each C. edule, three histological slides were randomly chosen through the antero- posterior axis. The different cell cate- gories present in gonadic follicles were counted on five ocular fields (magnifica- tion: x 100 for females, and x 400 for males) randomly selected in the visceral mass. In females, four categories: oogo- niae, vitellogenic oocytes, ripe oocytes, and atresic oocytes, were considered. In males, the cells were classified into the following three categories: protogoniae and spermatogoniae, primary and sec- ondary spermatocytes, and, lastly, sper- matids and spermatozoa. The mean per- centage of each cell category was calcu- lated in relation to the total number of cells counted. Mean valúes and corre- sponding S.D. were established for each cell category. For the second study, gonad smears and the determination of sex ratios were made by removing each bivalve from its shell and by rubbing soft masses against a histological slide. The microscopic examination was made using the scale of Lucas (1965) adapted for C. edule by Fernandez-Castro, Guillou, Le Pennec, and CARDENAS López (1989) (see table I). The sex ratio (number of females in relation with the total number of bivalvs with a recognizable sex) was performed when this identifi- cation was easy. A x2 test was used to determine levels of statistical signifi- cance. The condition Índex represents the variations of dry weight for a standard bivalve and aims to elimínate the effect of mollusc growth while revealing an accumulation or a loss of organic matter, associated with reproduction. A single 32 Bergayou ET AL.: Reproduction of Cerastoderma edule in Southwestern Morocco size class (20-30 mm) was considered and 30 bivalves were randomly chosen to determine this condition Índex. The índex selected was that proposed by Lucas and Beninger (1985): IC = [(dry weight of soft masses) / (dry weight of valves) x 1000]. The dry weight was obtained using a dehydration of soft masses (or shell) in a desiccator (60°C, 24 h). Population dynamics To study the dynamics of the popu- lation and to specify the period of juve- nile recruitment, samplings of C. edule were performed from January to December 2002. This study was also carried out in the station 2. Monthly, sixteen sediment samples (surface, 0.0625 m2, height, 20 cm) were collected at low tide according to the method of quadrats used by Elliott and Decamps (1973, in Bayed, 1982). These samples were sieved (square meshes, 1 mm) to recover and count cockles. The antero- posterior length of each cockle was mea- sured using a calliper rule (precisión, 0.1 mm). For mollusc sizes less than 5 mm, a stereomicroscope equipped with a micrometric ocular was used. The indi- vidual valúes recorded for the length of bivalves and sampling dates were com- pared using the FISAT software (Gayanilo, Sparke and Pauly, 1996) to draw size histograms in relation to the frequency of animáis and to make a modal analysis based on the algorithm according to the method by Battacharya (1967). This comparison, with the use of the x2 test (at P<0.05) allows to distin- guish the different size classes in each monthly sample, to determine mollusc densities, and to specify recruitment periods. Spatial distribution, density and biomass To determine the spatial distribu- tion, density, and biomass (ash-free dry weight, AFDW) of cockles, two surveys were carried out in 2002 and 2003 (during summer period). The replication of this study during these two periods (in 2002, when the estuary received the wastewater, and in 2003, when the dis- charge stopped) gives insights on the wastewater pollution impact and the reestablishment of the ecosystem when this pollution stopped. These investiga- tions were made in a total of 8 stations (27 sampling sites) located at 400-500 m intervals from the estuary mouth up to the wastewater outlet (Fig. 1). In each site, four to six sediment samples (surface, 0.0625 m2, height, 20 cm) were collected at low tide according to the method of quadrats used by Elliott and Decamps (1973, in Bayed, 1982). These samples were sieved (square meshes, 1 mm) to recupérate and count cockles, as above. Ash-free dry weight (AFDW) was determined by dipping molluscs in a solution of 10% HC1 until the shell was completely dissolved. Then, the soft parts were dried for 48 h at 80°C, weighed, placed at 600°C for 2 h and weighed again. The weight loss at 600°C was considered to represent the AFDW of individuáis and was expressed in g/m2 (Bachelet, Bouchet and Lissalde, 1980-1981). RESULTS Effects of pollution on the biology of the species Reproductive cycle: The study of gonad smears from August 2001 to August 2002 had allowed to recognize sex in 1138 bivalves (87.5%). Within this group, the sex ratio of C. edule was 51.1% ± 2.6%, and was well-balanced at 1:1 (x2 = 2.5, P < 0.05). No hermaphro- dite individuáis were found in this pop- ulation. In September, the A stage (undetectable sex) was observed in 75% of cockles and might correspond to a previtellogenic phase rather than a sexual rest, as the percentage of C. edule showing the C stage (gonad maturation) increased in October to reach about 50% in November. The frequency of C stage was above 50% between November and April (Fig. 2A), and subsequently decreased in May (from 75 to 20%), thus corresponding to a slight decrease of the 33 Iberus, 26 (1), 2008 Figure 2. Distribution of sexual-development stages (A) according to the scale of LUCAS (1965) and annual cycle of condition indexes (B) in the populations of C. edule studied from August 2001 to August 2002. The cumulated frequencies of the different sexual-development stages corresponded to the total of bivalves studied (100%). For details of each stage, see Materials and Methods (Table I). Figura 2. Distribución de los estadios de desarrollo sexual (A) según la escala de LUCAS (1965) y ciclo anual de índices de condición (B) en las poblaciones de C. edule estudiadas entre agosto 2001 y agosto 2002. Las frecuencias acumuladas de distintos estadios de desarrollo sexual corresponden al total de bivalvos estudiados ( 1 00%). Véase Material y Métodos (Tabla L) para los detalles de cada estadio. condition Índex (Fig. 2B). An episode of pardal spawning had thus occurred during this last period. In May, the fre- quency of the D stage (80%) proved that gametogenesis had resumed. In June and July, the C stage was the most fre- quent (65 to 75%) and was followed in August by a more marked decrease of the condition Índex (from 80 to 50 %o) thus indicating an important release of gametes. It may be concluded that two periods of gamete maturation (C stage), the first ranging from November to April and the other occurring in June- July, were differentiated. The histological study was carried out over a longer period of time (January 2001- March 2002) than the analysis of gonad smears. Mature gonads (IIIA stage) were often observed on more than 50% of cockles studied from January to April (Fig. 3). Episodes of pardal spawn- ing occurred from January to March; fol- lowed by another spawning in April and May (in May, the IIIB stage was noted in 60% of males, Fig. 3A). The reconstitu- tion of the gonad (IIIC stage) is more marked in females (Fig. 3B) in May (80%) and seems more precocious in males. In both sexes, this stage devel- oped in parallel from May to July so that pardal spawning occurred during these months. In August, the IIIB stage was preponderant in both sexes (80% of 34 Bergayou ET AL.: Reproduction of Cerastoderma edule in Southwestern Morocco A B 2001 Months 2002 2001 Months 2002 ■ Sta IIID m Sta MIC El Sta IIIB m Sta IIIA Sta II E Sta 1 □ Sta 0 ■ Sta IIID m Sta MIC E Sta IIIB m Sta IIIA a Sta II □ Sta □ Sta 0 Figure 3. Distribution of gonadic-development stages over 2001-2002 in the males (A) and females (B) of C. edule. The cumulated frequencies of the different developmental stages corresponded to the whole bivalves studied (100%). For details of each stage, see Materials and Methods (Table I). Figura 3. Distribución de los estadios de desarrollo gonadal sobre el periodo 2001-2001 en machos (A) y hembras (B) de C. edule. Las frecuencias acumuladas de distintos estadios de desarrollo sexual corre- sponden al total de bivalvos estudiados (100%). Véase Material y Métodos (Tabla I) para los detalles de cada estadio. males, 60% of females), proving that the main spawning episode had happened. The resorption of gonads (IIID stage) was observed in a few individuáis after the spawning of May and was less than 25% from June to August. Several females in IIID stage were noted during the other months, but they never exceeded 5%. In September, gametogen- esis started in both sexes (I and II stages were found in 95% of cockles). The mat- uration of the gonad (IIIA stage) was observed in the first bivalves from October to December and partial spawn- ing occurred from November to March. The highest valúes (2.7) of the gonadic Índex (Fig. 4) were noted between January and April, thus con- firming the presence of ripe gonads during this period. The lowest valúes (1.7) were found from June to August. In both sexes, the gonadic Índex was always higher than 1 so that the period of sexual rest was short. Gametogenesis in the population of C. edule: Figure 5 gives the results of stere- ological analysis. From January to March, the percentages of mature gametes increased to peak in April. Another episode of gametogenesis began in May, as demonstrated by the high frequencies of young germinal cells during this month in both sexes. In males, the effect of this developing sper- 35 Iberas, 26 (1), 2008 —°"“a — b Figure 4. Annual cycle of the gonadic Índex in the males (A) and females (B) of C. edule over 2001-2002. Figura 4. Ciclo anual del índice gonádico en machos (a) y hembras (b) de C. edule en el periodo 2001- 2002. matogenesis was high rates of sper- matids and spermatozoa in June or August (grouped frequencies, 75% and 85%, respectively). In females, an impor- tant vitellogenesis was observed in May and June so that the highest frequency of mature oocytes (55%) was noted in July and that of degenerated oocytes (60%) in August. In September, the highest percentages of spermatogoniae (29%) in males and of oogoniae (37%) in females were noted, thus indicating the starting of another gametogenetic wave. From September to December, the gonadic tissue of males was important, as the grouped frequencies of spermato- goniae and of spermatocytes were more than 47%. The same finding was noted in females during this period (grouped rates of oogoniae and of vitellogenic oocytes, >70%). Low percentages of atresic oocytes (5-25%) were always observed in females throughout the year. From the above study, it can be con- cluded that three successive gametoge- netic waves were differentiated: the first one from January to April, the second one from May to July, and the third one from September to March. Ripe cockles for both sexes were predominant from January to April, in July, and from January to March. Population dynamics: The lengths of C. edule , measured from January to December 2002, are given in Figure 6. The period of recruitment spanned from the end of spring to the onset of autumn. The first settlement of juvenile cockles occurred in June. A second cohort appeared in July, a third cohort in August, and a fourth one in October. At the end of autumn, the individuáis recruited in June-July had a length of 18.3 ±1.8 mm and constituted 71.6% of the population. By contrast, those origi- nating from the grouped cohorts of August and October only had a length of 11.1 ± 1.1 mm and represented 9.2% of the population. In addition to these recruitments; a few young cockles (2- 3%) were found for each date of sam- pling. In summer, there was a strong decrease in the number of cockles mea- suring 24.6 ±1.9 mm in length. This diminution is followed by the almost complete disappearance of adult cockles just after the period of recruitment (August 2002) so that the population of C. edule in 2002 was mainly composed of young individuáis which are being renewed each year. 36 BERGAYOU ET AL. : Reproduction of Cerastoderma edule in Sourhwestern Morocco ■ Spz Hfl Spc EH3 Spg B JFMAMJ JASONDJFM 2001 Months 2002 Figure 5. Frequencies of different cell categories in the male (A) and female (B) gonads of C. edule collected between January 2001 and March 2002. Male line: Spc (primary and secondary sperma- tocytes), Spg (protogoniae and spermatogoniae), Spz (spermatids and spermatozoa) . Female line: D (degenerated oocytes), Og (oogoniae), R (mature oocytes), V (vitellogenic oocytes). The cumu- lated frequencies of the different cell categories corresponded to the whole bivalves studied (100%). Figura 5. Frecuencia de distintas categorías de células en gónadas de machos (A) y hembras (B) de C. edule recolectados entre enero 2001 y marzo 2002. En machos: Spc (Espermatocitos primarios y secun- darios), Spg (protogonias y espermatogonias), Spz (espermatidios y espermatozoos); en hembras: D (ovoc- itos degenerados), Og ( oogonias ), R (ovocitos maduros), V (ovocitos vitelogénicos). Las frecuencias acu- muladas de distintos estadios de desarrollo sexual corresponden al total de bivalvos estudiados (1 00%). Spatial distribution, biomass, and structure of population of C. edule before and after wastewater discharges The spatial distribution of C. edule along the estuary, its density and its biomass are given in Table II for the summer surveys (2002 and 2003). During the period of wastewater dis- charge, the species was confined down- stream (stations 1 and 2). By contrast, in 2003, its distribution had extended with an upstream penetration of cockles up to the station 5. The density of C. edule did not exceed 320 individuals/m2 in 2002, and strongly increased in 2003 to reach valúes higher than 4,900 cockles/m2 in several places (these valúes were found during the periods of recruitment). The mean biomass (AFDW) calculated on the cockle samples collected during summer surveys, increased from 2.54 g/m2 in 2002 to 14.87 g/m2 in 2003. Table III shows the different groups of cockles constituting the population in summer surveys (2002 and 2003). 37 Iberus , 26 (1), 2008 Length (mm) 0 10 20 30 N=171 N=182 N=207 N=242 N=300 N=244 N=1 14 N=119 N=256 N=92 N=119 Figure 6. Size distribution of C. edule in the estuary of Oued Souss from January to December 2002 and the principal normal components. Month and numbers of individuáis measured (N) are given for each sampling. Figura 6. Distribución de tallas de C. edule en el estuario del Oued Souss entre enero y diciembre 2002 y componentes normales principales. Meses y número de ejemplares medidos (N) están indicados para cada muestra. During the wastewater discharges in the estuary (July 2002), individuáis are dis- tributed among four groups: two classes with lowest lengths (6.81 ± 1.53 mm and 9.97 ± 0.83 mm), deriving from a recruit- ment in May and June; and two classes of oíd cockles (24,72 ±2.17 mm and 27.5 ± 0,76 mm). In July 2003, the most fre- quent group (89.2%) was composed of young individuáis (length, 6.55 ± 1.17 mm; 11.76 ± 1.63 mm) originating from a recruitment in May and June. Another group of médium size (36.25 ± 1.73 mm) appeared for the first time and probably comprised cockles more than one year oíd, as the absence of growth lines on cockle valves did not allow to easily identify the different age subgroups constituting this last size group. DISCUSSION The results reported in the present study on the gonochorism of C. edule agree with the reports of several authors (Gimazane, 1971; Kingston, 1974). Several cases of accidental hermaphro- dism (4%0) were also noted by Fernan- dez Castro, Guillou, Le Pennec and Cárdenas López (1989). In the estuary of Oued Souss, the sex ratio of cockles was well-balanced so that pollution did not have an influence on the distribu- tion of males and females. This finding agrees with studies that some authors have performed in other populations of C. edule at different latitudes, such as the report by Kingston (1974) along the coasts of Kent (UK), that of Ivell (1981) 38 BERGAYOU ET AL.: Reproduction of Cerastoderma edule in Southwestern Morocco Table II. Spatial distribution of C. edule along the estuary, its density and its Biomass (AFDWg / m2) during wastewater contamination (summer 2002) and after stopping of pollution (summer 2003). Tabla II. Distribución espacial de C. edule a lo largodel estuario , su densidad y su biomasa (AFDW/m2) durante vertidos de agua contaminada ( verano 2002) y después de poner fin a la contaminación ( vera- no 2003). Sfátions Sampling sites 2002 2003 Density (ind/m2) 2002 2003 Biomass AFDW (g/m2) 0 OB 4 0.0012 ObisA 8/2 32.368 ObisB 24 0.08 1 1A 4 0.6492 IB 4 0.85 Ibis A 20 1.22 IbisB 4 24 0.582 3.12 TA 36 0.446 TB 16 0.48 2 2A 108 0.28 2B 6760 102.892 2bisA 320 5.539 2bisB 112 3.648 2'A 4972 83.768 3 3A 8 0.1384 3'A 432 3.7176 3'B 636 6.8256 3B 8 0.0416 4 4'A 16 0.074 4'B 124 2.0676 5 5B 68 0.5248 in the Limfjord (Denmark), that of Mejuto (1984) in Ria de Pasaxe (Spain), or still that of Fernandez Castro, Guillou, Le Pennec and Cárdenas López (1989) at Brouennou (France). Through methods used for repro- ductive cycle analysis, complementary and concordant results were noted. In the year, two successive gametogenetic waves, the first occurring from Septem- ber to April and the second in May-July, were differentiated in this population and spawning periods staggered over time. These phenomena can be explained by an effect of latitude (Sola, 1997; Rodriguez-Rua, Prado, Romeo and Bruzon, 2003) and, in particular, by water temperature (Hugues, 1971). When temperature was less than 10°C, it induced early spawning, followed by gonad reconstitution and a second period of spawning. The presence of atresic oocytes observed throughout the year during vitellogenesis, as mentioned by Lubet (1991) is apparently a frequent phenom- enon in bivalves noted in the start of gametogenesis (first mature oocytes degenerate), after partial spawnings and in the end of a reproductive cycle. The long period of cockle recruit- ment (from June to October) in 2002 can 39 Iberus, 26 (1), 2008 Table III. Distribution of shell lengths for C. edule during summer surveys (2002 and 2003): prin- cipal normal components. Tabla III. Distribución de longitudes de conchas de C. edule durante muéstreos de verano (2002 y 2003): principales componentes normales. Group N° Percentage (%) July 2002 Mean of shell length (mm) Structure of population S.D Percentage (%) July 2003 Mean of shell length (mm) S.D 1 19.5 6.81 1.53 11.4 6.55 1.17 2 16.4 9.9 7 0.83 77.8 11.76 1.63 3 55.5 24.72 2.17 10.68 36.25 1.73 4 8.4 27.5 0.76 be easily explained by spawning which occurred from April to August and this time was perfectly consistent with the gonadic-development stages deter- mined by the histological study of gonads and the examination of gonad smears. Contrary to juveniles recruited in June-July which had a better develop- ment, those settled from August to October showed a great mortality. This last result would not be related to wastewater pollution, in agreement with the reports by Madani (1989), Sauriau (1992), Bachelet, Guillou and Labourg (1992a), or with that by Bachelet, Desprez, Ducrotoy, Guillou, Labourg, Rybarczyk, Sauriau, Elkaim and Glémarec (1992b). According to these authors, high mortalities of cockles were noted in the recruitments which occurred at the end of summer, in autumn, and in winter. This mortality affected small- sized (<10 mm) cockles and may be explained by the almost complete absence of energetic reserves (Sauriau, 1992) whereas the metabolism of young cockles was changing (Gabbott, 1976). By contrast, the disappearance of medium-sized molluscs in 2002 coin- cided with the period of summer recruitment. This finding might be explained, either by a great predation of cockles by Haematopus ostralegus (this bird only tackled on médium sizes as reported by Atkinson, Clark, Bell, Daré, Clark and Ireland, 2003; John- stone and Norris, 2000), or by a high rate of mortality for these bivalves, as their vulnerability became more marked with increasing age and was also dependent on the quality of waters and/or an eventual eutrophication in the estuary, as demonstrated by Ducro- toy and Ibanez (2002). Indeed, as sea waters in winter and spring had high loads of mineral seston in the absence of pollution, the filter-feeders produced numerous pseudo-faeces and had a marked energetic expense for selective sorting of particles, mucus secretion, and cleaning of gills for Mytilus edulis (Widdows, Fieth and Worrall, 1979) or for Crassostrea gigas (Héral, 1986). According to Foster-Smith (1975a, b) and Newell (1977), the behaviour of adult cockles was different, as they adapted their pumping activity in the presence of high concentrations of mineral seston, with a passage by a rel- ative State of dormancy (Savari, Lock- wood and Sheader, 1991). However, such a behaviour did not seem to exist during wastewater discharges in summer, as cockles were in reproduc- tion and had to live in waters double loaded with matters in suspensión (pol- lution) and the phyto-planktonic bloom (with the increase of temperature), therefore placing these bivalves under stress conditions and inducing a high mortality. Since November 2002, the discharge of decanted wastewater in another site 40 Bergayou ET AL. : Reproduction of Cerastoderma edule in Southwestern Morocco beyond the estuary and the conversión of this ecosystem into a marine environ- ment had resulted, for C. edule, in a wider spatial distribution, a strong increase of density, and the appearance of médium sizes in this site. To comment on these changes, it is neces- sary to take into account the reproduc- tion of cockles living in this estuary, as it is stretched over all the year. Indeed, according to Gordo (1982), the repro- ductive cycle of Spanish and Portuguese populations of C. edule showed a period of sexual rest during summer months and the author explained it by the mean temperature of sea water which were more than 28°C and would inhibit the BIBLIOGRAPHY Ansell, A. D., Barnett, P. R. O., Bodoy, A. and Masse, H., 1981. Upper temperature toler- ances of some European molluscs. III. Cardium glaucum, C. tuberculatum and C. ed- ule. Marine Biology, 65: 177-183. Atkinson, P. W., Clark, N. A., Bell, M. C., Daré, P. J., Clark, J. A. and Ireland, P. L., 2003. Changes in commercially fished shell- fish stocks and shorebird populations in the Wash, England. Biological Conservation, 114: 127-141. Bachelet, G., Bouchet, J. M. and Lissalde, J. P., 1981. Les peuplements benthiques de la Gironde: biomasse, productivité et évolu- tion structurale. Oceanis, 6: 593-620. Bachelet, G., Guillou, J., Labourg, P. J., 1992a. Adult, larval and juvenile interactions in the suspension-feedingbivalve, Cerastoderma ed- ule: field observations and experiments. In Colombo, G., Ferrari, I., Ceecherelli, V.U., and Rossi, R., (Eds.): Marine eutrophisation and population dynamics. Fredensborg, 175- 182. Bachelet, G., Desprez, M., Ducrotoy, J. P., Guillou, J., Labourg, P. J., Rybarczyk, H., Sauriau, P. G., Elkaim, B. and Glé- marec, M., 1992b. Role de la compétition intraspécifique dans la régulation du re- crutement chez la coque, Cerastoderma edule (L). Anuales de l'Institut Océanographique, 68:75-87. Battacharya, C. G., 1967. A simple method of resolution of a distribution into Gaussian components. Biometrics, 23: 115-135. Bayed, A., 1982. Ecologie descriptive et dynamique des plages de la región de Rabat, Maroc. Doctoral Thesis, Brest, 122 pp. reproduction of this species, as they were cióse from the lethal temperature recorded for C. edule (Ansell, Barnett, Bodoy and Masse, 1981). As the cockles from the estuary of the Oued Souss river practically did not have sexual rest and showed a gametogenesis starting towards autumn, the results noted in this site might be interpreted as an adaptation of this cockle population to high temperatures of sea waters which exist in the South of Morocco. Under these conditions, it is logical to consider C. edule an opportunistic species which might quickly overtake new areas in presence of favourable environmental conditions. Ducrotoy, J. P. and Ibanez, F., 2002. Ecolog- ical groups of estuarine macrobenthic in- vertebrates in the Baie de Somme (France): changes in time and space. Journal ofthe Ma- rine Biological Association of the United King- dom, 5: 749-769. Fernández-Castro, N., Guillou, J., Le Pen- nec, M. and Cárdenas López, J. J., 1989. Le cycle sexuel de Cerastoderma edule L. (Bivalve: Cardiidae) á Brouennou (Finistére). Haliotis, 19: 325-334. Foster-Smith, R. L., 1975a. The effect of con- centration of suspensión on the filtration rates and pseudofaecal production for Mytilus edulis (L.), Cerastoderma edule (L.) and Venerupis pul- lastra (Montagu). Journal of Experimental Marine Biology and Ecology, 17: 1-22. Foster-Smith, R. L., 1975b. The role of mucus in the mechanism of feeding in three filter- feeding bivalves. Proceedings of the Malaco- logical Society ofLondon, 41: 571-588. Gabbott, P. A., 1976. Energy metabolism. In Bayne, P. A. (Ed.): Marine mussels: ecology and physiology. Cambridge University Press, Cambridge, 213-355. Gabe, M., 1968. Techniques histologiques. Masson et Cié, Paris, 1113 pp. Gayanilo, F. C. Jr., Sparke, P. and Pauly, P., 1996. FISAT Software. FAO-ICLARM-Stock- Assessment-Tools, Makati City, Metro Manila, The Philippines. Gimazane, J. P., 1971. Introduction a Vétude ex- périmentále du cycle sexuel d’un mollusque Bi- valve Cardium edule L. Analyse des populations, évolution de la gonade et action de quelquesfac- teurs: nutrition, température, photopériode. Doc- toral thesis, University of Caen, 112 pp. 41 Iberus, 26 (1), 2008 Gordo, L. S., 1982. Aspects of the growth and density of Cerastoderma edule in the Alvor Salt Marsh, South Portugal. Arquivos do Museu Bocage (Series A), 19: 435-452. Guillou, J., Bachelet, G. and Glémarec, M., 1992. Influence des fluctuations de tem- pérature sur la reproduction et le recrute- ment de la coque Cerastoderma edule (L). An- uales de l'Institut Océanographique , 68: 65-74 Héral, M., 1986. L'ostréiculture frangaise tra- ditionnelle. In Barnabe, G., (Ed): Aquacul- ture, 2nd ed. Lavoisier, París. 345-390. Hugues, R. N., 1971. Reproduction of Scrobic- ularia plana Da Costa (Pelecypoda: Semelidae) in North Wales. The Veliger, 14: 77-81. Ivell, R., 1981. A quantitative study of a Ceras- toderma-Nephthys community in the Lim- fjord, Denmark, with special reference to production of Cerastoderma edule. Journal of Molluscan Studies, 47: 147-170. Johnstone, I. and Norris, K., 2000. The influ- ence of sediment type on the aggregative re- sponse of oystercachers, Haematopus ostrale- gus, searching for cockles, Cerastoderma edule. Oikos, 89: 146-154. Kingston, P. F., 1974. Studies on the repro- ductive cycles of Cardium edule and C. glau- cum. Marine Biology, 28: 317-323. Lubet, P., 1991. Reproduction des mollusques In Barnabé, G., (Ed.): Bases biologiques et écologiques de l'aquaculture. Lavoisier, Tec and Doc, París, 167-210. Lubet, P., 1959. Recherches sur le cycle sexuel et l'émission des gamétes chez les Mytilidés et les Pectinidés. Doctoral Thesis, University of París, 159 pp. Lucas, A., 1965. Recherches sur la sexualité des Mollusques Bivalves. Bulletin Biologique de Trance et de Belgique, 99: 115-247. Lucas, A. and Beninger, P. G., 1985. The use of physiological condition indices in marine bivalve aquaculture. Aquaculture, 44: 187- 200. Madani, I., 1989. Dynamique des populations et processus de recrutement chez quatre espéces de bivalves appartenant aux genres Abra et Ceras- toderma, dans le bassin d'Ar cachón. Doctoral Thesis, University of Bordeaux 1, 152 pp. Mejuto, J., 1984. Primeros datos sobre la di- námica de la poblation de Cerastoderma edule L.) de la Ria do Pasaxe ( NW de Galicia), es- trategias de explotation. Actas IVo Simposio Ibérico de Estudos do Benthos Marinho, 2: 83-102. Newell, R. I. E., 1977. The ecophysiology of Cardium edule (Linné). Ph. D. Thesis. Uni- versity of London, 250 pp. Rodríguez-Rua, A., Prado, M. A., Romeo, Z. andBruzon, M., 2003. The gametogenic cy- cle of Scrobicularia plana (Da Costa, 1778) (Mollusca: Bivalvia) in Guadalquivir estu- ary (Cádiz, SW Spain). Aquaculture, 217: 157- 166. Sauriau, P. G., 1992. Les mollusques benthiques du bassin de Marennes-Oléron: estimation et cartographie des stocks non cultives, compétition spatiale et trophique, dynamique de population de Cerastoderma edule (L.). Doctoral Thesis, University of Nantes, 262 pp. Savari, A., Lockwood, A. P. M. and Sheader, M., 1991. Variations in the physiological State of the common cockle ( Cerastoderma edule (L.)) in the laboratory and in Southampton Water. Journal of Molluscan Studies, 57: 33-44. Seed, R., 1975. Reproduction in Mytilus edulis L. (Mollusca: Bivalvia) in European waters. Pubblicazioni delta Stazione Zoológica di Napoli, 39: 317-334. Seed, R. and Suchanek, T. H., 1992. Popula- tion and community ecology of Mytilus. Chapter 4. In Gosling, E., (Ed.): The mussel Mytilus: ecology, physiology, genetics and cul- ture. Developments in Aquaculture and Fish- eries Science, n° 25. Elsevier, Amster- dam/New York, 87-168. Sola, J. C., 1997. Reproduction, population dy- namics growth, and production of Scrobicu- laria plana Da Costa (Pelecypoda) in the Bida- soa estuary, Spain. Netherlands Journal of Aquatic Ecology, 30: 283-296. Widdows, J., Donkin, P., 1992. Mussels and environmental contaminants: bioaccumula- tion and physiological aspects. In Gosling, E., (Ed.): The mussel Mytilus: ecology, physiology, genetics and culture. Developments in Aqua- culture and Fisheries Science, n° 25. Elsevier, Amsterdam/New York, 383-424. Widdows, J., Fieth, P. and Worrall, C. M., 1979. Relationship between seston, available food and feeding activity in the common mussel Mytilus edulis. Marine Biology, 50: 195- 207. 42 © Sociedad Española de Malacología Iberus , 26 (1): 43-46, 2008 Xylodiscula wareni n. sp., una nuova specie per le coste orientali della Sicilia Xylodiscula wareni n. sp., una nueva specie para las costas orientales de Sicilia Cesare BOGI* e Stefano BARTOLINI** Recibido el 27-X-2007. Aceptado el 15-XII-2007 RIASSUNTO Si descrive un piccolo gasteropode planispirale, che non corrisponde a nessuna delle spe- cie attualmente viventi nel Mar Mediterráneo, trovato in campioni di sedimento raccolti a profonditá cómprese tra i 35 e i 52 m lungo le coste siciliane. La forma generóle della conchiglia ci ha indotto ad ascrivere questa specie, solo prowisoriamente, al genere Xylo- discula Marshall, 1988 anche se alcune caratteristiche della protoconca, non iperstrofica, lo escluderebbero. Questa specie si descrive come nuova, con il nome di Xylodiscula wareni anche se proba- bilmente non appartiene a questo genere, evitando di descrivere al momento un nuovo genere scarsamente definito. ABSTRACT A small planispiral gastropod, which could not be ascribed to any known species from the Mediterranean or adjacent seas, was found in sediment samples collected in depths between 35 and 52 m in the seas around Sicily. The general shape resmbles the genus Xylodiscula Marshall, 1 988, but the protoconch ¡s not heterostrophic and has a distinct sculpture not described from any gastropod. There- fore we describe this new species as Xylodiscula wareni, although it probably does not belong to the genus, instead of creating a new, poorly defined genus. RESUMEN Se describe un pequeño gasterópodo planiespiral recolectado entre 35 y 52 m de profun- didad en las costas Sicilianas, que no pudo ser asignado a ninguna especie conocida del Mediterráneo ni de mares adyacentes. La forma general se asemeja al género Xylodiscula Marshall, 1 988, aunque la protoconcha no sea heterostrófica y tenga una escultura dis- tinta que no se conoce en ningún otro gasterópodo. Se propone como nueva especie Xylodiscula wareni. Aunque probablemente no pertenezca a este género, se descarta de momento la descripción de un género nuevo escasamente definido. PAROLE CHIAVE: Gastropoda, Xylodisculidae, Xylodiscula wareni, nuova specie, Mar Mediterráneo, recente. KEY WORDS: Gastropoda, Xylodisculidae, Xylodiscula wareni , new species, Mediterranean Sea, recent. PALABRAS CLAVE: Gastropoda, Xylodisculidae, Xylodiscula wareni , nueva especie, mar Mediterráneo, reciente. * Via delle Viole 7 1-57124, Livorno. e-mail bogicesare@tiscali.it ** Via E. Zacconi,l6 1-50137, Firenze. e-mail stefmaria.bartolini@libero.it 43 Iberus , 26 (1), 2008 INTRODUZIONE Le nostre continué ricerche malaco- logiche e Pesame di numerosi campioni di sedimento raccolti lungo le coste della Sicilia Nord-Orientale e le isole antistanti, ci ha portato, questa volta, a studiare alcuni micromolluschi, in tutto quattro esemplari, isolati da piccole quantitá di detrito raccolto a profonditá variabili tra i 35 mt. e i 52 mt. II piccolo gasteropode di forma pla- norbide viene, per l'aspetto generale della conchiglia, solo provvisoriamente attribuito al genere Xylodiscula Marshall, 1988, in quanto le caratteristiche della sua protoconca, non iperstrofica, lo escluderebbero . SISTEMATICA Ordine Heterostropha Fischer P. , 1885 Famiglia Xylodisculidae Waren A., 1992 Genere Xylodiscula Marshall,1988 Xylodiscula wareni n. sp. (Fig. 1) Materiale esaminato: 1 esemplare (olotipo, fig.l a-e) proveniente da detriti raccolti nel Luglio 2005 alia base della "Secca dei 6 metri" presso l'isola di Filicudi a -50 m. , 1 esemplare (paratipo A) rac- colto nel Giugno 2006 a Cannizzaro, loe. Bellatrix, -35 m.di profonditá lungo una párete rocciosa, 2 esemplari (paratipi B e C) provenienti da campioni di detriti coralligeni raccolti a Scilla a -52 m. Materiale tipo: L'olotipo (Diam.= 1.1 mm), (Fig. 1A-E ) é stato depositato nella collezione malaco- logia del Museo di Storia Naturale del Mediterráneo di Livorno, Italia, con il numero: Malacologia Vol. V, n. 734. 1 paratipi sono conservad nelle seguenti collezioni: il paratipo A (Diam.= 1.0 mm), nella collezione di S. Bartolini (Firenze), il paratipo B (Diam.= 1,3 mm) , e il paratipo C (Diam.= 1,2 mm), nella collezione E. Perna (Napoli). Locus typicus: Detriti raccolti alia base della "Secca dei 6 metri" presso Pisóla di Filicudi a -50 m. di profonditá. Etimología: La specie é stata dedicata al Dottor Anders Warén, malacologo di fama mondiale e specialista, tra l'altro, nei piccoli "Skeneimorpha". Diagnosi (olotipo): Conchiglia piccola, fragüe, trasparente, di forma planor- bide.La protoconca (fig.l e) é costituita da 0.7 giri di spira di cui la parte iniziale é scolpita da piccole e irregolari depres- sioni.La teleoconca é costituita da circa 2 giri di spira attraversati da deboli linee di accrescimento. I giri si toccano solo per un breve tratto piú o meno alia peri- feria del giro precedente e quindi la sutura risulta profonda e leggermente canaliculata. Spira arrotondata con la protoconca di poco sporgente nella parte superiore della stessa. Alia base della conchiglia é ben visibile per tutta la lunghezza della teleoconca, una evi- dente carena posizionata internamente all'ombelico. La bocea é di forma roton- deggiante con la parte inferiore legger- mente espansa in prossimitá della carena. La larghezza dell'ombelico é circa il 30% del diámetro delPintera con- chiglia. Dimensioni : Diámetro^ 1.1 mm. Opercolo e partí molli ancora scono- sciute. Distribuzzione: La specie sembra avere una distribuzione ristretta a poche localitá delle coste siciliane, ed il suo habitat non sembra essere mai superfi- ciale. Discussione : L'attribuzione di questi esemplari al genere Xylodiscula é da rite- nersi provvisoria in quanto solo lo studio delle partí molli potranno dame un'inquadramento tassonomico piú congeniale. La forma generale della con- chiglia in realtá corrisponde alie caratte- ristiche tipiche di questo genere, ma la protoconca differisce in quanto non ha uno sviluppo iperstrofico típico della famiglia Xylodisculidae Warén, 1992. 44 Bogi E BartolinI: Xylodiscula wareni n.sp., una nuova specie per le coste della Sicilia Figura 1. Xylodiscula wareni n. sp., Filicudi -50 m, olotipo (Museo di Storia Naturale del Mediter- ráneo di Livorno, Coll. Malac. N° Malacologia Vol. V n. 734). A: vista dall’alto; B: vista basale; C: vista frontale; D: dettaglio dell’ombelico; E: dettaglio della protoconca. Figura 1. Xylodiscula wareni n.sp., Filicudi -50 m, holotype (Museo di Storia Naturale del Mediterrá- neo di Livorno, Coll. Malac. N° Malacologia Vol. V n. 734). A: apical view; B: basal view; C: frontal view; D: detail ofthe umbilicus; E: detall ofthe protoconch. II genere Xylodiscula é attualmente rappresentato nel Mar Mediterráneo da due specie: X. boucheti Warén, Carrozza e Rocchini, 1992, e X. leus Warén, 1992. Warén stesso nel suo articolo (Warén, 1992) evidenzia la difficoltá nel distin- guere le due specie che differiscono quasi esclusivamente per il diámetro 45 Iberus, 26 (1), 2008 dell'ombelico che é il 30-33 % del diáme- tro deH'intera conchiglia in X. boucheti mentre in X. lens é il 40%. La caratteri- stica principale che distingue X. wareni da queste due specie, come dalle altre non mediterranee appartenenti a questo genere, oltre alia diversa protoconca, é la presenza di una carena ben visibile alia base della conchiglia (Fig. IB). Abbiamo dubitato che gli esemplari potessero appartenere ad una specie dulciacquicola o terrestre, ma le nostre ricerche e l'opinioni di alcuni specialisti hanno escluso questa possibilitá. RINGRAZIAMENTI Un particolare ringraziamento al Dottor Ánders Warén del Museo Svedese di Storia Naturale di Stoccolma, per aver eseguito le foto al SEM e per i consigli dati nella rilettura critica dell' articolo, e all'a- BIBLIOGRAFIA Marshall, B.A., 1988. Skeneidae, Vitrinelli- dae and Orbitestellidae (Mollusca: Gastro- poda) associated with biogenic substraía from bathyal depths off New Zealand and New South Wales. Journal of Natural History, 22: 949-1004. La specie tipo del genere, Xylodi- scula vitrea Marshall, 1988, come puré X. eximia Marshall, 1988, originarle rispetti- vamente dell'Australia e della Nuova Zelanda, sono state tróvate su pezzi di legno affondati ed anche le specie descritte da Warén per il Mar Mediterrá- neo sono state raccolte viventi in un biotopo caratterizzato da fibre di Posido- nia e frammenti di legno affondati. Nei campioni di sedimento dove noi abbiamo trovato i pochi esemplari di X. wareni, non erano presenti frammenti di Posidonia. mico Edoardo Perna per aver messo a disposizione i suoi esemplari nonché le foto degli stessi. All'amico Ivano Niero per aver studiato gli esemplari come even- tuali appartenenti alia fauna continentale. Warén A., 1992. New and little known "Ske- neimorph" gastropods from the Mediterra- nean Sea and the adiacent Atlantic Ocean. Bol- lettino Malacologico, 27 (10-12):149-248. 46 © Sociedad Española de Malaca logia Iberus , 26 (1): 47-51, 2008 The colour white diminishes weight loss during aestivation in the arid-dwelling land snail Sphincterochila (Albea) can - didissima El color blanco disminuye la pérdida de peso durante la estivación en el caracol de medios áridos Sphincterochila (Albea) candidissima Gregorio MORENO-RUEDA* Recibido el 14-V-2007. Aceptado el 2-1-2008 ABSTRACT It has been suggested that white colour is beneficial for snails living ¡n arid environments. In this work, shell coloration in the arid-dwelling land snail Sphincterochila (Albea) can- didissima was manipulated during aestivation. Snails painted black lost more body weight than did control ones, presumably as a consequence of higher heat absorption. This sug- gests that light colour is advantageous for this land snail. RESUMEN Se ha sugerido que el color blanco es beneficioso para los caracoles en ambientes ári- dos. En el presente trabajo se manipula la coloración de la concha en el caracol de medios áridos Sphincterochila (Albea) candidissima durante la estivación. Los caracoles pintados de negro perdieron más peso que los caracoles que sirvieron como control, pre- sumiblemente como una consecuencia de una mayor absorción de calor. Esto sugiere que los colores claros son ventajosos para este caracol. KEY WORDS: Sphincterochila candidissima , arid environments, colouration. PALABRAS CLAVE: Sphincterochila candidissima , medios áridos, coloración. INTRODUCTION It is well established that tegument colour has important functions in camou- flage, as well as in communication (e.g., Badayev and Hill, 2000; Théry, Debut, Gómez and Casas, 2005; Exnerova, SVADORA, BARCALOVA, LANDOVA, Prokopova, Fuchs and Socha, 2006). The colour of teguments depends on the wavelengths that are reflected. Therefore, colouration affects the energy that is absorbed by the tegument, the amount being higher as the colour darkens. In this sense, animal colouration may also have a role in thermoregulation, espe- cially important for arid-dwelling animáis (Cloudsley-Thompson, 1978). Terrestrial molluscs are very susceptible to dehydration (Prior, 1985; Luchtel and Deyrup-Olsen, 2001), and therefore need adaptations to survive in arid envi- * Konrad Lorenz Institut für Vergleichende Verhaltensforschung, Ósterreischische Academie der Wissenschaften, SavoyenstraEe la, A- 1160, Wien (Austria) and Departamento de Biología Animal, Facultad de Ciencias, Universidad de Granada, E- 18071, Granada (Spain). 47 Iberus , 26 (1), 2008 ronments. White colour may favour the success of land-snails in warm environ- ments: White shells have a higher reflectance of sunlight (Schmidt- Nielsen, Taylor and Shkolnik, 1971), and, as a consequence, land snails with light-coloured shells register lower body temperatures (Heath, 1975), thereby improving survival under high tempera- tures (Richardson, 1974). This translates as a selective pressure for whiter shells in arid environments and, in fact, some studies have correlated shell colour with environment temperature (Jones, 1973; but there are exceptions: Heller, 1984). The snails of the genus Sphincterochila have white shells. In Sierra Elvira (SE Spain), Sphincterochila (Albea) candidísi- ma (Draparnaud, 1801) is the most abun- dant gastropod (Moreno-Rueda, 2002). Sierra Elvira has an arid environment, and sheltering in protective microhabi- tats may be a vital strategy against dehy- dration (Steinberger, Grossman, Du- binsky and Shachak, 1983; Arad, Goldenberg and Heller, 1989; Cook, 2001). However, S. candidísima does not use refuges during drought periods (spring and summer) (Moreno-Rueda, 2007; Moreno-Rueda and Collantes- Martín, 2007). Because the shell of S. candidísima is puré white, I hypothe- sized that shell colour might contribute to the survival of S. candidísima in Sierra Elvira, explaining why this snail does not need refuges against dehydration. I investigated this hypothesis by manipu- lating the shell colour of S. candidísima during aestivation (painting some snails), and by examining the effect of this manipulation on the amount of body mass lost. About 80-90% of fresh body weight (shell not included) of Sphincterochila land snails is water, and, therefore, a decrease in body weight sug- gests a loss of water (Schmidt-Nielsen et al., 1971; Yom-Tov, 1971; Steinberg- er, Grossman and Dubinsky, 1981). METHODS This study was performed in Sierra Elvira (SE Spain, 37° 15' N, 3o 40' W), a small mountain range with a dry mesomediterranean climate (UNESCO, 1963). The study area undergoes five months of drought each year, with an averáge annual precipitation of 600-1000 mm (Alonso, López- Alcántara, Rivas and IbáÑez, 1985). It is, therefore, a dry zone for land snails. Table I presents cli- matic data during the study period, measured from the meteorological station of Pinos Puente, about three kilo- metres from the study area, and approx- imately at the same altitude (630 m. a.s.l.). Sphincterochila candidísima is the only species of the genus Sphincterochila in Sierra Elvira (Ruiz Ruiz, Cárcaba Pozo, Porras Crevillén and Arrébola Burgos, 2006). In the study area, this species begins aestivation in April-May (Moreno-Rueda, 2007; Moreno-Rueda and Collantes- Martín, 2007). This snail adheres to rock or vegetation during aestivation. For manipulation, snails were not sepa- rated from the substrate, because this could provoke dehydration in the snails (Luchtel and Deyrup-Olsen, 2001). For this reason, I could not measure body mass before treatment, but I col- lected 75 additional individuáis in order to analyse the relationship between shell morphology and body mass. The experiment started on 26 June 2005. Each individual found was sequentially assigned to the control group (C), to the control of manipula- tion group (CM), or to the experimental group (E). Manipulation in control group was only a mark for recognition. Snails in the CM group were painted in yellow with a marker. Paint covered approximately 50% of shell surface. In the experimental group, the shell was painted black with a marker in the same way as in CM group. In total, 52 snails were used in each group (n = 156). The study area was prospected two months later (26 August 2005). Snails found were collected and mea- sured (shell height and width) with a calliper (accuracy 0.01 mm.) and weighed with a digital balance (accu- racy 0.1 g.). 48 MORENO Rueda: Colour white and aestivation weight loss in Sphincterochila candidissima Table I. Climatic data of the meteorological station of Pinos Puente, located near of the study area, for the study period (06/26/2005 to 08/26/2005). Tabla I. Datos climáticos de la estación meteorológica de Pinos Puente, próxima a la zona de estudio, para el período de estudio (2610612005 a 26/08/2005). Mean S.E. Mínimum Máximum Daily maximal íemperature (9C) 36.8 0.36 30.7 42.9 Daily minimal temperature (9C) 17.6 0.30 12.7 22.8 Average daily temperature (9C) 27.0 0.27 23.4 31.3 Daily radiation (MJ/m2) 27.6 0.51 13.9 32.5 Daily precipitation (mm.) 0.03 0.02 0.00 0.80 Table II. Average weight on the day 08/26/2005, and shell height and width for the snails in the experimental (E), control (C) and control of manipulation (CM) groups. The last row shows the average weight after statistically controlling for shell height and width. The last column shows the results of ANOVA and ANCOVA. In brackets is the standard error. Tabla II. Peso promedio el día 26/08/2005, y altura y anchura de la concha para los caracoles en el grupo experimental (E), control (C) y control de la manipulación ( CM). La última fila muestra el peso promedio después de controlar estadísticamente por la altura y anchura de la concha. La última columna muestra los resultados de los tests de ANOVA y ANCOVA. Entre paréntesis el error estándar. E group n= 29 C group n= 34 CM group /7 = 30 F ANOVA d.f. P Weight (g.) 2.70 (0.09) 3.02 (0.09) 2.87 (0.09) 3.15 2, 90 <0.05 Width (mm.) 21.49 (0.22) 21.31 (0.21) 21.33 (0.22) 0.21 2, 90 0.81 Height (mm.) 16.54 (0.18) 16.77 (0.17) 16.37 (0.18) 1.33 2, 90 0.27 Weight (controlled for shell size) 2.69 (0.07) 3.00 (0.06) 2.91 (0.07) 6.28 2, 88 <0.003 The variables had a distribution similar to normal (Kolmogorov-Smirnov test, p > 0.05), and parametric statistics were used. An ANOVA was used to test the effect of the treatment on body weight, and an ANCOVA was used con- trolling by snail body size (height and width). For post hoc comparisons the Fisher LSD test was used. The Chi- square was used to test the probability of survival according to treatment. RESULTS In August, I recaptured 34 snails alive in the control group, 30 for the CM group, and 29 of the experimental group. The frequency of recaptures did not differ significantly between the three groups (x22= 1.12; p= 0.57). When indi- viduáis were collected in August, there were significant differences for body mass between the treatments (Table II). Individuáis of the experimental group weighed less than those in the control one (post hoc Fisher LSD, p = 0.01), while the average weight in the CM group was intermediate between the other two groups (post hoc, CM vs. C, p = 0.22; CM vs. E, p = 0.22). In the addi- tional sample of 75 individuáis, body mass was strongly predicted by shell morphology (Múltiple Regression Model; R2 = 0.81; F 2,72 = 149.0; p < 0.001; equation: Body mass = -5.16 (SE = 0.46; t72 = 11.2) + 0.28 (SE = 0.03; t72 = 10.2) x Width + 0.14 (SE = 0.03; t72 = 5.1) x Height). There were no significant dif- ferences for body size (height and 49 Iberus , 26 (1), 2008 width) between the three groups (MANOVA, Wilks = 0.94; F4, ízs = 1.46; p = 0.22; Table II), suggesting that initial body mass did not differ among groups. When the analyses were repeated with shell height and width as a covariate, differences in weight between groups were accentuated (ANCOVA, Vi, 88 = 6.28; p < 0.003; Height effect: Fi, 88 = 10.90; p = 0.001; Width: Fi,88 = 21.98; p < 0.001; Table II). Differences between the control group and the experimental group increased in this analysis (post hoc, p < 0.001), while body weight in the CM group remained intermediate between the other two groups (CM vs. C, p = 0.08; CM vs. E, p = 0.08). DISCUSSION The findings of this experimental study show that shell colour alteration in Sphincterochila candidissima during aestivation had effects on weight loss, snails with shells painted black suffer- ing a quicker loss of weight than control snails. The most probable mechanism behind this result is that light reflection was lower in shells painted in black, and for this reason they trapped more heat, as shown in other studies with other species of snails (e.g., Heath, 1975). The higher the body temperature, the higher the water loss, decreasing body weight (Yom-Tov, 1971). Body size may affect the interaction between shell colour and heat absorption (Slotow, Goodfriend and Ward, 1993), but there were no differences in body size between the groups, and dif- ferences in weight remained significant after controlling statistically for shell morphology. This weight loss presum- ably harms fitness, increasing the risk of mortality, especially for the smallest individuáis (with less reserves), or in very dry years (Richardson, 1974). Moreover, the treatment lasted only two months (although the warmest), but S. candidissima aestivates for 5-7 months in the study area (Moreno-Rueda and Collantes-Martín, 2007), and there- fore, the effect should be more accentu- ated if the entire aestivation period is considered. The control of manipulation (CM) group, with shells painted in yellow, had weight valúes intermediate to the other two groups. In fact, this group is not a true control of manipulation, as colour was altered with respect to unmanipulated snails. Because their shells were darker than shells in the control group, but lighter than shells in the experimental group, the results support that weight loss is due to shell colour. As weight for the CM group was intermediate, if there was an effect of paint on weight, this cannot completely explain the differences between the control and the experimental group. Therefore, results presented here and in the literature strongly suggest that the white colour is advantageous for survival in arid-dwelling snails. The question arises as to why white colour is not more widespread in arid environ- ments (see, for example, Heller, 1984). Other selective mechanisms besides thermal selection act on shell colour, such as predation (Jones, Leith and Rawlings, 1977). Lighter shells, espe- cially those with a puré white colour as in Sphincterochila candidissima , are usually easier to detect by predators (Reed and Janzen, 1999). However, predation on Sphincterochila candidissima is rare due to its thick shell (Yanes, Suárez and Manrique, 1991), and thus the absence of a strong selection by predators in this species would favour the maintenance of puré white shells. In conclusión, this study, applying an experimental approach, supports the hypothesis that shell colour affects weight loss in arid-dwelling land-snails, and, as a consequence, fitness, by a mechanism mediated by thermoregula- tion. ACKNOWLEDGEMENTS Carlos Marfil Daza and José Luís Ros Santaella collaborated in the field work. David Nesbitt improved the English. 50 MORENO Rueda: Colour white and aestivation weight loss in Sphincterochila candidissima BIBLIOGRAPHY Alonso, M. R., López- Alcántara, A., Rivas, P. and IbáÑEZ, M., 1985. A biogeographic study of Iberus gualtierianus (L.) (Pulmonata: Helicidae). Soosiana, 13: 1-10. Arad, Z., Goldenberg, S. and Heller, ]., 1989. Resistance to desiccation and distribution pattems in the land snail Sphincterochila. Jour- nal ofZoology, 218: 353-364. Badyaev, A. V. and Hill, G. E., 2000. Evolu- tion of sexual dichromatism: contribution of carotenoid- versus melanin-based coloration. Biological Journal of the Linnean Society, 69: 153-172. Cloudsley-Thompson, J. L., 1978. Adaptive function of the colour s of desert animáis. Comparative Physiological Ecology, 1: 109-120. Cook, A., 2001. Behavioural ecology: On doing the right thing, in the right place at the right time. In Barker, G. M. (Ed.): The Biology ofTer- restrial Molluscs. CAB International. Walling- ford. Pp. 447-487. Exnerova, A., Svadora, K., Barcalova, S., Landova, E., Prokopova, M., Fuchs, R. and Socha, R., 2006. Importance of colour in the reaction of passerine predators to aposematic prey: experiments with mutants of Pyrrhocoris apterus (Heteroptera). Biological Journal ofthe Linnean Society, 88: 143-153. Heath, D. J., 1975. Colour, sunlight and inter- nal temperatures in the land-snail Cepaea nemoralis (L.). Oecologia, 19: 29-38. Heller, J., 1984. Shell colours of desert land- snails. Malacologia, 25: 355-359. Jones, J. S., 1973. Ecological genetics and natural selection in molluscs. Science, 182: 546-552. Jones, J. S., Leith, B. H. and Rawlings, P., 1977. Polymorphism in Cepaea: a problem with too many Solutions? Annual Review in Ecology and Systematics, 8: 109-143. Luchtel, D. L., Deyrup-Olsen, I., 2001. Body wall: form and function. In Barker, G. M. (Ed.): The Biology ofTerrestrial Molluscs. CAB International. Wallingford. Pp. 147-178. Moreno-Rueda, G., 2002. Selección de hábitat por Iberus gualtierianus, Rumina decollata y Sphincterochila candidissima (Gastropoda: Pul- monata) en una sierra del sureste español. Iberus, 20: 55-62. Moreno-Rueda, G., 2007. Refuge selection by two sympatric species of arid-dwelling land snails: Different adaptive strategies to achieve the same objective. Journal of Arid Enviro- ments, 68: 588-598. Moreno-Rueda, G. and Collantes-Martín, E., 2007. Ciclo anual de actividad del caracol Sphincterochila (Albea) candidissima (Dra- pamaud, 1801) en un medio semiárido. Iberus, 25: 49-56. Prior, D. ]., 1985. Water-regulatory behaviour in terrestrial gastropods. 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Some aspects of the ecology of the desert snail Sphincterochila prophetarum in re- lation to energy and water flow. Oecologia, 50: 103-108. Steinberger, Y., Grossman, S., Dubinsky, Z. and Shachak, M., 1983. Stone microhabitats and the movement and activity of desert snails, Sphincterochila prophetarum. Malaco- logical Review, 16: 63-70. Théry, M., Debut, M., Gómez, D. and Casas, J., 2005. Specific color sensitivities of prey and predator explain camouflage in differ- ent visual Systems. Behavioral Ecology, 16 : 25- 29. UNESCO, 1963. Recherches sur la zone aride. Etude écologique de la zone méditerraéenne. Cañe bioclimatique de la zone méditerranéenne. Notice explicative. UNESCO. Paris. Yanes, M., Suárez, F. and Manrique, J., 1991. La cogujada montesina, Galerida theklae, como depredador del caracol Otala lactea: com- portamiento alimenticio y selección de presa. Ardeola, 38: 297-303. Yom-Tov, Y., 1971. Annual fluctuations in the water contení of desert snails. Malacological Review, 4: 121-126. 51 1 Iberus, 26 (1): 53-63, 2008 © Sociedad Española de Malacologta Notes on the genus Anadema H. and A. Adams, 1854 (Gastropoda: Colloniidae) Notas sobre el género Anadema H. y A. Adams, 1854 (Gastropoda: Colloniidae) James H. MCLEAN* and Serge GOFAS** Recibido el 15-1-2008. Aceptado el 23-IV-2008 ABSTRACT Shell morphology and characters of the living animal of the poorly known, Atlantic Moroc- can species Anadema mocandrewii (Mórch, 1 864) are described and illustrated, based on beach collected specimens and a single live-collected specimen. The genus ¡s mono- typic and is assigned to the Colloniidae rather than Turbinidae because of the dome- shaped profile of the shell, open umbilicus, symmetrical tooth rows of the radula, lack of cephalic lappets, and the non-bicarinate ¡uvenile shell. Within the Colloniidae, it unusual for its relatively large mature shell, ¡uvenile shell with a keeled profile, and the lack of the secondary flap above the rachidian tooth. The species is regarded as sexually dimorphic, with the female shell having a raised periumbilical rim comparable to that of other tro- choideans modified for brooding by means of an enlarged umbilical cavity. RESUMEN Se describe e ilustra la morfología de la concha y del animal vivo de Anadema macan- drewii (Mórch, 1864), una especie poco conocida de la costa atlántica de Marruecos. El género es monotípico y se asigna a la familia Colloniidae, en lugar de a los Turbinidae por la forma abombada de la concha, el ombligo abierto, las filas de dientes radulares simétricas, la ausencia de lóbulos cefálicos y por su concha juvenil no bicarenada. Entre los Colloniidae, la especie es insólita por el tamaño relativamente grande de la concha adulta, el perfil de la concha juvenil con una quilla y la ausencia de un repliegue secun- dario sobre el diente raquídeo. Se considera que existe dimorfismo sexual en esta especie, pues la concha de la hembra tiene un reborde periumbilical elevado comparable con el de otros trocoideos modificados para incubar en una cavidad umbilical ampliada. KEY WORDS: Trochoidea, Morocco, brooding, endemism. PALABRAS CLAVE: Trochoidea, Marruecos, incubación, endemismo. INTRODUCTION The poorly known genus Anadema been regarded as a trochid, a liotiid, a H. and A. Adams, 1854 has at times colloniid, and a turbinid. Here we * Natural History Museum of Los Angeles County, California 90007, USA. ** Departamento de Biología Animal, Facultad de Ciencias, Universidad de Málaga, E-29071 Málaga, Spain. 53 Iberus , 26 (1), 2008 review the taxonomy and update what is now known about the type and only known species of this genus. Anadema was proposed for the com- bination Omphalius (Anadema) caelata A. Adams, 1854, an Atlantic Moroccan species then thought to be a trochid in the absence of data on a calcified oper- culum. The species was not to be illus- trated until 19 years later but it was described in sufficient detail that its identity has never been in question. In his remarks that followed the Latin description of the species, Adams (1855: 39) wrote: "The character of this shell is so peculiar, on account of the internal spiral callus of the umbilicus, and the absence of characters which constitute allied forms, that I propose to consider it a subgenus of Omphalius , under the ñame of Anadema." The generic ñame Omphalius Philippi, 1847 now pertains to the Tegulinae, as defined by Hickman and McLean (1990). Ten years after the introduction of Anadema , Mórch (1864: 46) reported that a living specimen had been col- lected at the type locality. Morch stated that: "Omphalius (Anadema) caelata is provided with a calcareous operculum, which proves that this species may be removed to Turbo. As there is already a T. caelata, L., I propose for this species Turbo macandrewii." Morch's placement of the species in Turbo predated most work on the genera of turbiniform gas- tropods, and was based on the assump- tion that any turbiniform species with a calcareous operculum could only be a Turbo. This was unfair to Arthur Adams, but the ICZN rules about secondary homonymy forcé us to abandon the original ñame and to use the replace- ment ñame. This ñame honors the re- discoverer Robert McAndrew, who dredged a living specimen at Mogador (now Essaouira), the type locality of the species. The first shell figure known to us was provided still later by P. Fischer (1873), who used the replacement ñame Turbo macandrewii; this is a drawing that has been copied by many subsequent authors. The operculum has never been figured and the repository of the oper- culate specimen examined by Morch is unknown. Pilsbry (1888) ignored the replace- ment ñame and called it Leptothyra caelata, a not unreasonable choice, as it hardly resembles a species of Turbo. Lep- tothyra is now assigned to Colloniidae. Wenz (1938: 340) recognized the genus Anadema and placed it in the turbinid subfamily Liotiinae, under a broad definition in which he also included genera with calcareous oper- cula related to Homalopoma. Keen (1960: 270) placed Anadema in the more restricted Homalopomatinae, a group with fully calcified operculum, now subsumed under the turbinid sub- family Colloniinae in the classification of Hickman and McLean (1990). Nordseick (1968: 33) overlooked Mórch (1864) and placed it in Liotiinae: "Deckel spiralig und mit Kalkbesatz, nicht verdickt", apparently having assumed that it must have the opercular definition of the now understood Lioti- idae, in which the operculum is multi- spiral with calcareous beads on the outer surface. Resolution of the uncertainty is here provided by a preserved immature specimen with operculum, which was collected by the second author in 1991 at Essaouira (formerly Mogador), the type locality. We therefore take this opportu- nity to illustrate the species and confirm its placement. MATERIALS This report is based on material col- lected by the second author, which is now in the malacology collection of the Muséum National d'Histoire Naturelle, Paris (MNHN). The external features of the single live-collected specimen were drawn with the animal fully extended. Subsequent preservation of the speci- men resulted in retraction within the shell. It was later critical-point dried for SEM examination, after which it was rehydrated for extraction of the radula for SEM analysis. 54 McLean AND GOFAS: Notes on the genus Anadema H. and A. Adams, 1854 SYSTEM ATICS Superfamily Trochoidea Rafinesque, 1815 Hickman and McLean (1990) divided the family Turbinidae into a number of subfamilies, including Lioti- inae, Colloniinae, and Turbininae. However, Williams and Ozawa (2007) have reported that their data toward a molecular phylogeny of the family Turbinidae indicates that there are two well-supported groups within the previ- ously defined Turbinidae, which there- fore preeludes the placement of all of the subfamilies in the same family. In view of the ongoing reconsideration of the relationships among these groups. we here treat the Liotiidae, Colloniidae, and Turbinidae at the family level within the superfamily Trochoidea, which simply raises the ranking in the existing classification. This has already been adopted by McLean and Kiel (2007). Additionally, and without dis- cussion, Warén and Bouchet in Bouchet and Rocroi (2005: 245) have separated Turbinoidea and Trochoidea at the superfamily level, which also indicates that the higher classification for trochi- form vetigastropods is currently unset- tled. Family Colloniidae Cossmann, 1916 Subfamily Colloniinae Cossmann, 1916 Hickman and McLean (1990) pro- vided an extensive treatment of the Col- loniidae (then as Colloniinae), distin- guishing them from Turbinidae (then as Turbininae) on their smaller size, non-bi- carinate juvenile shell, lack of cephalic lappets, symmetrical tooth rows and inner lateral teeth that are not greatly enlarged. This is in contrast with the Turbinidae, characterized by larger size, bicarinate ju- venile shell, asymmetrical tooth rows and enlarged inner lateral teeth of the radula. McLean and Kiel (2007) distin- guished two subfamilies within the Col- loniidae, based on opercular morphol- ogy: the basal and mostly extinct Petropomatinae Cox in Knight et al., 1960, having a calcified operculum that is conical on the inner surface, with a fully multispiral pattern, and the Col- loniinae, in which the operculum is fíat on the inner surface with a multispiral pattern that changes to broadly pau- cispiral on the final volution. There is one living genus ( Liotipoma McLean and Kiel, 2007) of Petropomatinae, whereas there are numerous living and fossil genera of Colloniinae. Genus Anadema H. and A. Adams, 1854 Anadema H. and A. Adams, 1854: 430 [as subgenus of Omphalius Philippi, 1847]. Type species (monotypy): Omphalius ( Anadema ) caelata A. Adams in H. and A. Adams, 1854 [= Turbo macan- drewii Morch, 1864; not Turbo caelata Linnaeus, 1758]. Anadema macandrewii (Morch, 1864) (Figs. 1-23) Omphalius (Anadema) caelata A. Adams, in H and A. Adams, 1854: 430 [as type of new subgenus]. A. Adams, 1855: 39 [more detailed description]. Turbo macandrewii Morch, 1864: 46 [new ñame for secondary homonym Omphalius (Anadema) caelata, not Turbo caelata Linnaeus, 1758]. Turbo macandrewi. Fischer, 1873: 98, pl. 29, fig. 3 [first illustration]. Pasteur-Humbert, 1962: 132 [listed]. Leptothyra coelata [sic]. Pilsbry, 1888: 255, pl. 48, fig. 38 [figure after Fischer], 55 Iberus, 26 (1), 2008 Turbo (Anadema) macandrewi. Pallary, 1920: 63. Anadema coelata [sic]. Wenz, 1938: 340, fig. 795 [figure after Fischer]. Nordesick, 1968: 33, fig. 17.00 Anadema caelata. Keen, 1960: 270 [no figure]. Trew, 1992: 19 [listing of species described by H.& A. Adams]. Material examined: BMNH, 2 syntypes, BMNH 1968183, Mogador (now Essaouira), Atlantic Morocco (height 12.3 mm, diameter 16.6 mm; height 12.1 mm, diameter 17.3 mm). MNHN, Essaouira (formerly Mogador), Atlantic Morocco (31° 31' N, 9o 47' W), 1 live-collected male specimen and several beach-worn shells, leg. Gofas, 23 September 1991. MNHN, El Jadida, Atlantic Morocco (33° 16' N, 8o 29' W), beach worn shells, leg. Gofas, 26 September, 1991. MNHN, Mohammedia (for- merly Fedala), Atlantic Morocco (33° 43' N, 7o 21' W), 10 beach-worn shells, leg. Gofas, 1970-71. Description : Because the genus is monotypic, the description that follows applies both to the genus and species. Shell composed of 5 whorls with a low spire; profile dome-shaped, broader than high, suture not impressed; whorls weakly rounded, periphery of immature shell strongly projecting, forming a keel; periph- ery spinose in early stages, but losing spination at maturity; mature shell with a subangulate base; axial sculpture of fine, raised lamellae; spiral sculpture of low, strongly beaded cords; cords between suture and beaded basal cord increasing from three in juvenile to 6 at maturity; base slightly convex, basal cords of com- parable strength and beading to those of body whorl, increasing from three in juve- nile to 8 at maturity; spiral sculpture of both body whorl and base separated by narro w interspaces; axial lamellae well developed in interspaces but not expressed on surface of nodular beads; umbilicus open in juvenile shell, closed in male shell at basal diameter of 9 mm; remaining open in female shell; umbilicus of mature female shell bordered by raised, unbeaded peripheral cord that partially obstructs final quarter whorl and connects directly at base of aperture; aperture oblique, thickened within, descending slightly on final whorl; interior nacreous, inner wall of aperture of female shell smooth, edge with U-shaped sinus; shell colour brick-red. Shell dimensions. Mature female shell (Figs. 1-3): height 13.0, diameter 17.1 mm; immature male shell (Figs. 4- 6): height 4.9, diameter 9.1 mm; máximum dimensions possible for male shell unknown; immature female shell (Figs. 7-9), height 5.9, diameter 9.1 mm. (Right page) Figures 1-15. Anadema macandrewii (Morch, 1864). 1-3: Mature, beach-worn female shell, from Essaouira, Morocco (MNHN), 3 views, height 13.0 mm, diameter 17.1 mm; 4-6: live- collected, immature male specimen with operculum in place, same locality (MNHN), 3 views, height 4.9 mm, diameter 9.1 mm; 7-9: immature, beach-worn female shell, from Mohammedia, Morocco (MNHN), 3 views, height 5.9 mm, diameter 9.1 mm; 10, 11: juvenile beach-worn shell, from Essaouira, Morocco (MNHN), 2 views, height 2.1 mm, diameter 5.0 mm; 12,13: juvenile shell with protoconch showing in umbilical view, from Essaouira, Morocco (MNHN), 2 views, diameter 2.7 mm; 14, 15: operculum of specimen shown in Figures 4-6 and 16, exterior and inte- rior views, máximum diameter 2.1 mm. (Página derecha) Figuras 1-15. Anadema macandrewii (Morch, 1864). 1-3: Concha adulta, explayada, de una hembra, de Essaouira, Marruecos (MNHN), 3 vistas, altura 13,0 mm, diámetro 17,1 mm; 4-6: ejemplar macho inmaduro, recolectado vivo, con el opérculo cerrando la concha; misma localidad (MNHN), 3 vistas, altura 4,9 mm, diámetro 9,1 mm; 7-9: concha inmadura, explayada, de una hembra, de Mohammedia, Marruecos (MNHN), 3 vistas, altura 5,9 mm, diámetro 9,1 mm; 10, 11: concha juvenil explayada, de Essaouira, Marruecos (MNHN), 2 vistas, altura 2,1 mm, diámetro 5,0 mm; 12, 13: concha juvenil con la protoconcha visible desde el ombligo, de Essaouira, Marruecos (MNHN), 2 vistas, diámetro 2,7 mm; 14, 15: opérculo del ejemplar representado en las Figuras 4-6 y 16, vistas exterior e interior, diámetro máximo 2,1 mm. 56 McLean AND GOFAS: Notes on the genus Anadema H. and A. Adams, 1854 57 Iberus , 26 (1), 2008 Figure 16. Anadema macandrewii (Morch, 1864). Drawing of living specimen shown in Figures 4-6. Figura 16. Anadema macandrewii (Morch , 1864). Dibujo del animal vivo, del mismo ejemplar de las Figuras 4-6. Juvenile shell (Figs. 10-13) not with raised axial lamellae, of low profile, exposing protoconch and early whorls in basal view; profile not equally bicari- nate, the upper carination spinose and more strongly projecting than lower, non-spinose carination; protoconch and first teleoconch whorl positioned slightly below level of second whorl. Operculum (Figs. 14-15) calcareous, thick, with a concavity at the base where the white outer surface is coloured light reddish-brown. There is a strong ridge and groove of similar width leading to the upper part, which bears strong pus- tules in the región adj acent to the col- umella; outer edge with a narrow groove; inner surface becoming broadly paucispiral in final volution. This oper- culum is chipped or somehow reduced at its inner edge, to the extent that the outline is not oval; the chitinous layer of the inner surface that shows at the lower left of Figure 15 is missing at the upper left. Flead-foot (Figs. 16-19). Head pro- portionally large, provided anteriorly with a snout terminating in a broad. (Right page) Figures 17-23. Anadema macandrewii (Morch, 1864). 17-19. Scanning electrón micrographs of critical-point dried specimen, same as shown in Figure 16. 17: general view of head-foot; 18: close-up of snout; note the absence of cephalic lappets; 19: close-up of right ante- rior bundle of epipodial tentacle. 20-23. Scanning electrón micrographs of the radula, same speci- men as Figure 16. 20: complete radula; 21: close-up of several complete rows; 22: detail of central and lateral teeth; 23: detail of marginal teeth. Abbreviations, ct: cephalic tentacle; et: epipodial tentacles; f: foot (pointing to propodium); sn: snout. (Página derecha) Figuras 17-23. Anadema macandrewii (Morch, 1864). 17-19. Micrografías elec- trónicas de barrido del ejemplar de la Figura 16 deshidratado por punto crítico. 17: vista general de la cabeza y del pie; 18: vista aumentada del hocico; nótese la ausencia de lóbulos cefálicos; 19: vista aumentada de un haz anterior derecho de tentáculos epipodiales. 20-23. Micrografías electrónicas de barrido de la rádula, ejemplar de la Figura 16. 20: radula completa; 21: vista aumentada de algunas filas completas; 22: detalle de los dientes centrales y laterales; 23: detalle de los dientes marginales. Abre- viaturas, ct: tentáculo cefálico; et: tentáculos epipodiales; f: pie (señalado el propodio); sn: hocico. 58 McLean AND GOFAS: Notes on the genus Anadema H. and A. Adams, 1854 59 Iberusy 26 (1), 2008 flattened area surrounding the mouth, laterally with two prominent, smooth bulges containing rather large, black eyes and, next to these on the anterior side, two slender, villose cephalic tenta- cles; cephalic lappets lacking. Neck tobes present, rather symmetrical, undi- vided. Foot rather small, less than half the diametre of the shell when extended; each side of the epipodium provided in its anterior part with a cluster of three tentacles, the foremost on hardly more than a rounded bulge, the next two tapering and villose, hardly one-tenth of the cephalic tenta- cles in size; in the posterior part with two more tentacles similar in size and shape to the latter. Radula (Figs. 20-23) strikingly sym- metrical, with broad rachidian having lateral extensions, and moderately long, tapered overhanging cusps; with four pairs of similar lateral teeth, which are elbowed like the rachidian and extend above the shaft of the next tooth; shaft of fifth lateral tooth not in cióse contact with shaft of fourth lateral tooth; this tooth broad on both sides, with a less prominent cusp than those of the inner lateral teeth or any of the marginal teeth; innermost pair of marginal teeth with short overhanging cusps, followed by four pairs of marginal teeth with longer cusps; outer marginal teeth with longer shafts, overhanging cusps shorter and deeply serrate on sides of the cusps. The live-collected specimen (Figs. 4- 6, 16) reported here (diameter 9.1 mm) has a nearly closed umbilicus and a gen- erally unmodified base, with no indica- tion of the incipient formation of the DISCUSSION Knowledge of the genus Anadema has been slow to develop because the species A. macandrewii lives in the sublit- toral zone on rocky bottoms exposed to strong surf, where there have appar- ently been few efforts at collecting by diving due to the exposure, low visibil- ity, and extensive muddy bottoms off- shore in relatively shallow water. The projecting rim of mature female shells; it is therefore identified as a male speci- men. There are immature beach-worn shells of about 10 mm in diameter (Figs. 7-9), which are considered to be female shells, having the beginning of a project- ing rim that will form the strong peri- umbilical angulation of the large female shell (Fig. 2). It is evident that male shells are much less frequent among the beach-worn shells. Because the male shell with an operculum is larger than any of the comparable beach-worn shells considered to be male shells, and because the lip is immature, there is no indication from the material at hand as to the possible size reached by male shells. It may be that male shells can reach a size similar to that of female shells. If so, they would differ from female shells in having a rounded umbilical wall, rather than the project- ing umbilical rim of female shells. Both male and female shells seal the umbilicus upon attaining a half-grown diameter of about 10 mm. The female shell then proceeds to form a secondary umbilicus for the brooding cavity. The initial sealing of the umbilicus may help to protect the shell from exposure due to erosión of the apical whorls, which is also avoided in most marine gastropods by shell deposition of a plug from within. There is variation in the peripheral spination of immature stages. The spination of the live-collected male specimen (Figs. 4-6) is stronger than that of any of the beach-worn shells of similar size, whether identified as male or female. single live-collected specimen reported here from the intertidal zone is an unusual record for the species. Systematic position: There is now no doubt that Anadema should be assigned to the family Colloniidae, on the basis of shell and opercular characters, charac- ters of the external anatomy, and the radula. The fine lamellar sculpture had 60 McLean AND GOFAS: Notes on the genus Anadema H. and A. Adams, 1854 brought to mind a comparison with Liotiidae, but that is ruled out by the calcareous operculum. There are many possibilities for opercular morphology in the Colloni- idae, just as there are in the Turbinidae (see Vermeij and Williams, 2007). There is no comparative work on the colloniid operculum, but the operculum of Anadema is within the range of possi- ble expression for the family. Immature shells of the Mediterranean turbinid species Bolma rugosa (Linnaeus, 1767) have a resemblance to Anadema macan- drewii in having a somewhat similar operculum and a spinose peripheral car- ination, but mature specimens of Bolma are much larger, have a higher profile and an impressed suture, a closed umbilicus and an expansive columellar callus that forms a columellar shield nearly as broad as the aperture. For a review of Bolma , see Beu and Ponder (1979). Anadema macandrewii differs from all illustrated species of Bolma in its low, dome-shaped profile and open umbili- cus at maturity. The keeled early juvenile shells indi- cate that early sculpture is not evenly bicarinate, which is a defining feature for Turbinidae, according to the restricted definition provided by Hickman and McLean (1990: 55) at the previously recognized subfamily level. Most colloniids have the juvenile shell with even spiral cords, but there are exceptions to that generalization. The keeled early stage brings to mind the recently described Liotipoma McLean and Kiel, 2007, in the basal subfamily Petropomatinae, which also has an early keel. The lack of cephalic lappets is consis- tent with the assignment to Colloniidae, in which lappets are lacking (Hickman and McLean, 1990). This is in contrast to the Turbinidae, in which the lappets are well-developed (Hickman and McLean, 1990), albeit they are small in the Mediterranean Bolma rugosa. The radular morphology of Anadema provides convincing evidence that it is colloniid rather than turbinid because the tooth rows are perfectly symmetri- cal, and the inner margináis are not greatly enlarged. In turbinids, the asym- metrical tooth row causes the tooth alignment to be skewed, because the large inner laterals must altérnate in zipper fashion when the radula is longi- tudinally folded (as detailed by Hickman and McLean, 1990). Anadema is highly unusual within Colloniidae for its large size and its keeled early stage. With its máximum shell diameter of 17 mm, it may well be the largest known colloniid. All of the colloniid radulae illustrated by Hickman and McLean (1990) have a secondary flap that projects above the rachidian tooth; the radula of Anadema is unusual in not having the secondary flap. In addition, the morphology of the fifth lateral of Anadema seems also to be unusual for the family. Larval brooding: Brooding of larvae within the umbilical cavity has been broadly reported among the Trochoidea (Hickman, 1992: 254). In the Trochidae it is known in Margantes vorticiferus, as shown by Lindberg and Doberteen (1981). In Liotiidae, it has been reported for Arene socorroensis by Shasky (1968) and again by Hertz (1998), and for "Munditia" subquadrata it has been reported by Burn (1976). In the Colloni- idae, it is known in the recently described genus Liotipoma, as reported by McLean and Kiel (2007). Its occur- rence in Anadema is therefore the second known example. In each of these cases, the umbilical rim of the female shell is raised to increase the volume of the umbilical cavity. Such a modification can be the only explanation for the strongly raised umbilical rim of what we interpret as the female shells of Anadema macandrewii. The size reached by the shells of males remains to be discovered; it is possible that mature male shells are smaller than female shells. In support of that possibility, the operculum illus- trated here shows the expansión of the final volution that is characteristic of a mature operculum. However, this shell is somewhat immature because the final lip is not thickened. All beach-worn 61 Iberus, 26 (1), 2008 male shells in the material on hand seem to be immature. The immature male shells are represented by fewer specimens than the larger shells attrib- uted to, for reasons unknown. ACKNOWLEDGMENTS We thank Philippe Bouchet and Vir- ginie Heros of the MNHN for arranging the loan of the Anadema specimens. Illustrations of shells (from photos by BIBLIOGRAPHY Adams, H., and Adams, A., 1854. Genera of Re- cent Mollusca, vol. 1. London, John Van Voorst, 484 pp. Adams, A., 1855. Further contributions towards the natural history of the Trochi- dae; with a description of a new genus, and of several new species, from the Cumingian Collection. Proceedings of the Zoological Society of London, for 1854: 37-41, pl. 27. Bellon-Humbert, C., 1974 ("1973")- Les Mol- lusques marins testacés du Maroc. Catalogue non critique. Premier supplément. Travaux de l'Institut Scientifique Chérifien, serie Zoologique, 37, 1-144, 23. pls. Beu, A. G., and Ponder, W. F., 1979. A revisión of the species of Bolma Risso, 1826 (Gas- tropoda: Turbinidae). Records ofthe Australian Museum, 32 (1): 1-68. Bouchet, P., and Rocroi, J. P., 2005. Classifi- cation and Nomenclátor of gastropod fami- lies. Malacologia, 47 (1-2): 1-397. Burn, R., 1976. Shell with a built-in nest. Aus- tralian Shell News, 16: 3. Fischer, P., 1873. Genre Turbo. Species General et Iconographie des Coquilles Vivantes [L. C. Kiener, continué par P. Fischer]. 128 pp., pls. 1-42. Hertz, C. M., 1998. Arene socorroensis (Strong, 1934) with nestling nepionic larvae. The Fes- tivus, 30 (5): 65. Hickman, C. S., 1992. Reproduction and de- velopment of trochacean gastropods. The Veliger, 35 (4): 245-272. Hickman, C. S. and McLean, J. H., 1990. Sys- tematic revisión and suprageneric classifi- cation of trochacean gastropods. Natural His- tory Museum ofLos Angeles County, Science Se- ries, 35: 1-169. the first author) were prepared in Photo- shop by Michelle Schwengel (formerly LACM) and píate preparation for the shells was completed by Ángel Valdés (formerly LACM). Critical-point dried preparation of the live-collected speci- men and the SEM views of the head- foot, as well as the SEM illustrations of the radula, were provided by Daniel L. Geiger of the Santa Barbara Museum of Natural FFistory. We thank the reviewers for their suggestions, which led to improvements in the paper. Keen, A. M., 1960. [Recent Archaeogastropoda]. In Knight, J. B., L. R. Cox, A. M. Keen, R. L. Batten, E. L. Yochelson, and R. Robertson. 1960. Systematic descriptions (Archaeogas- tropoda). In Moore, R. C. (Ed.): Treatise on Invertebrate Paleontology, Part I, Mollusca 1, pp. 169-310, Geological Society of America and University of Kansas Press. Lindberg, D. R. and Doberteen, R. A., 1981. Umbilical brood protection and sexual di- morphism in the boreal Pacific trochid gas- tropod, Margantes vorticiferus Dalí. Interna- tional Journal of Invertebrate Reproduction, 3: 347-355. Mórch, O. A. L., 1864. Notes on shells. Amer- ican Journal of Conchology , 4: 46. McLean, J. H. and Kiel, S., 2007. Cretaceous and living Colloniidae of the redefined sub- family Petropomatinae, with two new gen- era and one new species, with notes on op- ercular evolution in turbinoideans, and the fossil record of Liotiidae (Vetigastropoda: Turbinoidea). Palaontologische Zeitschrift, 81 (3): 254-266. Nordsieck, F., 1968. Die europaischen Meeres- Gehduseschnecken (Prosobranchia) von Eismeer bis Kapverden und Mittelmeer. Gustav Fischer Verlag, Stuttgart, 273 pp. Pallary, P., 1920. Exploration scientifique du Maroc organisée par la Société de Géographie de París et continuée par la Société des Sciences Na- turelles du Maroc. Deuxiéme fascicule. Mala- cologie (1912). Larose, Rabat and París, 108 p., 1 pl., 1 map. Pasteur-Humbert, C., 1962. Les mollusques marine testacés du Maroc. Catalogue non critique. I. Les Gasteropodes. Travaux de ITn- stitut Scientifique Chérifien, serie Zoologie, 23: 1-245. 62 McLean AND GOFAS: Notes on the genus Anadema H. and A. Adams, 1854 Pilsbry, H. A., 1888. Phasianellinae, Turbinidae, Delphinulinae. Manual of Conchology, vol. 10, 323 pp., 69 pls. [Neritidae, Adeorbiidae, Cy- clostrematidae, Liotiidae in this vol. by George W. Tryon, Jr.] Shasky, D., 1969. Observations on Rósenla nido- rum (Pilsbry) and Arene socorroensis (Strong). The American Malacological Union, Annual Report for 1967: 74. Trew, A., 1992. Henry and Arthur Adams's new molluscan ñames. National Museum of Wales, Cardiff, 63 pp. Vermeij, G. J., and Williams, S. T., 2007. Pre- dation and the geography of opercular thick- ness in turbinid gastropods. Journal of Mol- luscan Studies, 73: 67-73. Wenz, W., 1938. Gastropoda. Handbuch der Palao- zoologie, vol. 6. Teil 1: Allgemeiner Teil und Prosobranchia, Berlin, 1639 pp. Williams, S. T. and Ozawa, T., 2006. Molecu- lar phylogeny suggests polyphyly of both the turban shells (family Turbinidae) and the superfamily Trochoidea (Mollusca: Veti- gastropoda). Molecular Phylogenetics and Evo- lution, 39: 33-51. 63 Iberus, 26 (1): 65-68, 2008 © Sociedad Española de Malacología A new species of Mangelia (Turridae: Mangeliinae) from the Mediterranean Sea Una nueva especie de Mangelia (Turridae: Mangeliinae) del Mediterráneo Charles CACHIA* and Constantine MIFSUD** Recibido el 17-VII-2007. Aceptado el 12-III-2008 ABSTRACT A new species Mangelia melitensis nov. sp., from Malta, is introduced and compared with other similar congeneric species. RESUMEN Se describe una nueva especie Mangelia melitensis, de Malta, y se compara con especies congenéricas de aspecto similar. KEY WORDS: Mollusca, Conoidea, Turridae, Mangeliinae, Mangelia nov. sp., Mediterranean. PALABRAS CLAVE: Mollusca, Conoidea, Turridae, Mangeliinae, Mangelia nov. sp., Mediterráneo. INTRODUCTION The genus Mangelia is represented in the Mediterranean Sea by about 28 species (CLEMAM 2007). The shells are slender, fusiform in shape, with a sculp- ture consisting of axial, and often spiral sculpture. The siphonal canal is short and open and the columella is smooth without any processes. The mollusc has a wide foot, long tentacles with eyes sit- uated on the sides of the stalks and at about 2/ 3 their height, a long siphon and the body is mostly of a transparent white colouration, but sometimes with opaque coloured streaks or blotches, especially on the siphon. The radula consists of hollow marginal teeth, rarely barbed, with an irregular basal swelling (McLean, 1971). The animal lacks an operculum. The diet consists mainly of * 1, Alley 1, St. Catherine Street, Qormi QRM, Malta ** 5, Triq ir-Rghajja, Rabat RBT 2486, Malta polychaete worms, sometimes larger than the mollusc itself. (C. M. pers. obs.) Certain authors (Bouquoy, Dautzenberg and Dollfus, 1883; Powell, 1966; van Aartsen and Fehr de Wal, 1978) have proposed that the Mediterranean species classification should be separated into two genera, Mangelia (Risso, 1826) type species Man- gelia attenuata (Montagu, 1803) and Mangiliella (B.D.D., 1826) type species Mangiliella multilineolata (Deshayes, 1836). The proposition is based on the type of development of the protoconch. Mangelia has a multi-spiral or plank- totrophic type of protoconch, usually consisting of 2 l/i or more whorls while Mangiliella has a paucispiral or direct type of protoconch consisting of 1 1 /lio 65 Iberus, 26 (1), 2008 2 whorls. Although this arrangement is very convenient for separating the species, Bouchet (1990) advised against such methods for creating genera and subgenera in this already problematic family. The Mediterranean species have been dealt with by several authors (Nordsieck, 1977; van Aartsen and Fehr de Wal, 1978; Sabelli, Giannuzzi- -Savelli and Bedulli, 1990; Cachia, Mifsud and Sammut, 2001; Gruppo Malacologico Livornese and Gruppo Malacologica Romagnolo, 2005). During our ongoing research on the marine mollusca which inhabit the coasts of the Maltese Islands, we have come across many specimens of a dis- tinct Mangelia species which is new to Science. It will be introduced herein. SYSTEMATICS Superfamily Conoidea Vaught, 1989 Family Turridae Swainson, 1840 Subfamily Mangeliinae Vaught, 1989 Genus Mangelia Risso, 1826 Mangelia melitensis nov. sp. (Figs. 1, 2) Material studied: All studied material is from the Maltese Islands. Holotvpe. collection number BMNH 20070312 (H= 6mm, W= 2.5mm), and two paratypes, collection number BMNH 20070313, off Gnejna Bay, in sand and algae from 40 m., x-2006, The Natural History Museum, London. Three paratypes, off Ras il-Wahx, 35 m., ix-1992, collection number MNHN 9988, Muséum National d'Histoire Naturelle, París. Three paratypes, off Rdum id-Delli, 50 m., ix-2006, collection number. TAU MO 57001, National Collections of Natural History, Department of Zoology, Tel Aviv University. Four paratypes, off Ras il-Wahx, 25-30 m., ix-1992, and three paratypes, St Thomas Bay, 3-4 m., viii-1987, National Museum of Natural History (Mdina, Malta). Other Paratypes: Salina Bay, 4 m viii-2000, 21 shells; off Rdum id-Delli, 60 m, viii-2006, 5 shells; off Cirkewwa Point, 34 m, viii-1996, 14 shells, in prívate collection of Charles Cachia. [Coll. No 1185]. Off St. Paul's Bay, 40-50 m., ix-1986, 3 shells; Qammieh, Cumnija, 50 m., viii-1986, 4 shells; St. Thomas Bay, 3-4 m., viii-1987, 11 shells; Bahar ic-Caghaq, beached, i-1991, 5 shells; Gozo Island, Dwejra, Coral cave, in sand, 25 m., 1 shell, 9-vi-2001 (leg. Anthony Sammut); off Golden Bay, 40 m, iii-1991, 9 shells; off Ras il-Wahx, 25-30 m., ix-1992, 31 shells off Rdum id- Delli, 50 m., ix-2006, 9 shells, in prívate collection of Constantine Mifsud [coll. No. M1601]. Off Ras il-Wahx, 25-30 m., ix-1992, 2 shells, in prívate collection of Charles Sammut. Salina Bay, 4 m., viii-1998, 2 shells (C. Cachia leg.), in prívate collection of Gert Lindner. Type locality: Gnejna Bay, Maltese Islands. Etymology: Named after the type locality. Description : Shell small, rather fragüe, fusiform, typical of the genus. Proto- conch consists of 2 1 ¡i, rounded, trans- parent white, glassy whorls, the last with a few small, close-set axial riblets crossed by spiral micro-striae. Teleoconch con- sists of up to four whorls which are sub- angulated at their adapical third and curving regularly towards the suture. Sculpture of 8-9 narrow, flexuous, slight- ly opisthocline, axial ribs which continué to the base. Their interspaces are about twice as wide. The whole surface of the shell is covered with dense micro-spiral striae of about the same thickness and close-set axial growth lines, producing an iridescent pattern. There are about 45- 50 of these micro-spiral striae at the penultimate whorl. There are also 5-6 well spaced, thicker spiral chords at the base of the shell, near the siphonal canal. Aperture about half height of shell. Out- er lip sharp, thin and fragüe, thickened internally, with its topmost part clearly curved. Lip varix present in some speci- mens. Sinus shallow. Columella smooth. 66 Cachia AND MlFSUD: A new species of Mangelia from the Mediterranean Sea Figure 1. Mangelia melitensis nov. sp., shells. A: holotype from off Gnejna Bay, in sand and algae, 40 m (height 6 mm); B: paratype from the type locality (height 6 mm); C, D: paratype from off Ras il-Wahx, 25-30 m (height 6.1 mm). Figura 1. Mangelia melitensis nov. sp., conchas. A: holotipo, frente a Gnejna Bay, en arena y algas, 40 m (altura 6 mm); B: paratipo de la localidad tipo (altura 6 mm); C, D: paratipo, frente a Ras il—Wahx, 25-30 m (altura 6,1 mm). Siphonal canal short and wide. The shell colour is white or beige. Rarely, some specimens are decorated with four, faint, brownish spiral bands or with tiny, brown sub-sutural blotches on the penultimate whorl, while others have a dark blotch at the middle of the exterior of the outer lip. A single specimen is completely brown. DISCUSSION M. melitensis nov. sp. had already been introduced as Mangelia sp. A. in Cachia et al. (2001). It differs from all the other Mediterranean forms of the M. unifasciata (Deshayes, 1835) complex, to which it is very similar, by the lack of bold spiral cords on its whorls. It is similar in this respect to M. costulata Risso, 1826 [= M. smithii (Forbes, 1840)], but in that species the spiral striae are fewer, flatter, much bolder, and continuous, not crossed by growth lines, while their interspaces are deeply incised. Mangelia costulata never forms a thickened rib on the The animal is transparent white with large black eyes at about two-thirds of the height of the tentacle stems. The foot has opaque white spots, while the siphon has small opaque yellowish spots. There is no operculum. The species is found near or in Posi- donia oceánica (Linnaeus) Delile, substra- tum at depths of 3 to 50 metres. aperture, unlike the present species: Moreover, in M. costulata the last proto- conch whorl has a bold reticulate type of sculpture, whilst in M. melitensis nov. sp. this whorl has only close-set flex- uous axials with spiral micro-striae. M. costulata usually has a continuous wide brown band on the ultimate whorl and the animal has a bright orange spotted siphon. Mangelia melitensis nov. sp., also offers similarities to Mangelia costata (Donovan, 1804). In that species however the whorls are very evenly convex not somewhat subangulated as 67 ikrus, 26 (1), 2008 Figure 2. Mangelia melitensis nov. sp., scanning electrón micrographs, same paratype as Figures 1C-D. A: protoconch; B: detail of the sculpture on a spire whorl. Figure 2. Mangelia melitensis nov. sp., vistas en microscopía electrónica, mismo paratipo que las Figura 1 C-D. A: protoconcha; B: detalle de la escultura en una vuelta de la espira. in Mangelia melitensis nov. sp. The shell is also smaller, rather glossy, differently coloured, of a more fusiform aspect and with a slightly different protoconch. The animal is completely white. Distribution: The Maltese Islands. The authors have not come across any specimens of M. melitensis nov. sp. from any other Mediterranean locality. BIBLIOGRAPHY Buquoy, E., Dautzenberg, P. and Dollfus, G., 1882-1886. Les Mollusques marins du Rous- sillon. I: Gastropodes. París, Bailliére, 570 pp. Bouchet, P., 1990. Turrid genera and mode of development: The use and abuse of proto- conch morphology. Malacologia, 32 (1): 69-77. Cachia, C., Mifsud, C. and Sammut, P., 2001. The Marine Mollusca of the Maltese Islands, Part 3 Neogastropoda, Backhuys Publishers, 266 pp.. Leiden. CLEMAM, Unitas Malacologica; Checklist of European Marine Mollusca: Web Page: http: / / www.somali.asso.fr /clemam/bio- taxis.php [accessed 5-vi-2007] Gruppo Malacologico Livornese and Gruppo Malacologico Romagnolo, 2005. Nota sulle Mangelia Mediterranee. Notiziario S.I.M., 23 (9-12): 22-33. MacLean, J. H., 1971. A revised classification of the Family Turridae, with the proposal of new Subfamilies, genera and subgenera from the eastem Pacific. The Veliger, 14 (1): 114-130. ACKNOWLEDGEMENTS We would like to thank our colleague P. Sammut (Rabat, Malta) for revising the manuscript and G. Lindner (Germany) for his kind disposition. The scanning elec- trón micrographs for this paper were pro- duced at the University of Málaga with the help of Gregorio Martín Caballero. Mifsud, C., 1998. The molluscan species living in Maltese Posidonia meadows. La Conchiglia, 30 (287): 37-48, 61. Nordsieck, F., 1977. Turridae of the European Seas. "La Pirámide" for La Conchiglia, 131 pp. Rome. Nordsieck, F., 1982. Die europaischen Meeres- Gehauseschnecken. 2nd edition. 539 pp. Gus- tav Fischer, Stuttgart Powell, A. W. B. 1966. The Molluscan families Speightiidae and Turridae. Bulletin of the Auckland Institute and Museum, 5, 184 pp. + 23 plates. New Zealand. Sabelli, B., Gianuzzi-Savelli, R. and Bedulli, D., 1990. Catalogo annotato dei molluschi marini del Mediterráneo, S. I. M. Librería Nat- uralística Bolognese, Vol. 1. xiv, 1-348. Bologna. van Aarsten, J. J. and Fehr de Wahl, M.C., 1978. The sub family Mangeliinae Fischer, 1887 in the Mediterranean. Conchiglie, 14 (3- 6): 97-110. 68 © Sociedad Española de Malacologí Iberas , 26 (1): 69-80, 2008 Catálogo de los bivalvos marinos del sector central del Golfo de Valencia (España) Checklist of the marine bivalves in the central sector of the Gulf of Valencia (Spain) Jorge C. TAMAYO GOYA* Recibido el 12-11-2007. Aceptado el 31-III-2008 RESUMEN Se presenta una lista con 1 84 especies de Bivalvos marinos citados en la literatura y rea> lectados por el autor entre 1998 y 2008, en la costa del Golfo de Valencia, sector cen- tral: 39° 42' N, 0o 12' W (Playa de Canet de Berenguer, Sagunto) - 38° 55' N, 0o 03# W (Playa de Oliva), Mediterráneo occidental. Cari fervensis se cita por primera vez en ¡a zona y se confirma la existencia de poblaciones establecidas de Eastonia rugosa, Fulvia fragilis y Lentidium mediterraneum. ABSTRACT A list is presented with 1 84 species of marine Bivalves cited in the literature or collected by the author between 1 998 and 2007, along the coast of Gulf of Valencia, central sec- tor: 39° 42' N, 0o 12' W (Playa de Canet de Berenguer, Sagunto) - 38° 55' N, 0o 03' W (Playa de Oliva), Western Mediterranean. Gari fervensis is cited for the first time in the area and the occurrence of established populations is confirmed for Eastonia rugosa, Ful- via fragilis and Lentidium mediterraneum. PALABRAS CLAVE: Catálogo, moluscos, bivalvos, Golfo de Valencia, Mar Mediterráneo. KEY WORDS: Checklist, molluscs, bivalves, Gulf of Valencia, Mediterranean Sea. INTRODUCCIÓN En este trabajo se presenta una actua- lización de la lista faunística de moluscos bivalvos de la Bahía de Valencia. Estos complementan un catálogo reciente- mente publicado (Oliver Baldoví, 2007) donde se aporta una actualización de Gasterópodos marinos testáceos de la zona sur del Golfo de Valencia. La gran mayoría de las conchas reco- lectadas son explayadas, por lo que no se puede indicar su hábitat originario. En mucha menor medida, se han obtenido buceando a pulmón-con tubo (entre -2 y -3 m) o recogidas por una embarcación "marisquera" faenando a escasos metros de la costa (zona infralitoral). Las recogi- das de muestras se han ido realizando a lo largo de todas las estaciones entre los años 1998 y 2008 y no se ha seguido una metodología concreta. Los sustratos sedimentarios que constituyen el hábitat de las comunida- des presentes en el piso infralitoral son, las correspondientes a: arenas finas * C/Polo y Peyrolón, 37, izda 16, 46201, Valencia. 69 Iberus , 26 (1), 2008 Figura 1 . Localidades de muestreo en el Golfo de Valencia. Figure 1. Sampling sites in Gulf of Valencia. superficiales, arenas finas bien calibra- das (entre -2/ -3 m. y -20/ -25 m.), arenas finas fangosas (entre -20/-25 m. y -35/- 40 m.), arenas con proporciones peque- ñas de finos (<15%) (consideradas como transición entre ambos sustratos) y los fondos detríticos. El sustrato rocoso está presente en mucha menor medida, lo mismo que las praderas de Posidonia oceánica (en mucha mayor proporción de mata muerta que degradada, y que suele extenderse en una distribución batimé- trica entre -5 m y -20 m), así como la Cymodocea nodosa y la Caulerpa prolifera. Las arenas gruesas y gravillas, ocupan franjas a profundidades variables y sujetas a corrientes marinas. A continuación en profundidad, en la plataforma continental, está el piso circalitoral (entre -40 m. y -100 m.), de escasa pendiente, en el que están pre- sentes los fondos detríticos costeros. Los fondos fangosos terrígenos (a partir de - 100/-150 m.) se extienden por zonas muy amplias del fondo marino. El litoral en estudio, eminentemente sedimentario, se caracteriza por la suce- sión casi continuada de playas de arena fina, sólo interrumpidas por las instala- ciones portuarias, las desembocaduras fluviales (áreas de fondos fangosos) y el Cabo de Cullera (acantilado mediano) Las poblaciones en cuyas playas se han efectuado la recogida de ejemplares son colindantes entre sí y han sido agru- padas, de norte a sur, en las siguientes subzonas (Fig. 1), siendo la B y la D en donde mayor número de muéstreos se han realizado: A: (Canet d'En Berenguer), Sagunto, Puzol y El Puig. Predominan las arenas finas bien calibradas, en mucha menor medida las de arenas finas fangosas y las extensiones de mata muerta de Posi- donia. El puerto de Sagunto y el puerto deportivo de Canet de Berenguer, la planta siderúrgica de Sagunto, el emisa- rio de Canet de Berenguer, y la desem- bocadura del río Palancia, son agentes determinantes en la configuración sedi- mentaria de esta zona del litoral. 70 Tamayo GOYA: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia B: Puebla de Farnals, Massamagrell, Meliana, Alboraya y las playas al norte de Valencia. Las arenas finas fangosas predominan sobre las arenas finas bien calibradas. Es de mencionar el sustrato rocoso frente al Puerto de Valencia y las extensiones de mata muerta de Posidonia, en mayor superficie que las de la subzona A. Él puerto deportivo de Port sa Playa, el Barranco de Carraixet y el emisario de Vera (al norte del Puerto de Valencia), son los agentes de esta subzona. C: Playas al sur de Valencia. Hay mucho mayor predominio de las arenas finas bien calibradas sobre las arenas finas fangosas. El Puerto de Valencia, la desembocadura del río Turia, el emisa- rio del sur del Puerto y las Golas del Pujol y de El Perellonet (provenientes de La Albufera), son los agentes a destacar. D: Sueca y Cullera . Es la subzona con mayor proporción de arenas finas bien calibradas y con menor de finas fangosas en el área de estudio. A reseñar también, la presencia de mata muerta de Posidonia en el afloramiento rocoso frente a Cullera. Cabe mencionar en esta subzona, la desembocadura del Jucar y las Golas de El Perelló y la del Estany. E: Tavernes de la Valldigna, Xeraco, Xeresa, Gandía, Daimuz, Guardamar, Miramar, Piles y Oliva. Las arenas finas fangosas predominan sobre las finas bien calibradas. Frente a Gandía existe otro enclave rocoso con presencia de Posidonia. Los puertos de Gandía y Oliva, constituyen lo más destacable como agentes determinantes. SISTEMÁTICA En la Tabla I se presenta el listado de especies encontradas en el área de estudio, junto con el número de especí- menes encontrados, sus sectores de dis- tribución y citas bibliográficas. Esta lista ha sido ordenada de acuerdo con la cla- sificación propuesta por CLEMAM. Check List of European Marine Mollusca: chttp:// www.somali- asso.fr/ demarro . DISCUSIÓN Glycymeris violacescens (Lamarck, 1819) Es una especie muy presente en gran parte del litoral de Valencia que con fre- cuencia se muestra con un polimorfismo y policromía muy característicos (Fig. 2), predominando el contorno de la concha menos subcuadrangular, muchas veces más oblicua; el color gris negruzco- azulado, principalmente en la mitad superior y umbo, combinado con los colores tierra (beige y marrón) y con mucha mayor superficie de mancha en el interior de la valva; generalmente, los umbos más separados y con mayor área cardinal; cohabitando en proporciones similares con los ejemplares más comunes-tipo, más frecuentemente divulgados (Fig. 3). Fulviafragilis (Forsskal in Niehbur, 1775) Se confirma (Fig. 4) la presencia en la zona de esta especie, previamente citada en Cullera por Zenetos et al. (2004) a partir de ejemplares recolectados en 1991. Ello indica que esta especie está formando poblaciones perennes en la zona, al igual que sucede en otras partes del Mediterráneo (Crocetta, 2005; Vardala-Theodorou , 1999). Eastonia rugosa (Helbling, 1779) Especie presente ocasionalmente en este sector del Golfo de Valencia, salvo en la subzona B en la que si es habitual hallarla. En la playa de Meliana (subzona B) es donde se concentra la gran mayoría de los individuos de esta especie encon- trados en esta zona del litoral (Fig. 5). Decenas y decenas de conchas de indivi- duos jóvenes y adultos son depositadas todos los años en esta playa. En esta subzona B, también se localiza una impor- tante concentración de ejemplares, aunque en menor medida, de Lutraria magna (da Costa, 1778) y Venericardia antiquata (Linné, 1758). Es de notar que esta especie mani- fiesta una tendencia a ampliar su exten- sión en el Mediterráneo, siendo reciente- mente observada en abundancia en las costas italianas en localidades donde no se conocía anteriormente (La Valle, Vani, Liboni y Smriglio, 2007). 71 Iberus, 26 (1), 2008 Tabla I. Listado de especies encontradas en el área de estudio. Ej: concha recolectada explayada; Ej*: entre 1 y 3 ejemplares recolectados; Ej*: entre 1 y 3 ejemplares recolectados, deteriorados; L: especie presente en la colección de referencia del laboratorio de Biología Marina de la Universidad de Valencia (1982-1989, García Carrascosa, no publicado) o citada en la bibliografía reseñada, sin ejemplar recolectado por el autor. Las letras (ABCDE) corresponden a los sectores de la Figura 1 en los que se han recolectado ejemplares. Las referencias a las citas se indican como sigue: (1) Colección de referencia del laboratorio de la Universidad de Valencia; (2) ApáRICI SEGUER, Rowland, Taylor y García Carrascosa, 1996; (3) Bonnin y Rodríguez Babio, 1990; (4) Costa, García Carrascosa, Monzó, Peris, Stubing y Valero, 1984; (3) Generalitat Valenciana, Conselleria de Agricultura y Medio Ambiente, 1994-1997; (6) Giner Ponce, 1989; (7) Hidalgo, 1917; (8) Montero Agüera, 1971; (9) Ramón Herrero, 1993; (10) ROSELLÓ, 1910; (11) SÁNCHEZ Diana, 1980. Los nombres subrayados corresponden a las especies que están comentadas en este trabajo. Tal? le I. List of the species found in the studied urea. Ej: shell collected on the beach; Ej*: 1 to 3 speci- mens found; Ej*: 1 to 3 specimens found, damaged; L: species found in the reference collection deposited in the Marine Biology laboratory ofthe University of Valencia (1982-1989, García Carrascosa, unpu- blished data) or cited in literature, no specimens collected by the author. Letters (ABCDE) refer to Figure 1 sectors. Literature cited as follows: (1) reference collection deposited in the laboratory ofthe University of Valencia; (2) APARICI SEGUER, R.OWLAND, TAYLOR AND GARCIA CARRASCOSA, 1996; (3) Bonnin y Rodríguez Babio, 1990; (4) Costa, García Carrascosa, Monzó, Peris, Stubing and Valero, 1984; (5) Generalitat Valenciana, Conselleria de Agricultura y Medio Ambiente, 1994-1997; (6) Giner Ponce, 1989; (7) Hidalgo, 1917; (8) Montero Agüera, 1971; (9) Ramón Herrero, 1993; (10) Roselló, 1910; (11) Sánchez Diana, 1980. Underlined ñames are discussed in this paper. Especie Especímenes Localización Referencias Familia SOLEMYIDAE Solemya fogata (Poli, 1791) L 3, 7, 8,10,11 Familia NUCULIDAE Nucula nitidosa Winckworth, 1 930 E¡ BCD 3, 5, 7, 8,10,11 Nucula nucleus (Linné, 1758) E¡ ABD 1,3, 5, 6, 7, 8,10,11 Nucula sulcata Bronn, 1831 L 1,2, 3, 5, 6, 7, 8,11 Familia NUCULANIDAE Nuculana pella (Linné, 1767) E¡* ABCDE 2, 3, 5, 6, 7, 8,10,11 Saccella commutata (Phiíippi, 1 844) L 3, 5, 7, 8,10,11 Familia ARCIDAE Arca noae Linné, 1758 E¡ ABCDE 2, 3, 5, 6, 7, 8,10,11 Arca Terragona Poli, 1795 L U, 6, 7, 8,10,11 Barbada barbota (Linné, 1 7 58) E¡ ABCDE 3, 8,10,11 Barbada clatbrata (Defrance, 1816) L 3, Anadara corbuloides (Monterosato, 1 880) E¡* B 3, 7, 8,10,11 Anadara polil (Mayer, 1 868) E¡* B 3, 7, 8,10,11 Bathyarca pectunculoides (Scacchi, 1 835) L 3, 7, 8,11 Bathyarca phillppiana (Nyst, 1 848) L 3,8 Familia NOETIIDAE Striarca ladea (Linné, 1758) E¡ ABCDE 1,2, 3, 5, 6, 7, 8,10,11 Familia GLYCYMERIDAE Glycymeris blmaculata (Poli, 1 795) L 3, 5, 7, 8,10,11 Glycymeris glycymeris (Linné, 1758) E¡* D U, 3, 4, 6, 7,10,11 Glycymeris violascescens (Lamarck, 1819) E¡ ABCDE 1,3, 5, 6, 7, 8,10,11 Familia MYTILIDAE Mytilus edulis Linné, 1758 E¡ ABCDE 3, 7, 8,10,11 72 Tamayo Goya: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia Tabla I. Continuación. Table I. Continuation. Especie Especímenes Localización Referencias Mytilus galloprovincialis Lamarck, 1819 E¡ ABCDE 1,2, 3, 4,5 Mytilaster minimus (Poli, 1795) E¡* D 1,3, 7, 8,10,11 Gregariella petagnae (Scacchi, 1 832) E¡ DE 1,2, 3, 8,10,11 Musculus costalatus (Risso, 1 826) E¡ B 1,3, 6, 7, 8,10,11 Musculus subpictus (Cantraire, 1 835) Ej* B 1,2, 3, 8,10,11 Lithophaga lithophago (Linné, 1758) Ei ABCDE 1,3, 4, 5, 6, 7, 8,10,11 Myoforceps aristatus (Dillwyn, 1 81 7) L 3, 7, 8,10,11 Modiolus adriaticus (Lamarck, 1819) E¡ BD 3,5, 7, 8,10,11 Modiolus mortorelli (Hidalgo, 1 878) L 3, 4, 7, 8,10,11 Modiolus barbatus (Linné, 1 7 58) E¡ ABCDE U, 5, 6, 7, 8,10,11 Amygdalum agglutinans (Cantraine, 1 835) El D 5 Modiolula phaseolino (Phillipi, 1 844) L 3, 7, 8,10,11 Familia PINNIDAE Pinna nobilis Linné, 1758 L 3, 4, 7, 8,10,11 Pinna rudis Linné, 1758 L 3, 8,11 Atrina pectinata (Linné, 1 767) E¡ CDE 3, 7,10,11 Familia PTERIIDAE Pieria hirundo (Linné, 1758) Ei C 3, 4, 5, 7, 8,10, 11 Familia PECTINIDAE Pecten ¡acobeus (Linné, 1 7 58) Ei ABDE 3, 4, 5, 7, 8,10,11 Pecten maximus (Linné, 1758) L 3, 7, 8,11 Aequipecten opercularis (Linné, 1758) Ei DE 3, 5, 7, 8,10,11 íissopecten byalinus (Poli, 1 795) Ei ACD 3, 7, 8,10,11 Palliolum incomparable (Risso, 1 826) L 3, 7, 8,10,11 Pseudamussium clavatum (Poli, 1795) L 3, 7, 8,10,11 Hyalopecten similis (Laskey, 1811) L 3, 7, 8,10 Propeamussium fenestratum (Forbes, 1 844) L 3, Chlamys flexuosa (Poli, 1 795) Ei BCD 3, 4, 7, 8,10,11 Cblamys glabra Linné, 1758) Ei BDE 3, 5, 7, 8,10,11 Chlamys pesfelis (Linné, 1 758) Ei BD 3, 7, 8,10,11 Cblamys multistrata (Poli, 1795) Ei BCDE 3, 5, 8,11 Cblamys varia (Linné, 1758) Ei ABCDE 3, 5, 6, 7, 8,10,11 Familia Spondylidae Spondylus gaederopus Linné, 1 7 58 Ei ABCD 3, 5, 7, 8,10,11 Familia ANOMIIDAE Anomia ephippium Linné, 1 7 58 Ei ABCDE 2, 3, 5, 7, 8,10,11 Pododesmus patelliformis (Linné, 1761) L 3 Familia LIMIDAE tima lima (Linné, 1758) Ei ABCD 1,3, 5, 6, 7, 8,10,11 timaria bians (Gmelin, 1791) Ei ABCDE 1,3, 4, 5, 6, 7, 8,10,11 limaría tuberculata Olivi, 1792 Ei BCDE 1,3, 4, 6, 7, 8,10,11 Limea loscombii (Sowerby G.B., 1 824) L 3, 4,5 [¡matulo subauriculata (Montagu, 1808) L 3, 7, 8,10,11 Notolimea crassa (Forbes, 1 844) L 3, Familia OSTREIDAE Ostrea edulis Linné, 1758 Ei ABDE 3, 5, 7, 8,11 Crassostrea gigas (Thunberg, 1793) L 3, 8,11 73 Iberus , 26 (1), 2008 Tabla I. Continuación. Table I. Continuation. Especie Especímenes Localización Referencias Ostreola stentina (Payraudeau, 1826) E¡ ABCDE 1,3,8 Familia LUCINIDAE Ctena decussata (Costa O.G., 1 829) E¡ BCD U, 6, 7,10,11 toripes lacteus [ Linné, 1758) E¡ ABCDE 1,2, 3, 5, 6, 7, 8, 9,10,11 Lucinella dimicata (Linné, 1758) E¡ BD 1,2, 3, 6, 7, 8,10,11 Anodontia fragtts (Philippi, 1 836) L U, 7, 8,10,11 Myrtea spinifera (Montagu, 1 803) L U, 5, 7, 8,10,11 Lucinoma borealis (Linné, 1 767) E¡ DE 3 Familia THYASIRIDAE Thyasira flexuosa (Montagu, 1 803) L 3,11 Familia UNGULINIDAE Diplodonto rotundata (Montagu, 1 803) E¡ DE U, 7, 8,10,11 Familia CHAMIDAE Chama gryphoides Linné, 1 7 58 E¡ ABCDE 1,2, 3, 5, 6, 7, 8,10,11 Pseudochama gryphina (Lamarck, 1819) E¡ ABCD 3, 4, 7, 8,10,11 Familia GALEOMMATIDAE Galeomma Mw/Sowerby GB in Turton, 1 825 L 1,2, 3, 6, 7, 8,10,11 Familia KELLIIDAE Kellia suborbicularis (Montagu, 1 803) L 3, Bornia geoffroyi (Payraudeau, 1 826) L 3, 8,10,11 Bornia sebetia (Costa O.G., 1829) L 3, 7,10 Familia LASAEIDAE Lasaea rubra (Montagu, 1 803) L 3 Semierycina nítida (Turton, 1 822) L 3 Familia M0NTACUTIDAE Montacuta substríata (Montagu, 1 808) L 3 Tellímya ferruginosa (Montagu, 1 808) L 3, 7, 8,10,11 Mysella bidentata (Montagu, 1 803) L 1,3, 7, 8,10,11 Familia CARDITIDAE Cordita calyculata (Linné, 1758) L 3, 8,10,11 Glans acuieata (Poli, 1795) L 3, 8,10,11 Glans trapezia (Linné, 1 767) E¡ ABCD U, 5, 6, 7, 8,10,11 Venericardia antiguata (Linné, 1758) E¡ BD 3, 5, 6, 7, 8,10,11 Familia ASTARTIDAE Astarte fusca (Poli, 1795) L 3, 7, 8,10,11 715» sulcata (da Costa, 1778) L 3, Digitada digitada (Linné, 1758) L 3, 8,10,11 Goodallia triangulaos (Montagu, 1 803) L 3, 7, 8,10,11 Familia CARDIIDAE Acanthocardia acuieata (Linné, 1758) E¡ ABCDE 3, 5, 7, 8,10,11 Acanthocardia deshayesii (Payraudeau, 1 826) L 3, 7, 8,10,11 Acanthocardia echinata (Linné, 1758) E¡ ABCDE 3, 5, 7, 8,10,11 Acanthocardia paucicostata (Sowerby G.B. II, 1841) E¡ ABDE 3, 5, 7, 8,10,11 Acanthocardia spinosa (Solander, 1 786) L 3, 5, 7,10,11 Acanthocardia tuberculata (Linné, 1758) E¡ ABCDE 2, 3, 4, 6, 7, 9,10,11 Parvicardium exiguum (Gmelin, 1791) E¡ DE 1,3, 7, 8,10,11 Parvicardium mínimum (Philippi, 1836) L 1,3, 7, 8,10,11 74 TAMAYO GOYA: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia Tabla I. Continuación. Table I. Continuation. Especie Especímenes Localización Referencias Parvicardium scabrum (Philippi, 1844) L 3, 5, 7, 8,10,11 Popillicardium papillosum (Poli, 1791) E¡ ABCDE 1,2, 3, 5, 6, 7, 8,10,11 taevicardium crassum (Gmelin, 1791) E¡* BD 3, 5,11 Laevicardium oblongum (Gmelin, 1791) E¡ ABDE 3, 4, 5, 7, 8,10,11 Cerastoderma edule (Linné, 1758) El ABCDE 3, 5, 6, 7, 8,11 Cerastoderma glaucum (Poiret, 1 789) E¡ ABCDE 1,3, 5,10 Fulvia froQilis { ForskaL 1775) E¡ DE Familia MACTRIDAE Moctra glauca Born, 1778 Ei* D 3, 7, 8,10,11 Mactra stultorum (Linné, 1758) E¡ ABCDE 1,2, 3, 4, 5, 6, 7, 8, 9,10,11 Spisula subtruncata (da Costa, 1778) E¡ ABCDE 2, 3, 5, 6, 8, 9,10,11 Lutraria angustior Philippi, 1 844 E¡ BD Lutraria lutraria (Linné, 1 7 58) E¡ BCD 3, 8,10,11 Lutraria magna (da Costa, 1 778) E¡ BCD 3, 7, 8,10,11 Eastonia ruaosa (Helblina. 1 779) E¡ BCD 3, 8,11 Familia MESODESMATIDAE Donacilla cornea (Poli, 1791) L 3 Familia SOEENIDAE Solen marginatus Pulteney, 1799 E¡ ABD 3, 4, 7, 8,10,11 Familia PHARIDAE Pharus legumen (Linné, 1 758) E¡ ABCDE 3, 4, 5, 6, 7, 8, 9,10,11 Ensis arcuatus (Jeffreys, 1 865) L 3 Ensis ensis (Linné, 1758) E¡ ABCDE 3, 5, 7, 8,10,11 Ensis minor (Chenu, 1 843) L 3 Ensis siliqua (Linné, 1758) E¡ ABCDE 4, 7, 8,10,11 Phaxas pellucidus (Pennant, 1 777) E 2,5 Familia TELLINIDAE Teliina distorta Poli, 1791 E 1,3, 5, 7, 8,10,11 Tellina donacina Linné, 1 7 58 E¡* B 2, 3, 4, 5, 6, 7, 8,11 Jellina tabula Gmelin, 1791 Ei* BD 1,2, 3, 4, 6, 9,10 Tellina incarnata Linné, 1758 Ei ABCDE 1,2, 3, 5, 6, 7, 8,10,11 Tellina nítida Poli, 1791 Ei ABCDE 1,2, 3, 4, 5, 6, 7, 8, 9,10,11 Tellina planata Linné, 1758 Ei ABCDE 1,2, 3, 4, 5, 7, 8,10,11 Tellina pulchella Lamarck, 1818 Ei ABCDE 1,2, 3, 4, 5, 6, 7, 8, 9,10,11 Tellina serrata Brocchi, 1814 L 3, 5, 7, 8,10,11 Tellina tenuis da Costa, 1778 Ei ABCDE 2, 3, 4, 5, 6, 7, 8, 9,10,11 Arcopagia balaustina (Linné, 1 758) L 3, 5, 6, 7, 8,10,11 Macoma rumana (Costa O.G., 1 829) Ei ABCDE 1,2, 3, 5, 6, 7, 8, 9,10,11 Gastrana fragilis (Linné, 1 7 58) Ei BDE 1,3, 7, 8,10,11 Familia DONACIDAE Donax semistriatus Poli, 1 7 9 5 Ei ABCDE 1,2, 3, 4, 5, 6, 7, 8, 9,10,11 Donax trunculus Linné, 1758 Ei ABCDE 1,2, 3, 4, 5, 6, 7, 8, 9,10,11 Capsella variegatus (Gmelin, 1791) E 1,3,6 Capsella venustus Poli, 1795 Ei* D 1,3, 5,6 Familia PSAMMOBIIDAE Gari costulata (Turton, 1 822) L U,4 Garidepressa (Pennant, 1 777) Ei BD 3, 5, 6, 7, 8,10,11 75 Iberus, 26 (1), 2008 Tabla I. Continuación. Table I. Continuation. Especie Especímenes Localización Referencias Gari fervensis (Gmelin, 1791) E¡ BD - Familia SEMELIDAE Scrobicularia cottardi (Payraudeau, 1 826) L 3, 7, 8,10,11 Scrobicularia plana (da Costa, 1 77 8) L 3, 5, 8,10,11 Familia SEMELIDAE Abra alba (W. Wood, 1 802) E¡ BD 2, 3, 5, 7, 8,10,11 Abra prismática (Montagu, 1 808) E¡* D 1,3, 4, 7, 8,10,11 Abra segmentum (Récluz, 1 843) E¡ BD 3, 7, 8,10,11 Abra tenuis (Montagu, i 803) E¡ B 2, 3, 7, 8,10,11 Familia SOLECURTIDAE Solecurtus scopula (Turton, 1822) E¡* D 3, 7, 8,10,11 Solecurtus strigilatus (Linné, 1758) E¡ BCD 3, 7, 8,10,11 Azorinus cbamasolen (da Costa, 1 778) E¡ CDE 3, 5,10,11 Familia KELLIELLIDAE Kelliella abyssicola (Forbes, 1 844) L 3 Familia TRAPEZIIDAE Coralliophaga lithophagella (Lamarck, 1819) L 3 Familia GLOSSIDAE Glossus humanus { Linné, 1758) E¡* D 3, 7, 8,10,11 Familia VENERIDAE Venus casino Linné, 1758 E¡ BCD 3, 4, 5, 6, 7, 8,10,11 Venus nux Gmelin, 1791 E¡ BCD 3, 7, 8,10,11 Venus verrucosa Linné, 1758 E¡ ABCDE 3, 4, 5, 6, 7,10,11 Cbamelea gallina (Linné, 1758) E¡ ABCDE 2, 3, 5, 6, 7, 8, 9,10,11 Clausinella fasciata (da Costa, 1778) L 2, 3, 5,11 Timoclea ovata (Pennant, 1 777) L 1,3, 5, 8,10,11 Gouldia mínima (Montagu, 1 803) E¡ BD 13, 5, 6, 7, 8,10,11 Dosinia exoleta (Linné, 1758) E¡ BD 1,3, 4, 5, 6,11 Dosinia lupinus (Linné, 1/58) E¡ ABCDE 1,2, 3, 5, 6, 7, 8, 9,10,11 Pitar ruáis (Poli, 1795) E¡ ABCDE 1,2, 3, 5, 8,10 Callista cbione (Linné, 1758) E¡ ABDE 3, 5, 7, 8,10,11 Ruditapes decussatus (Linné, 1758) E¡ BCDE 1,3, 7, 8,10,11 Tapes rhomboides (Pennant, 1 777) E¡ BD 3, 7, 8,10,11 Irus ¡rus (Linné, 1758) E¡ ABCD 1,3, 7, 8,10,11 Venerupis aurea E¡ ABCDE 1,3, 4, 5, 6, 7, 8, 9,10,11 Venerupis corrugata (Gmelin, 1791) E¡ AD 1, 2, 3, 6, 7, 8,10,11 Familia PETRIC0LIDAE Petricola litbopbaga (Philippson, 1 788) E¡ BCD 1,2, 3, 5, 7, 8,10,11 Lalonkairia lajonkairii (Payraudeau, 1 826) E¡ BCD 3, 5,10 Petricola substriata (Montagu, 1808) L 3,10 Mysia undata (Pennant, 1 777) E¡ DE 2, 3, 8,10,11 Familia C0RBULIDAE Corbula gibba (Olivi, 1/92) E¡* BCD 2, 3, 5, 6, 7, 8, 9,10,11 Lentidium mediterraneum (Costa O.G., 1 829) E¡ BCD 1,2, 3, 4, 6, 8,10,11 Familia GASTROCHAENIDAE Gastrocbaena dubia (Pennant, 1 777) L 1,2, 3, 5, 6, 7, 8,10,11 76 Tamayo Goya: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia Tabla I. Continuación. Table I. Continuation. Especie Especímenes Localización Referencias Familia HIATElllDAE Hiatella árctico (Linné, 1 767) L 1,2, 3, 5, 6, 7, 8,10,11 Hiatella rugosa (Linné, 1 767) L 3,5 Familia PHOLADIDAE Pholas dactylus Linné, 1758 E¡ BCDE 3,10 Bornea candida (Linné, 1758) E¡ BCD 3 Familia THRACIIDAE Thracia papyracea (Poli, 1791) El BCDE 1,2, 3, 6, 7, 8, 9,10,11 Thracia pubescens (Pulteney, 1 799) E¡ BCD 3, 5, 7, 8,10,11 Familia PANDORIDAE Pandora inaequivalvis (Linné, 1758) E¡ ABCDE 1,2, 3, 5, 6, 7, 8, 9,10,11 Pandora pinna (Montagu, 1 803) L 3, 5,11 Familia LYONSIIDAE Lyonsia norwegica (Gmelin, 1791) E¡* D 3,5 Familia POROAAYIDAE Poromya granúlala (Nyst & Westendorp, 1 839) L 3, 8,10,11 Familia CUSPIDARIIDAE Cuspidaria cuspidata (Olivi, 1 792) L 3, 8,10,11 Cuspidaria rostrata (Spengler, 1 793) L 3, 8,10,11 Cardiomya costellata (Deshayes, 1 835) L 3, 8,10,11 Gari fervensis (Gmelin, 1791) Esta epecie tampoco está citada en los trabajos anteriormente relacionados, pero también habita de forma perenne en la Bahía de Valencia (Fig. 6). No es frecuente su presencia en el litoral exa- minado, aunque tampoco lo es de forma ocasional. Se pueden localizar, año tras año, individuos jóvenes y adultos en las mismos emplazamientos y con similar frecuencia y abundancia que su congé- nere, Gari depressa (Pennant, 1777). Lentidium mediterraneum (Costa O.G., 1829) Diversos trabajos escalonados en el tiempo, (recientes: Aparici et al., 1989; García Carrascosa, 1988; y menos reciente. Montero Agüera, 1971) además de los ejemplares localizados por el autor en Puebla de Farnals, Valen- cia, Sueca y Cullera, entre 2004 y 2007 (Fig. 7), evidencian la presencia no de forma esporádica, de esta especie a lo largo de todo el litoral en estudio. AGRADECIMIENTOS Mi más sincera gratitud y reconoci- miento por su generosa y desinteresada colaboración, a Celso Rodríguez Babío y Manuel García Carrascosa, ambos de la Facultad de Ciencias, Universidad de Valencia, por su permanente ánimo, apoyo y por la mucha bibliografía pro- porcionada. Mención especial merece el artículo escrito por Bonin y Rodríguez (1991) que ha sido punto de referencia obligado para la elaboración de este trabajo. A Carmen Salas Casanova y Serge Gofas, ambos de la Facultad de Ciencias, Universidad de Málaga, por su decisiva colaboración, ayuda y correc- ciones, sin las cuales no hubiera sido posible este artículo. A Javier Segarra Julve y colaboradores, Francisco Ruiz Sánchez y Emilio Valero Alfaro, todos ellos de la Generalitat Valenciana, Con- sellería de Agricultura Pesca y Alimen- tación, por su colaboración en propor- cionar los documentos relacionados en 77 Iberus , 26 (1), 2008 Figura 2. Glycymeris violacescens (Lamarck, 1819), ejemplares hallados en Mareny Blau (Sueca). Longitud de la mayor concha encontrada: 67 mm. Figura 3. Glycymeris violacescens (Lamarck, 1819), ejemplares hallados en Cullera. Longitud de la mayor concha encontrada: 64 mm. Figure 2. Glycymeris violacescens ( Lamarck , 1819), specimens collected in Mareny Blau (Sueca). Length ofthe biggest shell found: 67 mm. Figure 3. Glycymeris violacescens (Lamarck, 1819), speci- mens collected in Cullera. Length ofthe biggest shell found : 64 mm. 78 Tamayo GOYA: Catálogo de los bivalvos marinos del sector central del Golfo de Valencia Figura 4. Fulvia fragilis (Forsskal in Niehbur, 1775), ejemplares hallados en Cullera (izquierda) y Oliva (derecha). Longitud de la mayor concha encontrada: 42 mm. Figura 5. Conchas de Eastonia rugosa recolectadas en Meliana. Longitud de la mayor concha encontrada: 80 mm. Figura 6. Gari fervensis , ejemplar hallado en Cullera. Longitud de la mayor concha encontrada: 35 mm. Figura 7. Conchas de Lentidium mediterraneum recolectadas en Mareny Blau (Sueca). Longitud de la mayor concha encontrada por el autor de esta especie: 8 mm. Figure 4. Fulvia fragilis (Forsskal in Niehbur, 1775), specimens collected in Cullera (left) and Oliva ( ñght). Length ofthe biggest shell found: 42 mm. Figure 5. Eastonia rugosa shells collected in Meliana. Length ofthe biggest shell found: 80 mm. Figure 6. Gari fervensis, specimen collected in Cullera. Length ofthe biggest shell found: 35 mm. Figura 7. Lentidium mediterraneum shells collected in Mareny Blau (Sueca). Length ofthe biggest shell found: 8 mm. 79 Iberus, 26 (1), 2008 la bibliografía. A Rafael Araujo Armero, del Museo de Ciencias Naturales de Madrid, Alberto Martínez-Ortí, del Museo de Historia Natural de Valencia; Margarita Belinchón García, del Museo de Ciencias Naturales de Valencia y Francesc Uribe Porta, del Museo de Ciencias Naturales de la Ciutadella (Barcelona), por su colaboración en faci- litar la consulta de ejemplares de las res- pectivas colecciones y, en algún caso, bibliografía. A Isabel Morón Marchante y Mercedes González de Quevedo, de la BIBLIOGRAFÍA Aparici Seguer, V., Rowland, R. A., Taylor, S. y García Carrascosa, M., 1996. Molus- cos infralitorales de la playa del Pinedo-El Sa- ler (Valencia, Mediterráneo occidental). Lbe- rus, 14 (2): 93-100. Bonnin J., Rodríguez Babio C., 1991. Catá- logo provisional de los moluscos bivalvos marinos de la plataforma continental de las costas mediterráneas de la Península Ibérica y de las islas Baleares. Iberus, 9 (1-2): 97-110. Bosca Seytre, A., 1916. Fauna valenciana. Moluscos. Casa Editorial de Alberto Martín, 132 p. Cejalvo Lapeña, M. D., 1998. Peces, crustáceos y moluscos de la Comunidad Valenciana. Con- selleria d'Agricultura, Pesca i Alimentació, 239 p. Costa, M., García Carrascosa, A. M., Monzó F., Peris, J. B., Stubing, G. y Valero, E., 1984. Estado actual de la flora y fauna marinas en el li- toral de la Comunidad Valenciana. Ayunta- miento de Castellón. 239 p. Crocetta, F., 2005. Prime segnalazioni di Ful- via fragilis (Forskal in Niebuhr, 1775) (Mo- llusca: Bivalvia: Cardiidae) per i mari ita- liani. Bollettino Malacologico, 41 (5-8): 23-24. García Carrascosa, M., Silvestre Martínez, R. Y Pérez Yuste, C., 1989. Los Fondos ma- rinos. En Sanchis Molí, E. J. (Ed.): Guía de la Naturaleza de la Comunidad Valenciana. Le- vante - El Mercantil Valenciano, 611-652. Generalitat Valenciana, Conselleria de Agricultura y Medio Ambiente, Direc- ción General de Producción Agraria y Pesca, Servicio de Pesca. 1994-1997. Carac- terísticas bionómicas y cartografiado del bentos en el litoral de Valencia, Castellón y Alicante. Giner Ponce, I. M., 1989. Moluscos y comunidades bentónicas de la costa de Alboraya-Albuixech (Golfo de Valencia, Mediterráneo occidental). Te- sis de Licenciatura. Universitat de Valencia, 225 pp Biblioteca del Museo Nacional de Cien- cias Naturales de Madrid, Sandra Valen- tín Monte y Montserrat Navarro Codina, de la Biblioteca del Museo de Ciencias Naturales de la Ciutadella (Barcelona), por su colaboración en faci- litarme, a pesar de la distancia, la biblio- grafía solicitada. A Francisco Figueres López y a Alfredo His Rocher, ambos de la Cofradía de Pescadores de Cullera por haber permitido el acompañamiento durante el marisqueo en la embarcación MAYEL propiedad de F Figueres. Hidalgo, J. G., 1917. Fauna malacológica de Es- paña, Portugal y las Baleares. Trabajos del Museo Nacional de Ciencias Naturales, serie Zoología, 30: 752 pp. La Valle, P., Vani, D., Liboni, A., y Smriglio, C., 2007. Eastonia rugosa (Helbling, 1799) (Bi- valvia, Mactridae) occurring along the Latium coasts. Basteria, 71(1-3): 93-95. Montero Agüera, I., 1971. Moluscos bivalvos españoles. Anales de la Universidad Hispalense, serie Veterinaria, 5: 1-358 + 95 láminas no nu- meradas. Pla, E., 2006. Guía submarina de La Marina Alta e Ibiza. Publicado por el autor, 350 pp. Ramón Herrero, M., 1993. Estudio de las po- blaciones de Chamelea gallina (Linnaeus, 1758) y Donax trunculus (Linnaeus, 1758) (Mollusco: Bivalvia) en el Golfo de Valencia (Mediterráneo Occidental). Tesis doctoral, Univ. Barcelona: 395pp. Roselló, E., 1910. Los moluscos de Valencia. Asociación Española para el Progreso de las Cien- cias. Congreso de Valencia, Sección 4 (Ciencias Naturales ): 1-7. Sánchez Diana, A., 1980. Los moluscos marinos del Reino de Valencia. Instituto de Estudios Alicantnos. Diputación Provincial de Ali- cante. 154 p. Vardala-Theodorou, G. E., 1999. The occur- rence of the Indo-Pacific molluscan species Fulvia fragilis (Forsskal, 1775) and Bulla am- pulla L., 1758 in Elefsis Bay. Newslletter ofthe Hellenic Zoological Society, 31: 10-11. Zenetos, A., Gofas, S., Russo, G. y Templado, ]., 2004. CIESM Atlas ofexotic species in the Me- diten anean. Vol. 3: Molluscs (F. Briand, ed.). Monaco: CIESM, 376 pp.. 80 Iberus, 26 (1): 81-170, 2008 © Sociedad Española de Malacología New data on the Caribbean Triphoridae (Caenogastropoda, Triphoroidea) with the description of 26 new species Nuevos datos sobre los Triphoridae ( Caenogastropoda , Triphoroidea) del Caribe , con la descripción de 26 nuevas especies Emilio ROLÁN* and Raúl FERNÁNDEZ-GARCÉS** Recibido el 24-IX-2007. Aceptado el 31-III-2008 ABSTRACT In the present work 68 taxa of the family Triphoridae from the Caribbean and adjacent areas are studied. Comments are also provided on one invalid taxon and on another that has been erroneously reported from this area. Among these, 30 were previously known species and 26 are described as new ( Metaxia propinqua , M. propria, M. prompto, Isot- riphora guanahacabibes, Eutriphoro ouffenbergi , Marshallora ostenta , Marshallora apex- diversus, Inella pseudolongissima , I. noduloides, I. apexbilirata , I. harryleei, I. undeber- muda, I. pseudotorticula, I. differens, I. slapcinskyi , I. faberi , Sagenotriphora candidula, "T." inaudita, "T." pseudonovem, " T. " yociusi, " T. " guadaloupensis, "T." amicorum, "T." turtlebayensis, "T." grenadensis, "T." abacoensis a nd "T." portoricensis). Twelve species remain unnamed or are tentatively identified, due to scarcity or ¡nadequacy of available material, or doubtful specific assignment. New information is reported for Monophorus oli- vaceus (Dalí, 1889) and Sagenotriphora osclausum (Rolán and Fernández-Garcés, 1995) already treated in previous works. Lectotypes are designated and figured for the following species Inella bigemma (Watson, 1880) , Inella ¡nflata (Watson, 1880), Inella longissima (Dalí, 1881), Inella trlserialis (Dalí, 1881) , Inella intermedia (Dalí, 1881), Inella ibex (Dalí, 1881), Inella torticula (Dalí, 1881), Inella colon (Dalí, 1881), Inella compsa (Dalí, 1927), Inella enopla (Dalí, 1927), Inella meteoro (Dalí, 1927), Inella pompona (Dalí, 1927), Inella dinea (Dalí, 1927), Inella sentoma (Dalí, 1927), Inella gaesona (Dalí, 1927), "Triphora" cylindrella (Dalí, 1927), "Triphora" abrupta (Dalí, 1881), "Triphora" caracca Dalí, 1927, "Triphora" georgiana Dalí, 1927, "Triphora" indígena Dalí, 1927, "Triphora" lilacina (Dalí, 1889), "Triphora" pyrrha Henderson and Bartsch, 1914, Triphora" atlántica (E. A. Smith, 1890) and for the western Atlantic species Triphora aspera Jeffreys, 1 885. A list of the ñames employed for Caribbean species of Triphoridae is given, mentioning mistakes and synonymies. RESUMEN En el presente trabajo se estudian 68 taxones de la familia Triphoridae del Caribe y aguas próximas. También se aporta información sobre un taxon que carece de validez y otro erró- neamente mencionado para el área. Entre ellos, 30 corresponden a especies previamente conocidas, 26 se describen como nuevas para la ciencia [Metaxia propinqua, M. propria, M. prompta, Isotriphora guanahacabibes, Eutriphora auffenbergi, Marshallora ostenta , Mars- hallora apexdiversus, Inella pseudolongissima, I. noduloides, I. apexbilirata, I. harryleei, I. * Museo de Historia Natural, Campus Universitario Sur, 15782, Santiago de Compostela, Spain ** Centro de Estudios Ambientales de Cienfuegos (CEAC), Grupo de Gestión Ambiental (GGA), calle 17, esquina Ave. 46, Cienfuegos, Cuba. 81 Iberus , 26 (1), 2008 undebermuda , I. pseudotorticula, I. differens, I. slapcinskyi , I. faberi, Sagenotriphora candi- dula, "T." inaudita, "T." pseudonovem, “T." yociusi, "T." guadaloupensis, "T." amicorum, "T." turtlebayensis, "T." grenadensis, "T" abacoensis y "T." portoricensis) y o 1 2 de ellas no se les asigna un nombre definitivo debido a la escasez del material, o al mal estado del mismo, y también a la existencia de dudas sobre su determinación. Además, se aporta nueva infor- mación sobre Monophorus olivaceus (Dalí, 1 889) y Sagenotriphora osclausum (Rolán y Fer- nández-Garcés, 1 995), especies ya citadas en trabajos previos. Se designan y figuran lectotipos de las siguientes especies: Inella bigemma (Watson, 1 880), Inella inflata (Watson, 1 880), Inella longissima (Dalí, 1881), Inella triserialis (Dalí, 1881), Inella intermedia (Dalí, 1881), Inella ibex (Dalí, 1881), Inella torticula (Dalí, 1881), Inella colon (Dalí, 1881), Inella compsa (Dalí, 1927), Inella enopla (Dalí, 1927), Inella meteoro (Dalí, 1927), Inella pompona (Dalí, 1927), Inella dinea (Dalí, 1927), Inella sentoma (Dalí, 1927), Inella gaesona (Dalí, 1927), "Triphora" cylindrella (Dalí, 1927), "Triphora" abrupta (Dalí, 1881), "Triphora" caracca Dalí, 1927, "Triphora" georgiana Dalí, 1927, "Triphora" indígena Dalí, 1927, "Triphora" ¡Hacina (Dalí, 1889), "Triphora" pyrrha Henderson y Bartsch, 1914 y también de la especie atlántica Triphora aspera Jef- freys, 1885. Al final se aporta una lista de los nombres empleados para las especies de Triphoridae del Caribe, señalando los errores habidos y las sinonimias. KEY WORDS: Triphoridae, Caribbean, new species. PALABRAS CLAVE: Triphoridae, Caribbean, new species. INTRODUCTION Triphoridae J. E. Gray, 1847 is a very diverse family of marine gastropods that occurs world-wide, with about 600 ñames proposed for the Recent species (Marshall, 1983). Triphorids live on rocky substrates from the intertidal fringe to a depth of more than 1000 m. A complete general information on the family Triphoridae is given by Mar- shall (1983) and Wells (1998), who noted that it is a highly species-rich group, most species being sinistral, with a high spire, numerous whorls, a short to long anterior canal and a posterior apertural notch or canal. The teleoconch sculpture of triphorids is notably vari- able, and can be beaded, spinose or reticulate, and the shells are small (usually less than 10 mm, although some can be up to 40 mm or more). Marshall (1983), in his revisión of the Indo-Pacific species, emphasized the importance of the shape and sculpture of the protoconch for taxonomy, and pointed out that "under absolutely no circumstances should further new species be proposed unless a complete, unworn protoconch can be illustrated". The family has its máximum diver- sity in the tropical Indo-Pacific where, according to Marshall (1983), there are more than 1000 species. An impression of the wealth of species in this area is afforded by the 174 species found during an extensive sampling in the coral reef lagoon of Koumac, New Cale- donia (Bouchet et al., 2002), Triphori- dae being the second richest family of molluscs, after Turridae. Although notably less diverse than in the Indo-Pacific, this family contains many species in the Atlantic Ocean, poorly known in most areas. Bouchet and Guillemot (1978) and Bouchet (1985, 1997) revised the species of this family occurring in the Mediterranean and neighbouring Atlantic, where CLEMAM data base recognised 19 species in 10 genera. Some additional species were described by Fernándes and Rolán (1988, 1993) and Van der Linden (1998) in the Cape Verde Islands, 82 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae and Rolán and Peñas (2001) from the Canary Is. and the Mediterranean. Information on triphorids from many Caribbean areas is contained in a number of works, such as those for Jamaica (C. B. Adams, 1850a, 1850b, most species represented in Clench and Turner, 1950), Virgin Islands (Nowell-Usticke, 1959 and 1971), Yucatán Península, México (Vokes and Vokes, 1983), Puerto Rico and nearby Caribbean areas (Warmke and Abbott, 1961), Curasao, Aruba and Bonaire (De Jong and Coomans, 1988), Oceanic Islands off Brazil (Leal, 1991), Colom- bian Caribbean (Díaz Merlano and Puyana Hegedus, 1994), the Atlantic and Gulf coasts and West Indies (Morris, 1973), and Brazil (Ríos, 1994). Rolán and Fernández-Garcés (1993a, 1993b, 1994 and 1995) and Rolán and Espinosa (1994) showed the known species from Cuba and described some new species. Besides, other authors described new species from other Caribbean areas, such as De Jong and Coomans (1988), Moolenbeek and Faber (1989), Faber and Moolenbeek (1991), Rolán and Crúz-Ábrego (1996) and Rolán and Luque (1999). More recently a large new species has been described from Brazil (Simone, 2006). Many other species were recorded from deep water, and described by Watson (1880, 1886), Dall (1881, 1889, 1927) and other authors. Fossil species were mentioned in Olsson and Harbison (1953). Colour photographs of 33 species, including most of the known shallow- water species and those described during the last 20 years in previously mentioned papers, are provided in a recent paper (Rolán and Fernández- Garcés, 2007) together with a list of ñames assigned to the Caribbean Triphoridae. Henee, the present work has the fol- lowing objectives: 1- To give available information on the deeper water species, showing the type material whenever possible; 2- To provide addi- tional colour photographs of the Caribbean species which were not figured in our previous work (Rolán and Fernández-Garcés, 2007); 3- To supply additional information that has recently been obtained on other species already known from shallow water; 4- To describe some new species which have been collected from shallow water; 5- To present an updated list of all the taxa mentioned for the Caribbean and nearby areas, making corrections to our previous list. MATERIAL AND METHODS Some of the species presented here are only known from the type material, loaned by several museums (mentioned in Abbreviations and Acknowledge- ments), or viewed on photographs, pro- vided by their Curators or other person- nel. In the collected material, the collec- tor ñame and the data of collection were referred when both were known. Besides, material loaned by Harry G. Lee from his collection, mainly from deep water, is included in this study. Finally, a few species were collected by the authors and persons who have cooperated with them. Most of the early described species, some of which had never been figured previously or had only been represented by drawings, are photographed here in colour, and when- ever possible these figures inelude the types and also illustrate intraspecific variability. Some photos taken by Sally Diana Kaicher (found in ANSP) were included. The order of presentation is basically a grouping based on similarity. It was also the authors' intention to inelude most of the available information on protoconchs, radulae and opercula, but unfortunately this was only possible with a few species. Most of the type material excluded this possibility, and much of the studied material consisted of shells without soft parts. An attempt has been made also to solve taxonomic problems relevant to some of the studied taxa. None of Dall's descriptions included a designated holotype, although some 83 Iberus , 26 (1), 2008 seem to have been based on a single shell. Under these circumstances, Dall's material placed in some museums must be considered as syntypes. After exami- nation of available material from these museums, it was therefore decided to designate as lectotypes the shells that were closest to the original description or the original figure (shown in Dall, 1889). In the description of shells, the spiral beaded cords are frequently referred to as "spiral 1, 2, 3 etc.", spiral 1 being the subsutural cord on the adapical part of the whorl, with subsequent cords on the teleoconch whorls referred to as spirals 2 and 3, folio wing Marshall (1983). Other important characters in the description are, those of the protoconch, such as the diameter, the shape and size of the nucleus (elevated, depressed, large, small, ...), the number of spiral cords (none, 1 or 2), and the presence or absence of axial ribs. We have followed the method of Verduin (1976) for count- ing the number of whorls of the proto- conch. On the teleoconch, the number and size of the spiral cords is noted on the first and last whorls, where they are frequently different. Also noteworthy is the position of spiral 2 (closer to spiral 1, equidistant between spirals 1 and 3) and its size (nodules smaller, elongate, etc.). It is important to point out that the nodules, which are formed by the inter- section of spirals and axial ribs, are sometimes simply spherical, while other times they are concave above and convex below; in this case they are crossed by a line that represents the spiral cord and appears to cut the nodule: these are referred to as cut nodules or nodules that are cut in the middle. Due to the fact that the radula of most species was unavailable, and as this character is very important for generic assignment, it was necessary in many cases to employ generic ñames very much in a sensu lato as "Triphora" or "Inella". For generic arrangement we mainly follow the works of Laseron (1958), Marshall (1983) and Bouchet (1985). Abbreviations: AMNH: American Museum of Natural History, New York ANSP: Academy of Natural Sciences, Philadelphia BMSM: Bailey-Matthews Shell Museum, Sanibel Is., Florida FLMNH: Florida Museum of Natural History, Gainesville, Florida IES: Instituto de Ecología y Sistemática, La Habana MCZ: Museum of Comparative Zoology, Cambridge MHNS: Museo de Historia Natural, Santiago de Compostela MNCN: Museo Nacional de Ciencias Naturales, Madrid USNM: United States National Museum, Washington CCR: collection of Colin Redfern CHL: collection of Harry G. Lee CFG: collection of Raúl Fernández- Garcés CMK: collection of Mario Krisberg sp: specimen with soft parts s: shell j: juvenile f: fragment RESULTS I. Taxonomic part Family Triphoridae Gray, 1847 Subfamily Metaxiinae Marshall, 1977 Genus Metaxia Monterosato, 1884 Type species: Cerithium rugulosum C. B. Adams, 1850. According to Marshall (1983), this sub- family is formed by dextral triphorids. 84 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Metaxia propinqua spec. nov. Rolán and Fernández-Garcés (Figs. 1C-H) Type material: Holotype (Figs. 1C, ID) in FLMNH (154988). Paratypes: 4 s, Louisiana, 27.98568° N 92.6472° W, 65-91 m (FLMNH UF 291348); 5 s, Hillsborough Co., SW Egmont Key, Florida 73-91 m (collector Steger) (FLMNH 238632); 2 s (Figs. 1E, 1F), Palm Beach Co., Florida, off Boynton Inlet, 84-106 m (M. Glockstein, 1980) (FLMNH 47382). Other material examined: 2 s (with broken protoconch), Louisiana, 27.98568° N, 92.6472° W, 65- 91 m (FLMNH UF 291348); 4 s (protoconch lost or eroded), Hillsborough Co., SW Egmont Key, Florida 73-91 m (Steger) (FLMNH 238632); 1 s (protoconch eroded), 2 miles off Virginia Key, Florida, 9 m (FLMNH UF 365086); 1 s (protoconch eroded) (FLMNH). Type locality: Florida, Monroe Co, WNW Dry Tortugas, 76 m. Etymology: The specific ñame derives from the Latín word propinquus -a -um which means “cióse, similar", alluding to the similarity with other species of this genus. Description: Shell conical, very elon- gate, relatively solid. Protoconch (Figs. 1G, 1H) white, paucispiral, with a little more than two whorls and about 300 pm in diameter; several spiral threads begin at the apex. These are well defined, a little irregular, four on the first whorl decreasing to two on the second, where numerous axial ribs run from the suture to the upper thread. The teleoconch can be white or light brown. Four spiral cords are present from the beginning, with the subsutural upper one less prominent than the other three. Axial ribs are present on the entire shell, about 10-11 on the first whorls and eighteen on the final one. Suture depressed with a very small spiral thread above. On the base, the profile is slightly concave and one additional cord is present. Aperture almost round with an open and short siphonal canal. Dimensions : The holotype measures 6.3 mm. Distribution : Known from Florida and Louisiana, USA. Remarks: In order to facilítate com- parison of the protoconchs of the Metaxia species mentioned here, they have all been presented together in Figure 3. M. rugulosa has a protoconch (Figs. 3A- 3D) with 2 1/i whorls (see Rolán and Redfern, 1996), sometimes reaching three whorls, with a spiral sculpture on the first whorl formed by undulating or zigzag- ging lines; the second protoconch whorl has axial ribs which termínate at a single spiral thread (only exceptionally double); the teleoconch is white. M. excelsa and M. taeniolata have multispiral protoconchs. Metaxia propria spec. nov. Rolán and Fernández-Garcés (Figs. 2A-C) Type material: Holotype (Figs. 2A, 2B) in FLMNH (UF 393603). Type locality: Florida, Key Largo, 228 m. Etymology: The specific ñame derives from the Latín word proprius -a -um which means “special, characteristic", alluding to the differential characters of the protoconch that distinguish it from other species of this genus. Description : Shell conical, very elon- gate, relatively solid. Protoconch white (Fig. 2C) with a little more than 2 3A whorls and about 400 pm in diameter; two well defined spiral cords are crossed by 13-15 axial ribs per whorl which are continuous with those of the subsequent whorl. Teleoconch white. Four spiral cords are present from the beginning, the subsutural upper one being less promi- nent than the other three. Axial ribs present on the entire shell, with about 10- 11 on the first whorls and 13 on the final one. Suture depressed with a very small spiral thread above. On the base, the profile is slightly concave and one addi- tional cord is present. Aperture almost round, with an open siphonal canal. Dimensions : The holotype measures 5.9 mm. 85 Iberus, 26 (1), 2008 Distribution : Only known from Florida. Remarks : Characters of the proto- conch provide the main differences between this species and those previ- ously known: M. rugulosa has a protoconch (Figs. 3A-3D) with 2 l/i whorls, sometimes reaching 3, with a spiral sculpture on the first whorl formed by undulating or zigzagging lines; the second protoconch whorl has axial ribs which terminate at a single spiral thread (only exception- ally double). M. propincua spec. nov. lacks axial ribs on the first whorl, and they reach only to the upper spiral cord on the second whorl. Metaxia prompta spec. nov. Rolán and Fernández-Garcés (Figs. 2D-J) Type material: Holotype (Figs. 2D, 2E) and 2 paratypes (Figs. 2G, 21) (FLMNH UF 359136). Type locality: Bermuda, Hamilton Parish, Shelly Bay. Etymology: The specific ñame derives from the Latin word promptus -a -um which means "evident, available", alluding to the characters of the protoconch that are different from other species of this genus. Description : Shell conical, very elon- gate, relatively solid. Protoconch (Figs. 2F, 2H, 2J) white, paucispiral with about 2 V2 whorls and between 300 and 400 pm in diameter; a single spiral cord is poorly defined on the first whorl and a little stronger on the second. The axial ribs are numerous but not continuous; they descend from the upper suture and almost disappear just before reaching the spiral cord, reappearing and strengthen- ing on the cord before fading again towards the lower suture. Teleoconch white or light brown. Four spiral cords are present from the beginning, the sub- sutural upper one being less prominent than the other three. Axial ribs are present on the entire shell, varying in number on the early whorls and increas- ing to about 18 on the final one. Suture depressed with a very small spiral thread above. On the base, the profile is slightly concave. Aperture almost round with an open siphonal canal. Dimensions : The holotype measures 4.1 mm. Distribution : Only known from Bermuda Archipelago. Remarks : Characters of the proto- conch easily differentiate this species from related ones: M. rugulosa has more rounded whorls with zigzagging spiral lines on the first whorl and numerous axial ribs on the second, which only reach the middle of the whorl. M. espinosai has more angulate, wider whorls, with prominent elongate nodules and is always milky white. M. propincua has rounded whorls, with straight spiral cords beginning at the apex and two cords on the second whorl, with axial ribs absent from the lower half of the whorl. M. propria has almost three whorls, rounded with two spiral cords and with a few widely separated and prominent axial ribs on the first whorls. Metaxia sp. 1 (Figs. 1A-B) Material examined: 1 s. Barbados, 183 m (USNM, labeled "Cerithiopsis abrupta Watson, 1880"). Remarks : The shell from Barbados in USNM (without any type label) has a paucispiral protoconch of 450 pm in diameter, differing from M. abrupta (Watson, 1880), which has a proto- conch that is more depressed and also has axial ribs (figured in Bouchet, 1985). Dimensions : The shell studied mea- sures 5.0 mm. 86 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Bouchet (1985) was unable to confirm the existence of M. abrupta in the western Atlantic. Probably this species is endemic to the Azores Archipelago. The species studied here could be the one referred to by Dall (1889) as Cerithiopsis abruptus (Fig. 36L), but we are not sure. This species is different from the other two Caribbean species with pau- cispiral protoconch: M. espinosai (described by Rolán and Fernández- Garcés, 1993a) has a paucispiral proto- conch (Fig. 3E) with a prominent nodu- lous keeled border; and M. rugulosa (C. B. Adams, 1850), which has a narrower protoconch (Figs. 3A-3D) with a differ- ent sculpture (Rolán and Redfern, 1996). However, it must be pointed out here that the protoconch of the taxon M. rugulosa is not known, as the apex is missing from the holotype of this species, represented in Clench and Turner (1950). Another fragment in BMNH with the label "figured speci- men" had no protoconch. In several works it has been assumed that the protoconchs from Cuba and the Bahamas figured in Rolán and Redfern (1996) represent the protoconchs of M. rugulosa , probably due to the proximity of Cuba to Jamaica, which is the type locality. Similar proto- conchs were figured in Redfern (2001). The other two known species of Caribbean Metaxia ( M . excelsa and M. taeniolata) have multispiral protoconchs. The present species is probably unknown and unnamed, but as only one shell has been examined, and its proto- conch is not in very good condition, it is kept unnamed pending the availability of more material in the future. Subfamily Triphorinae Gray, 1847 Genus Monophorus Granata-Grillo, 1877 Type species (by monotypy): Trochus perversus Linnaeus, 1758. Recent, Mediterranean. Remarks: The main characters (shown in Marshall, 1983, Bouchet, 1985 and Rolán and Fernández-Garcés, 1994) are the following: multispiral protoconch, teleoconch whorls flat-sided or shallowly convex, well defined spiral cords and axial ribs, nodular intersections and a typical radula with central tooth with 3-5 cusps, lateral with 5-6 and numerous margináis with only 2-3 short cusps. Monophorus olivaceus (Dall, 1889) (Figs. 4B-F, 4F4-K) Triforis decorata var. olivácea Dall, 1889. Bull. M. C. Z., 18: 243. [Type locality: Gulf of México, west of Florida, 91 m, Key West, Hemphill]. Triphora ornata auct. non Deshayes (1832). Type material: One syntype, fragment of 10 mm, here designated as lectotype (Figs. 4C, 4D) (MCZ 7379). Other material examined: México: 1 s, Cancun, Quintana Roo (Heilman, March, 1963) (ANSP 285517). Florida: 3 s, WSW of John's Pass, 50 m (Steger, 1956) (ANSP 306254); 1 s, Palm Beach Co., Bath and Tennis (McGinty coll/ Apr. 1951), 36-55 m (FLMNH 176649); 4 s, Collier Co., SW of Naples, 26 20' N (Powlus/Aug., 1960), 55 m (FLMNH 158202); 9 s, Palm Beach Co., off Palm Beach, 100 m (FLMNH 154886); 3 s, Palm Beach, WOE Radio Towers to Bath and Tennis (McGinty, May/1951) 36-55 m (FLMNH 250132); 1 s, Palm Beach Co., (R/V Tritón, Apr. 1950) 40-50 m (FLMNH 204877); 1 s, off Marathón (M. and S. Snyder, Jul. 1966) (ANSP 309744). Bahamas: 1 s, reef NE of North Point, Elbow (Little Guana) Cay, Abaco (Robertson, Aug. 1955) (ANSP 298409); 4 s, Grand Bahama Island, 26° 31' 00" N, 78° 46' 30" W (J.N. Worsfold) (ANSP 373939); 1 s, Grand Bahama Island, 26° 29' 45" N, 78° 37' 15" W (G. Taylor and D. Nilsen) (ANSP 273605); 2 s, Tamarind, Grand Bahama Island 26° 30' 45" N, 78° 36' 00" W (J. Worsfold) (ANSP 368896); 1 s, Bootle Bay, Grand Bahama Island 26° 87 Iberus , 26 (1), 2008 Figure 1. A, B. Metaxia sp. 1. A: shell, 5 mm, Barbados, 183 m (USNM); B: protoconch. C-H. Metaxia propinqua spec. nov. C, D: holotype, 6.3 mm (FLMNH); E, F: paratype, 3.4 mm (FLMNH); G: protoconch of the paratype; H: detail of the protoconch of the holotype. Figura 1. A, B. Metaxia sp. 1. A: concha , 5 mm, Barbados, 183 m (USNM); B: protoconcha. C-H. Metaxia propinqua spec. nov. C, D: holotipo, 6,3 mm (FLMNH); E, F: paratipo, 3,4 mm (FLMNH); G: protoconcha del paratipo; H: detalle de la protoconcha del holotipo. 88 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 2. A-C. Metaxia propria spec. nov. A, B: holotype, 5.9 mm (FLMNH); C: protoconch of the holotype. D-J. Metaxia prompta spec. nov. D, E: holotype, 4.1 mm (FLMNH); F, G: proto- conch and shell of paratype, 3.3 mm (FLMNH); H, I: protoconch and shell of paratype, 4.0 mm (FLMNH); J: protoconch of holotype. Figura 2. A-C. Metaxia propria spec. nov. A, B: holotipo, 5,9 mm (FLMNH); C: protoconcha del holotipo. D-J. Metaxia prompta spec. nov. D, E: holotipo, 4,1 mm (FLMNH); F, G: protoconcha y concha de un paratipo, 3,3 mm (FLMNH); H, I: protoconcha y concha de un paratipo, 4,0 mm (FLMNH); J: protoconcha del holotipo. 89 Iberus, 26 (1), 2008 39' 30" N, 78° 57' 00" W (J. Worsfold) 80 m (ANSP 371846); 1 s, Schooner Cays, NW of Powell Point, Eleuthera 24° 56' 45" N 76° 25' 00" (R. Robertson) (ANSP 363405); 3 s (G. Buchanan) (BMNH 90.11.24.428-34). Venezuela: 4 s, distrito Federal (Berthier, 1935) (ANSP 264204). Grenada: 1 s, mouth of St. George Harbour (Ostheimer, Jan. 1964) 44-66 m (ANSP 297306). Virgin Islands: 3 s, St. Croix (R.E. Griffith) (ANSP 18201). Dutch Antilles: 5 s, (Gray coll.) Saint Vincent (BMNH). No exact local- ity: 3 s (McAndrew coll.) (BMNH Acc n° 1563); 2 s (R. Sykes coll.) (BMNH acc. n° 1825). Description : See Dall (1889). It is important to point out some characters: the large size of the beads, spiral 2 being absent from the early whorls and smaller on most of the shell, the colour forming blotches, predominantly on spirals 1 and 2. Spiral 3 is white, but the cord between the beads is usually dark. Dimensions : The lectotype measures 10 mm. Distribution: Known throughout the Caribbean. Remarks: There is considerable con- fusión regarding this taxon: the species was referred to as Triphora ornata by many authors (Warmke and Abbott, 1961; Morris, 1973; Ríos, 1994) until Moolenbeek and Faber (1991) showed the correct ñame to be M. olivaceus (Dall, 1889). The type material of Triphora ornata was examined by Faber (pers. comm.) in 1983 from MNHN (Ecole des Mines collection where the Deshayes types should have been). This lot con- sists of several poor shells, none of which matches the Caribbean species, and one of them in better condition cor- responds to a Red Sea species. The only known type specimen of Triforis decor ata var. olivácea is an atypical and faded shell. It can be confused with some species with axial colour blotches: Latitriphora albida (Fig. 4A) has very characteristic smaller nodules, and no confusión is possible; Nototriphora deco- rata may be more similar and requires careful comparison. We have compared a typical shell of Nototriphora decorata (Fig. 4G) with representative shells of M. olivaceus (Figs. 4D-F) and with the lectotype of the latter species (Figs. 4B, C). It can be seen that the early whorls of N. decorata are narrower than those of the type of M. olivaceus and have smaller nodules, whereas the early whorls of representative M. olivaceus are very similar to those of the type specimen. As it is necessary for nomenclatural stabil- ity to designate a lectotype, we propose as such the only known syntype. The species M. olivaceus is very vari- able, with large, wide shells sometimes found in the same population as small, narro w ones. Also, the dark colour may vary from almost black to light brown and can be variably distributed, but it is always present on spiral 1 and on a few nodules of spiral 2. Examination of a shell from Florida (Figs. 4H-4J) revealed a dark subsutural spiral and a dark base. It appeared slightly different from average examples of M. olivaceus (like Fig. 4K), as the colour is not only darker but is also slightly differently distributed; spiral 1 has only isolated white beads and the brown color does not form blotches, spiral 2 is mostly white and spiral 3 is totally white, without any brown colour between the beads. Also the beads on spiral 2 become comparatively larger several whorls before the final whorl. However, it is considered to be only a variety. Genus Isotriphora Cotton and Godfrey, 1931 Type species (by original description) Triforis tasmanica T. Wood, 1875. Recent, South Australia, Tasmania. Description : The most important cióse to that of the genus Eutriphora character of this genus is the paucispiral Cotton and Godfrey, 1931, but the latter protoconch with nodular spiral cords, has axial riblets. Operculum multispiral. 90 RoláN AND FERNÁNDEZ-GARCÉS: New data on the Caribbean Triphoridae Figure 3. Protoconchs of Metaxia species. A-D. M. rugulosa (C. B. Adams, 1830) from Cuba (A) and Bahamas (B-D). E. M. espinosai Rolán and Fernández-Garcés, 1993, Cuba. Figura 3. Protoconchas de especies de Metaxia. A-D. M. rugulosa (C. B. Adams, 1850) de Cuba (A) y Bahamas (B-D). E. M. espinosai Rolán y Fernández-Garcés, 1993, Cuba. Isotriphora guanahacabibes spec. nov. Rolán and Fernández-Garcés (Figs. 5A-H) Type material: Holotype (Fig. 5A) in MNCN (15.05/15.05/47054) and paratypes in the following collections: MHNS, 2 s (Fig. 5B); IES, 1 s (Fig. 5C); CFG (3 s, 8 f), all from the type locality; 1 f and 1 j, from Cienfuegos, Cuba (MHNS); 1 s, outer slope of beach, W of Georgetown, Cayman Is. 100- 115 fms (Tyler, Oct. 1964) (ANSP 300623). Other material examined: 2 f, Cienfuegos Bay, Cuba 40 m; 1 j, S shore of island, 3 miles E of South- west Point, Cayman Brac Island (ANSP 397055). Type locality: From sediment at Cueva de Pipo, Maria la Gorda, Guanahacabibes, Pinar del Río, Cuba, at 35 m. Etymology: The specific ñame comes from the type locality. Description : Shell (Figs. 5A-5C) whitish, cortical elongate, solid. Proto- conch (Figs. 5D-5F) paucispiral with about 300 pm in diameter, the number of whorls not appreciable due to indetermi- nate separation from the teleoconch. The protoconch begins with a very narrow nucleus and the following two whorls have two prominent nodular spirals, with another very small one below the suture. Teleoconch with about 10 whorls and with a straight profile; each 91 Iberus, 26 (1), 2008 whorl with two widely separated nodu- lous spiral cords (spirals 1 and 3), the lower one more prominent; about the 6- 7^ whorls, a small cord (spiral 2) appears closer to spiral 1 (Fig. 5G). Spiral 2 con- tinúes onto the subsequent whorls, becom- ing narrow, undulating and slightly nodu- lous towards the end. Spiral 2 is always smaller than spiral 1 and 3 and is closer to the one above. On the last two whorls an additional very small, smooth spiral 4 appears at the suture. The aperture is ovoid, the outer lip fine and prominent. Near the end of the final whorl there is a small hole, located cióse to the apertural border on the upper part of the whorl. Siphonal canal closed, curving towards the rear. The colour is totally white, but some specimens can have small areas of light brown (Fig. 5C). Under high magni- fication spiral striae (Fig. 5H) can be seen between the spirals. Dimensions : The holotype measures 5.5 x 1.6 mm. Distribution: Known from the type locality and from the Cayman Islands, down to 183-210 m. Remarles: This species must be com- pared with the other congeneric species with a similar protoconch: Isotriphora peetersae (Moolenbeek and Faber, 1989) is dark brown, except for the protoconch and first whorls of the teleoconch, and the shell is more obese and with fewer whorls. Furthermore, the anal sinus is at the end of the spire and is not closed. Isotriphora taenialba Rolán and Espinosa, 1994, is similar to I. peetersae but a little larger, and always has dark and lighter bands of colour; also the anal sinus is open at the end of the spire. At first sight it might be confused with species of the genus Iniforis Jousseaume, 1884 such as I. immaculata, which is totally white, but that species has a protoconch with only one smooth fine cord. Genus Eutriphora Cotton and Godfrey, 1931 Type species (o. d.): Triphora cana Verco, 1909, Recent, Australia. Description: After Marshall (1983), similar to the genus Isotriphora but with multispiral protoconch with axial riblets. Radula with the rachidian tooth having three cusps, lateral teeth with 4-5 and numerous marginal teeth with an elongate central cusp. Operculum pau- cispiral. Eutriphora auffenbergi spec. nov. Rolán and Lee (Figs. 6A-I) Type material: Holotype (Figs. 6A-6B) in FLMNH. Paratypes: 1 s (Fig. 6C) (BMSM); 1 s (USNM); 1 s (CHL), all from type locality and ex CHL. Other paratypes: 3 s, 1 j, off Egmont Key, Hillsbor- ough Co., Florida, 76 m (Jim Moore /Cavalier, 1962) (CHL); 1 j, 65 miles E St. Augustine, St. Johns Co., Florida (T. Yocius, Jun. 1972), 54 m (CHL); 1 j, 32 miles E St. Augustine, St. Johns Co., Florida (T. Yocius, Jun. 1981), 30 m (CHL); 4 s (Fig. 6D), Florida, (MCZ 356088); 1 s (Fig. 6E), Florida, 5 m (MCZ 356096); 1 s, Hillsborough Co., W of Egmont Key, Florida, 274 m (FLMNH 169703); 1 sp, Palm Beach Co., E of Palm Beach, Florida, 183-578 m (FLMNH 257207). Other material examined (in bad condition): 1 s, 10 f, Pinellas Co., WSW John's Pass, Florida, (D. Steger) 68 m (FLMNH 238662). Type locality: West of Dry Tortugas, Monroe Co., Florida, USA (Jim Moore/ Cavalier, 1972), 90 m. Etymology: The species is named after Kurt Auffenberg, former Collections Manager in Malacol- ogy. Florida Museum of Natural History, USA. Description : Shell (Figs. 6A-6E) spiral, with about 4 whorls, the surface sharp-pointed, elongate, solid, brown or of the apex with small tubercles and the light brown. Protoconch (Fig. 6G) multi- subsequent whorls with two small 92 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae spiral cords crossed by numerous fine axial ribs that have a slight S-shaped curve. Teleoconch with about 17-18 whorls in large specimens, beginning with spirals 1 and 3, crossed by proso- cline axial ribs which form small nodules at the intersection points. On whorls 4-5, spiral 2 appears cióse to spiral 1; on the subsequent whorls, the three spirals are similar in size but spiral 3 is a little more prominent. An addi- tional spiral cord, appearing on about whorls 10-12, is slightly nodulous, always smaller, and is located just below the suture (Fig. 6F). This cord is more evident on the final whorl, where it occurs near the periphery. Below it there are two more smooth spiral cords, the lower one on the base of the siphonal canal (Fig. 6C). Aperture rounded, cól- umella curved, siphonal canal elongate and recurved, closed at its base by an extensión of the aperture. Dimensions: The holotype measures 21.8 mm, with 20 whorls. Other shells are smaller. Operculum (Fig. 6H): paucispiral, subcircular, yellowish, rather transpar- ent, with the nucleus subcentral. Radula (Fig. 61): relatively large, with a formula 16-1-1-1-16, the rachid- ian tooth having three prominent cusps; the lateral has 5 cusps of which cusps 2 and 4 are smaller. The marginal teeth have a filiform central elongate cusp, with the other two short and similar in size. Distribution : The species has been found only in Florida. Remarks: The generic assignment is based on the similarity of the radula, operculum and protoconch to the type species of the genus Eutriphora : E. armil- lata (Verco, 1909) from Australia (see Marshall, 1983). The holotype of E. auffenbergi spec. nov. resembles the figure of T. triserialis in Dall (1889: pl. 20, fig. 6a) (Fig. 36E), but the following characters of the present species are not coincident: the shell is brown and the protoconch also is darker, typically with 4 whorls and with tubercles on the apex; the siphonal canal is elongate. All these characters are dis- similar from DalTs species (see below). Comparison and differentiation must be made with the following Caribbean species described here: "Triphora" abrupta has a shorter shell, with the nodules less spherical, spiral 1 is very small and cióse to the suture, almost similar to spiral 4, while in T. auf- fenbergi spiral 1 is similar to the other main two. "T. " georgiana has a very small spiral 1 which remains reduced on the final whorls, and the protoconch probably has fewer whorls. "T. " caracca has two main spirals and between them a very narrow spiral 2; the axial ribs are very prosocline. "T. " indígena has a more convex profile to the whorls; the spiral cords are wider and the nodules at the intersec- tion points are very small, resulting in rectangular pits in the interspaces between cords and ribs. E. auffenbergi is somewhat similar to Sagenotriphora osclausum (Rolán and Fer- nández-Garcés, 1995) (see below), but the latter species is shorter (usually with no more than 9 teleoconch whorls), and the siphonal canal is also very short and curved. Other brown species are Marshallora nigrocincta (C. B. Adams, 1839) and M. modesta (C. B. Adams, 1850), but these species have a shorter spire (not more than 10 whorls), a more convex profile and a very short siphonal canal. Genus Marshallora Bouchet, 1985 Type species: Murex adversus Montagu, 1803 (o.d.). Diagnosis: After Bouchet (1985), protoconch of 4-5 whorls with two spiral threads, granules on its apical part, teleoconch with the second cord appearing later, operculum paucispiral and radula with central and lateral teeth 93 Iberus, 26 (1), 2008 with two groups of cusps. Caribbean Garcés (1995) and Rolán and Crúz- species in Rolán and Fernández- Ábrego (1996). Marshallora ostenta spec. nov. Rolán and Femández-Garcés (Figs. 7A-N, 8E, 8F) Type material: Holotype (Figs. 7A, 7B) in MNCN (15.05/47055) and a paratype (Fig. 7C) in IES, both from type locality. Other paratypes in the following institutions: 1 s (Fig. 7E), off Mayport, Duval Co., Florida (Yergin, Jan. 99), 30 m (FLMNH, ex CHL); 2 s (Figs. 7F, 7G), off Big Pine Key, Monroe Co., Florida, 30-50 m, (K. Sunderland, Aug/91) (BMSC ex CHL); 1 s, off Big Pine Key, Monroe Co., Florida, 30-50 m (K. Sunderland, 1991) (CHL, Fig. 71); 3 s, off Egmont Key, Hillsbor- ough Co., Florida (J. Moore/ Cavalier , 1962), 72 m (CHL); 1 s (Fig. 7H), 32 miles E. St. Augustine, St. Johns Co., Florida (T. Yocius, 1972) 30 m (USNM ex CHL); 8 s, Palm Beach Co., Florida, off Boynton Inlet (D. and H. Akers, Dec. 1969) 33 m (FLMNH 177040); 1 s (Fig. 7J) Miami, Florida, 73 m (FLMNH UF363887); 1 s (Fig. 7K), Palm Beach Co., Florida, off Dodge Estate (R/V Tritón, Dec. 1951) (McGinty coll.) (FLMNH 178201). Other material examined: Cuba: 2 sp (Fig. 7D)(destroyed for radular study), Cienfuegos (MHNS). USA: Florida: 1 sp (destroyed for radular study) and 2 s. Anclote Key, near Tarpon Springs, Pasco Co. (CHL); 1 s, off Dry Tortugas, Monroe Co., Riley Black (ex J. Dawley, 1/4/1983), 55 m (CHL) (the last two lots not in good condition); 4 s, off Destin, 25 m (ANSP 252157); 1 s, Palm Beach Co., off Yamato Rocks, (F. B. Lyman/ Apr. 1939) 11 m (FLMNH 10242); 3 s, St. Augustine (FLMNH 286017). Type locality: Cienfuegos Bay, Cuba. Etymology: The specific ñame derives from the Latín past participle ostentus -a, -um which means "shown", indicating that the review of material from several collections allowed us to rec- ognize this species. Description : Shell (Figs. 7A-7K) light, solid, conical elongate. Protoconch (Figs. 8E, 8F) dark brown, multispiral, with between 3 l/i and 4 whorls; the apex is covered by small tubercles; fine axial ribs are crossed by two spiral cords on the lower whorls and sometimes by a single cord on the first whorl. Teleo- conch with 7-10 whorls, the first 4-5 with only two beaded cords (spirals 1 and 3); spiral 2 appears between them on about the fifth or sixth whorl, increasing in size until the three spirals are the same size on the final whorl. Another very small, smooth spiral occurs at the suture. The colour is usually white at the beginning of the teleoconch, sometimes including the first 1-2 whorls; subsequently the colour of the shell is more variable: frequently uniform cream or light brown, darker at the suture; the beaded cords may be dif- ferent, whitish, cream or light brown, and either the lower or upper cord can be lighter. Aperture rounded; base brown, sometimes only light brown, the darker colour extending to the col- umella; siphonal canal short and open. Dimensions: The holotype measures 4.2 mm; some paratypes reach 5.5 mm. The soft parts (examined in only two specimens) are whitish, slightly translu- cent, with milk-white spots on the head and a few on the tentacles. Operculum (Fig. 7N) rounded- ovoid, multispiral, with the nucleus somewhat deviated from the centre. Radula (Figs. 7L, 7M) studied in one specimen from Cuba and another from Florida, has a formula n-l-l-l-n. The rachidian tooth has two parts with 5-6 sharp cusps at each side; the lateral ones are very similar to the central one; the marginal ones are sharp pointed, very elongate and narrow. Distribution: Known from Cuba and Florida. Remarks : In spite of the amount of material examined, it was not possible to arrive at a description that represented all the shells, suggesting that this must be a variable species. There are some dif- ferences in the protoconch (number of whorls, one or two spiral threads on the first whorl), but the main differences concern the very variable colour distrib- 94 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 4. A. Latitriphora albida (A. Adams, 1854) (BMNH). B, C. Monophorus olivaceus (Dalí, 1889), lectotype, 10 mm (USNM). D-F. M. olivaceus typical shells from Cuba, all for comparison with the shell of the lectotype. G. Nototriphora decorata (C. B. Adams, 1850), Cuba. H-K. Monophorus olivaceus (Dalí, 1889). H-J: shell, 7.0 mm, Peanut Island, Florida (MHNS); K: shell, 8 mm, Florida (FLMNH). Figura 4. A. Latitriphora albida (A. Adams, 1854) (BMNH). B, C. Monophorus olivaceus (Dalí, 1889), lectotipo, 10 mm (USNM); D-F. M. olivaceus, conchas típicas de Cuba, todas para compara- ción con la concha del lectotipo. G. Nototriphora decorata (C. B. Adams, 1850), Cuba. H-K. Monophorus olivaceus (Dalí, 1889). H-J: concha, 7,0 mm, Peanut Island, Florida (MHNS); K: concha, 8 mm, Florida (FLMNH). 95 Iberus , 26 (1), 2008 ution on the teleoconch and the initial appearance of spiral 2 between the 5th and 7th teleoconch whorls. All of this suggested to us that more than one species might be involved in the studied material. Anyway, some shells with clear differences ha ve been separated from the initial lot, and these are described below (the following species). The remainder were considered to represent a single variable species, which future study of live collected material may confirm through observation of the colour of the soft parts and examination of radulae. Similiphora intermedia (C. B. Adams, 1850) always has the spiral cords white and brown, with a rather uniform color pattern of a darker upper cord and a dark or light brown middle one; the soft parts are black. The radula has numer- ous teeth all similar in size. Marshallora modesta (C. B. Adams, 1850) and M. nigrocincta (C. B. Adams, 1839) have similar shells but they are darker brown and never whitish. The protoconchs are somewhat different, being wider and shorter (as can be seen in Figures 8H and 81). Marshallora apexdiversus spec. nov. Rolán and Lee (Figs. 8A-D, 8G) Type material: Holotype (Fig. 8A) and 3 s (Figs. 8B, 8C), paratypes (FLMNH UF363887) from type locality; 1 s, 1 s (Fig. 8D), Palm Beach Co., Florida, off Dodge Estate (R/V Tritón, Dec. 1951) (McGinty coll.) (FLMNH 178201). Type locality: Miami, Florida, 73 m. Etymology: The specific ñame derives from the Latin words diversus, which means "different", and apex, referring to an important character for separation from other cióse species. Description : Shell (Figs. 8A-8D) light, solid, conical elongate. Protoconch (Fig. 8G) light brown, with 5 whorls; the apex is covered by small tubercles and is very narrow; the whorls with fine axial ribs that are crossed by two spiral cords, the upper cord almost in the middle of the whorl and the lower one closer to the suture be- low. Teleoconch with 8-11 whorls, the first 2-3 with only two beaded cords (spirals 1 and 3); spiral 2 appears between them on about the fourth whorl, increasing in size until the three cords are of equal size on about the fifth or sixth whorls. Another very small, smooth spiral occurs at the su- ture on the final whorls. The colour is usu- ally white on the first 2 whorls of the teleoconch; on subsequent whorls spiral 1 is light brown, and on some shells (but not on the holotype)this colour also reach- es the suture. The nodules are always lighter than the ground colour. Aperture rounded-ovoid; base cream, siphonal canal short and closed at the aperture by an extensión of the outer lip. Dimensions: The holotype measures 6.2 mm. Distribution : Only known from Florida. Remarks : The present species is tenta- tively included in Marshallora due to sim- ilarity with the other Caribbean species in this genus. At the beginning of this study the shells included in this taxon were placed in the variable M. ostenta spec. nov. Finally, both species were considered dif- ferent on the basis of the following char- acters: M. ostenta almost always has a dark brown base, the protoconch has fewer whorls (see Figs. 8E, 8F in comparison with Fig. 8G), the apex is wider, the spiral threads on the protoconch are in the middle of the whorl; on the teleoconch, spiral 2 never begins before the fifth whorl, sometimes even later. Similiphora intermedia (C. B. Adams, 1850) always has very dark colour on spiral 1, while spiral 2 is light or dark brown. Marshallora modesta (C. B. Adams, 1850) and M. nigrocincta (C. B. Adams, 1839), have similar shells but in both cases they are darker brown and never whitish or cream. The protoconchs of both species are rather different, being wider at the apex and shorter (as can be seen in Figures 8H and 81, in compari- son with Fig. 8G). 96 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 5. Isotriphora guanahacabibes spec. nov. A: holotype, 5.5 mm (MNCN); B: paratype, 4.7 mm (MHNS); C: paratype, 4.35 mm (IES). D-F: protoconchs of paratypes (MHNS); G: detail of the end of the last whorl, paratype in figure B; H: detail of the microsculpture. Figura 5. Isotriphora guanahacabibes spec. nov. A: holotipo, 5,5 mm (MNCN); B: paratipo, 4,7 mm (MHNS); C: paratipo, 4,55 mm (IES). D-F: protoconchas de paratipos (MHNS); G: detalle del final de la última vuelta, paratipo de la Figura B; H: detalle de la microescultura. Genus Inella Bayle, 1879 Ino Hinds, 1843, p. 17. Type species (subsequent designation of Jousseaume 1844: 230): Triforis (Ino) gigas Hinds, 1843. Recent, New Guinea. Inella Bayle, 1879, p. 35. New ñame for Ino Hinds (not Ino Samsuelle, 1817). Remarks : The type species (see Marshall, 1983) has a blunt and wide protoconch with between two and three whorls, each with two spiral cords; the shells included in this genus are elongate and usually have two spiral beaded cords at the beginning of the teleoconch (three exceptionally) and three on the subsequent whorls, frequently with a fourth cord appearing at the suture. Marshall (1983) mentioned that the limits of the genus are uncertain, and the radula has numerous small and similar teeth except the rachidian, that has symmet- rical cusps. 97 Iberus , 26 (1), 2008 Inella bigemma (Watson, 1880) (Figs. 9A-B, 36B) Cerithium (Triforis) bigemma Watson, 1880. Prelim. Report., pt 5. Journ. Linnean Soc. London, 15: 101, sp. 2. [Type locality: 18° 38' 30" N, 65° 5' 30" W, Off Culebra Island, 713 m]. Type material: Syntype (BMNH 1887.2.9.1762) (Fig. 9A) here designated as lectotype. Other material examined: 1 s (Fig. 9B), (BMNH 1915.12.31.199) from Irish Fishery Board. Description : See Watson (1881, 1886). The protoconch is unknown. Two spiral cords are present on the first whorls, subsequently with an additional smaller, less prominent cord appearing above them. About 17 whorls on the lec- totype. Dimensions: The lectotype measures 13.5 mm. Distribution : Only known with cer- tainty from the lectotype. Culebra Island (between Puerto Rico and Virgin Islands). Remarks: The shell mentioned in the original description as having 0.6 inch (=15 mm) and the original figure (Fig. 36B) are coincident in size and number of whorls with the shell studied by us, and probably it is the same. The second shell studied (Fig. 9B) has a different profile and could be a different species (Bouchet and Warén, 1993 are of the opinión that this shell is Strobiligera brychiá). Without protoconchs it is not pos- sible to make a correct comparison. This species is very similar to I. torticula (see below, in Remarks for that species). This taxon is mentioned in Dall (1981, 1989), who suggested that the species Triforis torticula could be a variety of T. bigemma. As the shells are not in perfect condition and lack proto- conchs, not enough information was obtained to decide if they represent dif- ferent species, but the whorls seem to expand more slowly on the type of T. torticula, although not on the other shells studied. Inella inflata (Watson, 1880) (Figs. 9C, 36G) Cerithium (Triforis) inflatum Watson, 1880. Prelim. Report., pt 5. Journ. Linn. Soc., 15: 103. [Type locality: 18° 38' 30" N, 65° 5' 30" W, St. Thomas, N of Culebra Island, Danish West Indies, 713 m; indicating: Habitat.- Yucatán Strait, 1170 m (Dalí).]. Triforis inflatum Watson: Dall, 1881. Bull. Mus. Comp. Zool., 9: 6. Triforis (Sychar) inflata Watson: Dall, 1889. Bull. Mus. Comp. Zool., 18: 249. Type material: Syntype (BMNH 1887.2.9.1766) (Fig. 9C) here designated as lectotype. Description : See Watson (1881, 1886). The protoconch is pupoid and perhaps with about two whorls, the first one wider than the second. It may be about 500 pm in diameter. The teleoconch is in bad condition, but it is possible to see the two main beaded spirals, and one more near the suture. The nodules are concave above and pointed. Dimensions: The lectotype measures 4.8 mm. Distribution: Only known from the type locality. Remarks: No other syntype of this species was found in other museums and the shell is nearly identical to that represented in the original figure of I. Inflata (Fig. 36G), for which 13 whorls are mentioned (really 12 including the protoconch) and the correct size of 4.8 mm. Therefore, probably it could be the only type specimen but, as no holotype was mentioned in the original descrip- tion, this shell is here designated as lec- totype. Its protoconch is very different from most others from the Caribbean area, and is therefore a differential char- acter. Dall (1889) placed his species T. ibex as a variety of I. inflata Watson. Both 98 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 6. Eutriphora aujfenbergi spec. nov. A, B: holotype, 21.8 mm, off Dry Tortugas, Florida (FLMNH); C: paratype, 8.4 mm, same locality (BMSM); D: paratype, 13.7 mm, Fia., USA (station 4)(MCZ 336088). E: paratype, 12.1 mm, 14 whorls, Fia., USA, 5.5 m (MCZ 356096). F: detail of sculpture of the holotype; G: protoconch of the paratype in fig. C; H: operculum (of a specimen from Palm Beach, Florida, 100-300 fms (FLMNH); I: radula (from the same specimen). Figura 6. Eutriphora auffenbergi spec. nov. A, B: holotipo, 21,8 mm, de fuera de Dry Tortugas, Florida (FLMNH); C: paratipo, 8,4 mm, la misma localidad (BMSM); D: paratipo, 13,7 mm. Fia., USA (estación 4) (MCZ 356088). E: paratipo, 12,1 mm, 14 vueltas, Fia., USA, 5,5 m (MCZ 356096). F: detalle de la escultura del holotipo ; G: protoconcha del paratipo en la fig. C; H: opérculo (de un ejemplar de Palm Beach, Florida, 100-300 brazas (FLMNH); I: rddula (del mismo ejemplar). 99 Iberus, 26 (1), 2008 taxa do not appear to be similar: I. ibex lacks a protoconch and the type, with 14 whorls, is 10.4 mm; the description of I. inflata mentions 13 whorls and 4.8 mm, but the shell is less mature than the type specimens of I. ibex. Comparison of the types shows a different sculpture, the suture is deeper on I. ibex and the sutural threads are less prominent. It is more difficult to distinguish it from other species of Inella and this shall be done below. Inella longissima (Dalí, 1881) (Figs. 10A-G, 36D) Triforis (Ino) longissimus Dalí, 1881. Bull. Mus. Comp. Zool., 9: 80. [Type locality: Not stated]. Triforis (Inella) longissima Dalí, 1889. Bull. Mus. Comp. Zool., 18: 246, pl. 20, fig. 10. [Habitat: Sigsbee knoll, off Havana, in 320-823 m]. Type material: One syntype, here designated as lectotype (Fig. 10A). Off Havana, Cuba 22° 9' N 82° 21' 30W, 175 m. Blake 1877-1878 Exped. Sta. 56 (MCZ 7381). Other material studied: Bahamas: 1 f, Tamarind, Grand Bahama Island 26° 30' 45"N, 78° 36' 00" W (J. Worsfold) (ANSP 368279); Florida: 1 j (Fig. 10F) with protoconch, off Alligator Reef Light, lower Florida Keys (J. Moore, 1965) (ANSP 312592); 1 s (Fig. 10B), 1 j, SW of Egmont Key, (D. Steger, 1966) 366-420 m (ANSP 306391); 1 s, Monroe Co., 205° off Sombrero Light, (McGinty, Jul. 1951), 335-367 m (FLMNH 249518). Description: See Dall (1881). Shell (Fig. 10A) elongate, solid, whitish. The protoconch is not known from the syn- type, but a juvenile (Fig. 10F), which could be this species, shows a paucispi- ral protoconch of about 400 pm, with a pupoid apex and three whorls with two spiral cords. On this juvenile, the begin- ning of the teleoconch has only two spi- rals, but after a few whorls the third very small spiral appears between them. The teleoconch in large shells can reach up to 27 or more whorls (Fig. 10B). The profile of the whorls is straight, with four spirals: spirals 1 and 3 are almost similar in size and between them is spi- ral 2, which is initially slightly smaller but is subsequently of similar size; very cióse to the suture there is one more smooth spiral cord (Fig. 10D). Spiral 3 is a little more prominent than spirals 1-2. The nodules are always rounded. Dimensions: The lectotype measures 26.7 mm. A larger specimen, supposedly of this species, has a more prominent spiral 3 on the final whorls (Fig. 10B) and the nodules are a little larger. Distribution: Known from Bahamas, Florida and Cuba. Remarks: The syntypes with this label from USNM and MCZ were exam- ined. The problem is that the two lots apparently represent two different species. In this situation, after the description of Dall (1881) was carefully revised, it was decided to desígnate as lectotype the syntype in MCZ, due to the fact that the characters of this shell more closely matched the original description. In this description, the author pointed out that: "all the shells were decapitated" (which is not the case in the USNM syntype), and he only mentioned three spiral cords; further- more, the shell designated as lectotype has characters that are more similar to those of the figure of I. longissima repre- sented in Dall (1889: pl. 20, fig. 10)(Fig. 36D), and is cióse to 26 mm in size. The syntype in USNM (with the label T. longissimus ) has a size of 22 mm, differ- ent to the dimensions mentioned by Dall (1881), has a perfect protoconch and has only two spirals on the first 6-7 whorls of the teleoconch; additionally, a third and fourth cord appear, but these differ from the figure in Dall (1889). Also the siphonal canal is larger. For all these differences it was considered to be a sepárate species, and will be described below as new ( I . pseudolongissima). The shell from ANSP (Fig. 10B) may be the same species, but spiral 2 is very small 100 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 7. Marshallora ostenta spec. nov. A, B: holotype, 4.2 mm, MNCN; C: paratype, 3.6 mm (IES); D: shell, 3.9 mm, specimen destroyed for radular studies; all from Cienfuegos, Cuba; E: paratype, 3.9 mm, off Mayport, Florida (FLMNH); F, G: paratypes, 3.3 and 5.4 mm, off Big Pine Key, Florida (BMSM); H: paratype, 3.5 mm, off St. Augustine, St. Johns Co., Florida (USNM); I: paratype, 5.5 mm, off Big Pine Key (CHL); J: paratype, 5.5 mm, Florida (FLMNH); K: paratype, 5.3 mm, Florida (FLMNH); L: radula from a specimen from Cienfuegos; M: radula from a speci- men from Florida; N: operculum, from Cienfuegos. Figura 7. Marshallora ostenta spec. nov. A, B: holotipo, 4,2 mm, MNCN; C: paratipo, 3,6 mm (IES); D: concha, 3,9 mm, ejemplar destruido para estudios radulares; todos de Cienfuegos, Cuba; E: paratipo, 3,9 mm, fuera de Mayport, Florida (FLMNH); F, G: paratipos, 5,5 y 5,4 mm, fuera de Big Pine Key, Florida (BMSM); H: paratipo, 3,5 mm, fuera de St. Augustine, St. Johns Co., Florida (USNM); I: paratipo, 5,5 mm, fuera de Big Pine Key (CHL); J: paratipo, 5,5 mm, Florida (FLMNH); K: paratipo, 5.3 mm. Florida (FLMNH); L: rádula de un ejemplar de Cienfuegos; M: radula de un ejemplar de Florida; N: opérculo, de Cienfuegos. 101 Iberus, 26 (1), 2008 from the first teleoconch whorls, and on the final whorls spiral 3 is more promi- nent than on the lectotype. However, as these minor differences probably only indícate some variation in the species, the shell has been kept in this taxon. In spite of there being no information about the protoconch of the lectotype of this species, it has been considered to be- long to the genus Inella on the basis of the protoconch present on a juvenile, which probably represents the same species. Inella pseudolongissima Rolán and Femández-Garcés spec. nov. (Figs. 11 A-H) Type material: Holotype (Fig. 11 A), a paratype (Fig. 11D) and a fragment (USNM 87316: syntypes of Triforis longissimus ) from the type locality; 1 paratype (Fig. HE), 26° 40' N (M.E. Powlus, July 1960) (McGinty coll.) 77 m, (FLMNH 171183); 1 paratype (Fig. 11G), Hillsborough Co., Tampa, Florida, West of Egmont Key (J. Moore, ]un. 1962) (McGinty coll.) 366 m (FLMNH 158180). Type locality: Cuba, off Havana, 823 m. Etymology: The specific ñame alludes to the similarity with 1. longissima, with which this species was confused. Description : Shell (Figs. 11A, 11D, 11E, IIG) very elongate, irregularly white or creamish. Protoconch (Figs. 11B, 11F, IIH) multispiral with nearly 4 whorls, almost 1 mm in height and with a diam- eter of about 500 pm, sculptured with 2 spiral cords that are more evident on the last whorls; these two cords are unap- parent or only insinuated on the first two whorls. Teleoconch with between 18 and 24 whorls, the first whorls with two beaded spiral cords (spirals 1 and 3), the nodules being rounded; below, on whorls 3-4, a small, very narrow cord (spiral 2) appears between them; this new cord is always smaller than the other two and closer to the upper one; spiral 3 subsequently increases in size and is very clearly the largest on the last 4-5 whorls, the other two being smaller; also an additional small, unbeaded cord appears at the suture. Axial ribs are almost not appreciable; the growth lines are prosocline. Aperture rhomboid, siphonal canal short and open. The colour of the holotype is totally white. Paratypes are cream with some areas of light brown. Dimensions : The holotype measures 22.0 mm. Distribution : Known from Florida and Cuba. Remarles: I. pseudolongissima may be distinguished from I. longissima by teleo- conch sculpture, the latter species having three cords of equal size on most of the spire. Also spiral 2 appears on the first whorls of I. pseudolongissima but on about the seventh whorl in I. longissima. The protoconch of I. longissima has three whorls with two well defined cords, while the protoconch of I. pseudolongis- sima has four smooth whorls, and the apex is smaller. For differentiation from other cióse species, see below. Inella noduloides spec. nov. Rolán and Fernández-Garcés (Figs. 12 A, B) Type material: Holotype (Figs. 12A, 12B) in ANSP (374588). Type locality: Tamarind, Grand Bahama Island, Bahamas 26° 30' 45"N, 78° 36' 00" W, 500 m (J. Worsfold, May, 1985). Etymology: The ñame alludes to the prominent nodules of the whorls. Description: Shell (Figs. 12A, 12B) small, subcylindrical, elongate, whitish and solid. Protoconch with a pupoid apex of about 380 pm diameter and 3 whorls, each with two smooth spiral cords; the lower cord becomes more prominent near the end, where another smooth, less prominent cord appears above the other two, very 102 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 8. A-D. Marshallora apexdiversus spec. nov. A: holotype, 6.2 mm, Florida (FLMNH); B-D: paratypes, 6.9, 6.5, 4,6 mm, Florida (FLMNH). E-I. Protoconchs of Marshallora species. E, F: M. ostenta , Florida; G: M. apexdiversus , Florida; H: M. modesta, Cuba; I: M. nigrocinta, Cuba. Figura 8. A-D. Marshallora apexdiversus spec. nov. A: holotipo, 6,2 mm, Florida (FLMNH); B-D: paratipos, 6,9, 6,5, 4,6 mm, Florida (FLMNH). E-I. Protoconchas de diferentes especies de Marshal- lora. E, F: M. ostenta. Florida; G: M. apexdiversus. Florida; H: M. modesta, Cuba; I: M. nigrocinta, Cuba. 103 IberuSy 26 (1), 2008 cióse to the suture; under high magnifi- cation numerous tubercles can be seen. On the first two whorls of the teleoconch there is one prominent cord in the middle and another smaller one below the suture; both ha ve prominent nodules numbering about 10-11 per whorl. On the subsequent whorls there is a small nodulous cord below the suture with another small, slightly nodu- lous one below it; these are followed by the most prominent cord, with rather sharply pointed nodules, and one more below it, cióse to the lower suture. The aperture is rounded but irregular, as the shell is juvenile. Dimensions : The holotype measures 2.1 mm. Distribution : Only known from the type locality. Remarks: No other species has these characters, with one spiral cord with very prominent nodules and 3 weaker spirals. The closest species may be the new species I. harryleei (see below). Inella apexbilirata spec. nov. Rolán and Fernández-Garcés (Figs. 12C-E) Type material: Holotype (Fig. 12C) (ANSP 367841); 4 paratypes (ANSP 367840) (Fig. 12D). All from the type locality. Type locality: Lucaya, Grand Bahama Island, Bahamas 26° 29' 45" N, 78° 37' 15" W (J. Wors- fold), 300 m. Etymology: The specific ñame alludes to the beginning of the protoconch, with two spirals com- mencing from the nucleus. Description : Shell (Figs. 12C, 12D) small, subcylindrical, elongate, shiny, whitish and solid. Protoconch (Fig. 12 E), shining white, with a very small nucleus, a diameter of nearly 400 pm and a height of about 660 pm, 3 whorls with two smooth spiral cords which begin just on the nucleus. The teleo- conch has two main beaded cords, the lower one slightly larger; one smooth smaller cord occurs at the suture. The nodules on the main spirals are pointed and cut at the middle. The axial ribs which connect the nodules are scarcely evident and are slightly prosocline. There are about 13 on the first whorls and 17 on the last. Aperture ovoid, siphonal canal very short and open. Dimensions : The holotype measures 3.5 mm. Distribution : Only known from the type locality. Remarks: The species with two main spirals are the following: I. bigemma and I. inflata have whorls that increase quickly in width, and the latter species has a pupoid protoconch. I. harryleei spec. nov. (see below) has spirals 1 and 3 at the beginning, spiral 2 appearing later and being smaller. I. ibex has the same number of nodules in spite of being a more devel- oped shell. Besides the two main spirals there is only one more near the suture. I. torticula has three spirals on the first whorls and 4-5 on the last ones. I. colon has two spirals but equidis- tant from both sutures. I. compsa also has two main spirals, with spiral 1 being smaller; its proto- conch is unknown, but the whorls on this shell increase rapidly in width. I. enopla and I. meteora have a very different wide, pupoid protoconch. Inella pinarena Espinosa, Ortea and Fernández-Garcés, 2007 (Figs. 12F-J) Inella pinarena Espinosa, Ortea and Fernández-Garcés, 2007. Avicennia, 19: 73, fig. 46. [Type local- ity: Yemayá, Maria la Gorda, Guanahacabibes, Pinar del Rio, Cuba], Description: See Espinosa, Ortea and 12F. 12G), are shown in SEM pho- Fernández-Garcés (2007). Shells (Figs. tographs. We add some informations on 104 ROLÁN AND FERNÁNDEZ-GARCÉS: New data on the Caribbean Triphoridae Figure 9. A. Cerithium bigemma Watson, 1880. lectotype, 13.5 mm (BMNH). B. Inella bigemma , shell, 13.0 mm (BMNH). C. Cerithium inflatum Watson, 1880, lectotype, 4.8 mm (BMNH). Figura 9. A. Cerithium bigemma Watson, 1880. lectotipo, 13,5 mm (BMNH). B. Inella bigemma, concha, 13,0 mm (BMNH). C. Cerithium inflatum Watson, 1880, lectotipo, 4,8 mm (BMNH). the protoconch (Figs. 12H, 121): small tubercles are present on the protoconch surface between the spiral ribs. The microsculpture of the teleoconch (Fig. 12J) is formed by very small spiral striae, more evident on the subsutural part. Distribution : Only known from the type material. Remarks : This species was recently described but, in the original descrip- tion, only one optical photo was pre- sented. More details on the shell, proto- conch and microsculpture are here illus- trated, showing that there is no similar- ity with other species of the group in the Caribbean. Inella harryleei spec. nov. Rolán and Fernández-Garcés (Figs. 13A-K) Type material: Holotype (Fig. 13A) and 2 (s) paratypes in FLMNH; one more in BMSM (Fig. 13B) all ex CHL; another in CHL (Fig. 13C); all from the type locality. Other paratypes: 9 j and f, 28° 4.57' N, 90° 59.99' W, 87.9 m (E.F. Garcia /Pelican, 9/1-5/01) (CHL); 1 j, from the same locality (USNM); 2 j more 26° 06.866' N, 91° 02.418' W, off Louisiana, 59-65 m (E.F. Garcia /Pelican, 6/30/01)(CHL); 105 Iberus, 26 (1), 2008 1 f, off Alligator Reef Light, Lower Florida Keys (J. Moore, 1965) (ANSP 312593); 3 s, SW of Egmont Key, Florida (D. Steger, 1966) 366-420 m (ANSP 306391); 3 j, W of Tampa Bay, Florida (Richardson) 500 m (ANSP 335492); 1 s, Hillsborough Co., 110 miles SW of Egmont Key, Florida (D. Steger, 1953) 165 m (FLMNH 151609); 1 s (Fig. 13D), Hillsborough Co., SW of Egmont Key, Florida (J. Moore, 1967) 46 m (FLMNH 158178). Other material examined: 2 f, Stn. 15, Florida (MCZ 356092) which are not in sufficiently good con- dition to examine all their characters and attribute them with certainty to this species; 1 f (A.M. Norman coll.) (Blake Expedition), West Indies (BMNH acc. n° 2283). Type locality: off Dry Tortugas, Monroe Co., Florida, 90 m (J. Moore/ Cavalier, 1992). Etymology: The species is named after Harry G. Lee, American malacologist who lent us the mate- rial on which this species is based. Description : Shell (Figs. 13A-13D) elongate, very large for the genus, solid, well sculptured. Protoconch (Figs. 13G- 13K) with between 3 and 3 V4 whorls, bulbous, white, with barely discernible separation from the teleoconch, the first whorl smooth and the next with two spiral cords that are sometimes a little irregular; on the last whorl the lower cord is more prominent, and the upper one may divide in two. The teleoconch has many whorls, 22 in the holotype and 17 in a paratype (Fig. 13B). The first whorls have two spiral nodulous cords (spirals 1 and 3), connected by very narrow prosocline ribs; below, around the fourth or fifth whorls, spiral 2 appears between those cords and remains very narrow for several whorls; after the tenth whorl spiral 3 becomes a little larger, and on the subsequent whorls it is more clearly different and prominent (Fig. 13E). Spiral 2 strength- ens but continúes to be the smallest cord until the end. The suture is very deep. The axial ribs are slightly prosocline. Aperture (Fig. 13F) ovoid, siphonal canal narrow and large, dark in colour. Shell colour light brown or white with irregular brown axial flammules. Dimensions : The holotype measures 18.3 mm. The paratypes are smaller. Distribution : Only known from the type locality. Remarks: The present species must be compared to and differentiated from the following: I. longissima is uniformly white and more cylindrical, having three equal spi- ral cords throughout most of the spire, and another smaller one on the suture which is relatively shallower; the shell sculpture is very similar from the begin- ning to the end with only a little more prominence of the lowermost cord. I. pseudolongissima spec. nov. is white with 4 cords, the upper one smooth at the suture, the following two very small, and only the lowermost is clearly prominent; the suture is not deep; the protoconch has more whorls. I. noduloides spec. nov. has one prominent beaded spiral cord with three more smaller ones which are absent from the first whorls of the teleoconch of I. harryleei. Other elongate deep water species are Inella colon Dalí, 1881, and I. ibex Dalí, 1881; both have only two spiral beaded cords per whorl. Inella aff. harryleei (Figs. 14A, B) Material examined: 1 s, Manatee Co., West of Anna María Key, Florida (J. Moore, 1961) (McGinty coll.) 55 m (FLMNH 158177). Remarks : The specimen studied is in perfect condition and appears rather similar to I. harryleei spec. nov., but it differs in the following characters: a more pointed protoconch, with about two whorls and an angulation on the second whorl, without visible cords; on the teleoconch, spiral 1 is not larger than spiral 2, but is less prominent; on the base, below spiral 3, there is only one 106 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 10. Inella longissima (Dalí, 1881). A: lectotype, 26.7 mm, off Havana, 175 m (MCZ 7381); B: shell, 27.9 mm (ANSP); C, D: detail of the spire of lectotype; E: fragment of first whorls, 6.3 mm (ANSP); F: juvenile with protoconch (ANSP 312592); G: detail of last whorls of the shell from Figure B. Figura 10. Inella longissima (Dalí, 1881). A: lectotipo, 26,7 mm, fuera de La Habana, 175 m (MCZ 7381); B: concha, 27,9 mm (ANSP); C, D: detalle de la espira del lectotipo; E: fragmento de las primeras vueltas, 6,3 mm (ANSP); F: juvenil con protoconcha (ANSP 312592); G: detalle de las últimas vueltas de la concha de la Figura B. 107 Iberus , 26 (1), 2008 smooth cord and another very small one immediately below. The siphonal canal is shorter. Dimensions : The only shell studied measures 28.8 mm. All these differences indícate that this shell may represent a séparate species, but they are not enough to justify naming it. Additional material is required. Inella triserialis (Dalí, 1881) (Figs. 14C-E, 36E) Triforis (Inella) triserialis Dalí, 1881. Bull. Mus. Comp. Zool., 9: 84. [Yucatán Strait, 640 fms (1171 m). Off Cape San Antonio, 640 fms (1171 m). Station 2, 805 fms (1472 m)]. Triforis (Inella) triserialis Dalí, 1881. In Dalí, 1889. Bull. Mus. Comp. Zool., 18: 249, pl. 20, figs. 5a, 6a. Triphora aspera auct. non Jeffreys, 1885. Type material: Two syntypes (USNM 87319), one of them (Fig. 14C) measuring 6.4 mm is here des- ignated as lectotype; the other shell (5.6 mm) would be a paralectotype (Fig. 14D). Both from Cape San Antonio, Cuba, 1170 m; probably, 1 syntype (BMNH Acc. n° 2283), off Havana, 823 m (in very bad condition). Other material studied: 1 j, SW of Egmont Key, Florida (Steger, 1966) 366-421 m (ANSP 306391). Description : See Dall (1881). Proto- conch paucispiral (Fig. 14E), wide, with about 3 whorls which have two spiral cords, the apex depressed and smooth. The teleoconch begins with three spiral nodulous cords, spiral 2 being smaller and closer to spiral 1. On the following whorls spiral 1 increases in size, with nodules that are larger than those on the other cords and are cut at the middle. These differences continué up to the end of the shell. Dimensions : The lectotype measures 6.4 mm. Distribution: Known from Yucatán (México) and Cuba. Remarks: As with other taxa, the syntype material found in USNM and MCZ has been examined. The material from USNM is coincident in protoconch and teleoconch sculpture with the description of Dall (1881) and with one of the figures in Dall (1898: píate 20, fig. 5a) (Fig. 36E) but not with the other figure (Dall, 1898: píate 20, fig. 6a), which seems to be a different species with a sharply pointed apex. The size of the lectotype is slightly smaller than that mentioned in the original description (8.25 mm), but it may be that this type could be partially broken. The material from MCZ seems to be what Dall named as var. intermedia (see below). Dall (1989: 246) considered this taxon in relation with Triphora aspera Jef- freys, 1885 (preoccupied, renamed Triphora brychia Bouchet and Guillemot, 1978; assigned to genus Strobiligera Dall, 1924 by Bouchet and Warén, 1985). He therefore referred to Triforis (Inella) trise- rialis var. aspera Jeffreys, 1885, which perhaps can correspond with the figure previously mentioned (Fig. 36E). A syntype of Jeffreys' taxon (Fig. 14F, BMNH 85.11.5.2673) has been examined, and is here designa ted as lectotype. It is a larger and sharply pointed shell with a multispiral protoconch, and there is no indication that this species occurs in the Caribbean. I. triserialis is different from most of the Caribbean species referred to in this work, mainly due to its spiral 1 having larger nodules. Comparison will be made in the Remarks of some species described below (such as I. sarissa). Inella intermedia (Dall, 1881) (Figs. 14G-J, 36F) Triforis intermedius Dall, 1881. Bull. Mus. Comp. Zool., 9: 85. [Yucatán Strait, 1170 m; off Cape San Antonio, 1170 m; station 2, 1472 m]. 108 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 11. Inella pseudolongissima spec. nov. A: holotype, 22.0 mm, off Havana, 823 m (USNM); B: protoconch of the holotype; C: detail of the spire; D: paratype, 10.4 mm (USNM); E: paratype, 18.5 mm (FLMNH); F: protoconch of this paratype; G: paratype, 23.4 mm (FLMNH); H: protoconch of this paratype. Figura 11. Inella pseudolongissima spec. nov. A: holotipo, 22,0 mm, fuera de La Habana, 823 (USNM); B: protoconcha del holotipo ; C: detalle de la espira; D: paratipo, 10,4 mm (USNM); paratipo, 18,5 mm (FLMNH); F: protoconcha de este paratipo; G: paratipo, 23,4 mm (FLMNH); protoconcha de este paratipo. 109 Iberas, 26 (1), 2008 Tñforis triserialis var. intermedia Dalí, 1881. In Dall, 1889. Bull. Mus. Comp. Zoo/., 18: 287, píate 20, fig. 8. [Habitat: 805 fms (1473 m). Barbados, 183 m]. Triphora clenchi Aguayo, 1935. Unnecessary replacement ñame pro Triforis intermedius Dall, 1881. Type material: Syntype (Fig. 14G) here designated as lectotype (MCZ 7384) (off Cuba, Yucatán Strait and Cape San Antonio. Depth: 1171 m. Blake 1877-1878 Exped.); one paralectotype (Fig. 14H)(MCZ 7382) (Havana, off Morro Light. Depth: 1473 m. Blake 1877-1878 Exped.). Both shells were labeled T. triserialis. Description : See Dall (1881). No pro- toconch was known and the author refers to "nucleus missing". Two spirals of similar strength (spirals 1 and 3) are present on the first whorls of the shell; spiral 2 appears below and is clearly smaller up to near the end of the spire, where the three spirals are almost similar in size (Fig. 141). Axial ribs prosocline, and the nodules are spherical. Dimensions : The lectotype measures 7.9 mm. Distribution : From Yucatán, México, Barbados and Cuba. Remarks: The lectotype here selected is very similar to the shell represented in Dall (1989: pl. 20, fig. 8) (Fig. 36F) but it is not coincident with the description, which refers to a size of 11 mm and 23 whorls; also the figure does not agree with the description, because in spite of the fact that a height of 11 mm is men- tioned, there are not more that 15-16 whorls. The shell here selected as lecto- type is only 7.9 mm and has 12 whorls, and it may be the original shell which lost some part of the spire. Anyway, it must be kept as lectotype in order to attain nomenclatural stability. The para- lectotype is 7.2 mm and has about 13 whorls (MCZ 7382). The ñame Triphora triserialis clenchi Aguayo, 1935 is an unnecessary replace- ment ñame for Triforis intermedius Dall, 1881 (which is the same taxon as Triforis triserialis var. intermedia Dall, 1881 (in Dall, 1889)). This is due to the fact that this ñame is not preoccupied by Cerithium intermedium C. B. Adams, 1850; initially both belonged in different genera and finally the first one is placed in Inella and the second was recently placed by Rolán and Fernández- Garcés (1995) in the genus Similiphora Bouchet, 1985. Lack of knowledge of the proto- conch is a problem when comparing this species to others from the Caribbean. Basing this comparison on the teleo- conch, the following species must be taken into account: Inella longissima has a more cylindri- cal, more elongate shell, spiral 2 is of similar size to the other two spirals on most of the spire, while in I. intermedia it is always smaller. Inella pseudolongissima spec. nov. has spiral 3 more prominent and with larger nodules. Inella harryleei spec. nov. has a typi- cally deep suture, spiral 3 being the most prominent, and spiral 1 is also larger than spiral 2, these differences being maintained to the end of the shell. Inella triserialis has spiral 1 more prominent and with larger nodules. Inella ibex (Dall, 1881) (Figs. 14K-M, 36G) Triforis ibex Dall, 1881. Bull. Mus. Comp. Zool., 9: 86. [Type locality: off Cape San Antonio, 1171 m; Yucatán Strait, 1171 m; Sigsbee, off Havana, 823 m]. Triforis (Sychar) inflata var. ibex Dall, 1889. Bull. Mus. Comp. Zool, 18: 249, pl. 20, fig. 12b. Type material: A syntype (f) of 10.4 mm here designated as lectotype (Fig. 14K), off Cuba, Yucatán Strait and Cape San Antonio, 1171 m (MCZ 7391); another syntype, now paralectotype, f of 7.7 mm, off Havana, Cuba 823 m (MCZ 7392). Another paralectotype (Fig. 14L) (USNM 87313), off San Antonio, Cuba, 1189 m. 110 RoláN AND FernÁNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 12 A, B. Inella noduloides spec. nov. holotype, 2.1 mm, Grand Bahama Island (ANSP). C- E. Inella apexbilirata spec. nov. C: holotype, 3.5 mm, Grand Bahama Island (ANSP); D: paratype, 3.2 mm, Grand Bahama Island (ANSP); E: protoconch of the paratype. F-J. Inella pinarena Espinosa, Ortea and Fernández-Garcés, 2007. F: paratype, 6.0 mm (CFG); G: shell, 5.0 mm (MHNS); H: protoconch of the paratype; I: detail of the protoconch; J: microsculpture. Figura 12. A, B. Inella noduloides spec. nov. holotipo, 2, 1 mm, Grand Bahama (ANSP). C-E. Inella apexbilirata spec. nov. C: holotipo, 3,5 mm, Grand Bahama (ANSP); D: paratipo, 3,2 mm, Grand Bahama (ANSP); E: protoconcha del paratipo. F-J. Inella pinarena Espinosa, Ortea and Fernández- Garcés, 2007. F: paratipo, 6,0 mm (CFG); G: concha, 5,0 mm (MHNS); H: protoconcha del paratipo; I: detalle de la protoconcha; J: microescultura. 111 Iberus, 26 (1), 2008 Type locality: The three localities mentioned in the original description are written on the label of the lectotype, and henee there is no exact information about which locality applies to this type. Description : See Dall (1881). The orig- inal description mentions a nucleus twice as large as that of I. colon , smooth, inflated, rounded on top, larger than the succeed- ing nuclear whorl, which has two incon- spicuous narrow keels which are wavy and almost tuberculate at first, and merge almost imperceptibly into the usual sculp- ture of the shell. The type material studied has no protoconch, which obviously was lost. The most important characters of the teleoconch are: shell (Figs. 14K, 14L) with two rows of nodules throughout the spire and with one more, smaller and not nodu- lous, occurring at the suture at the end the spire (Fig. 14M). The nodules are cut at the middle. The axial ribs are scarcely appreciable and slightly prosocline. Dimensions : The lectotype measures 10.4 mm. Distribution: Known from Yucatán (México) and Cuba. Remarles : Triforis ibex is the type species of the genus Strobiligera Dall, 1924. The problem is that we have no image or a good description of the pro- toconch of this species, and the original description allows us to think that the protoconch is almost smooth, the first whorl wider than the second and with two whorls, which is typical of a proto- conch of an Inella species. In this case; Strobiligera could be a synonym of Inella. The assignment of this species to the genus Inella is tentative, The lectotype here designated is similar to the shell represented in Dall (1889, pl. 20, fig. 12b) (Fig. 36G) and also its dimensions are closer to those referred to in the orig- inal description. Comments on the differences of I. ibex from other species with two spiral beaded cords can be seen below in I. colon, which is the most similar species. Inella torticula (Dall, 1881) (Figs. 15, 36H) Triforis torticulus Dall, 1881. Bull. Mus. Comp. Zoo/., 18: 82. [Type locality: Yucatán Strait, 1170 m], Triforis (Sychar) torticula Dall, 1881. In Dall, 1889. Bull. Mus. Comp. Zool., 18: 249, pl. 20, fig. 11b. Type material: One syntype (Fig. 15A) (f) here designated as lectotype, 6.7 mm, off Cuba, Yucatán Strait, 1171 m, Blake 1877-1878 Exped. (MCZ 7390); the syntypes in USNM have been lost (Ellen Strong, pers. comm.). Other material examined: 1 f (Fig. 15B) (labeled as Triphora bigemma Watson, 1880), 9.7 mm, 9 whorls, Yucatán Strait, 1171 m (MCZ 7388); 1 s (Figs. 15C, 15D), 25.0 mm, Straits of Florida, 1000- 1200 m (CHL). Description: See Dall (1881). The most important characters are: 4 spirals, the lowermost spiral 4 very narrow and very cióse to the suture; spiral 1 smaller than spirals 2 and 3, which are equal in size and prominence. Colour whitish. The study of an almost complete adult shell (without protoconch) with 17.5 whorls (Figs. 15C, 15D) (CHL), allows us to add some information: at the beginning of the shell there are only three spirals, almost similar in size, but the uppermost slightly smaller; later, the differences between spiral 1 and spirals 2-3 are more evident, and at same time spiral 4 appears; on about whorl 13 (on this shell) the uppermost spiral 1 is divided in two, there being 5 spirals at the end of the shell (Fig. 15E). The ribs are orthocline. Dimensions: see below. Distribution: Known from deep water off Florida and Cuba. Remarks: The placement of this species in the genus Inella is tentative, as the protoconch and the aperture are not known. The syntype in MCZ (Fig. 15A), here designated as lectotype, is a shell frag- ment that seems to be the one figured in 112 RolAn and FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 13. Inella harryleei spec. nov. A: holotype, 18.3 mm, off Dry Tortugas, 90 m (CHL); B: paratype 11.7 mm (BMSM); C: paratype, 12.8 mm (CHL); D: paratype, 14.6 mm (FLMNH). E: detail of the spire of the holotype; F: detail of the aperture of the paratype in figure B; G-K: proto- conchs of several paratypes (CHL). Figura 13. Inella harryleei spec. nov. A: holotipo, 18,3 mm, fuera de Dry Tortugas, 90 m (CHL); B: paratipo 11,7 mm (BMSM); C: paratipo, 12,8 mm (CHL); D: paratipo, 14,6 mm (FLMNH). E: detalle de la espira del holotipo; F: detalle de la abertura del paratipo de la figura B; G-K: protoconchas de varios paratipos ( CHL). 113 Iberus , 26 (1), 2008 Dall (1889, pl. 20, fig. 11b) (Fig. 36H), although a little more cylindrical, having only 4 whorls instead 6 and measuring 6.7 mm instead of 10.5, which could be due to the loss of two whorls of the shell. As it is compatible with the description, it was thought that it should be designated as lectotype in order to maintain nomenclatural stabil- ity. The other shell examined (Fig. 15B) measured 9.7 mm and had 9 whorls, the label without any indication of "type" and with the ñame Triphora bigemma; it is undoubtedly the same species. The differences with I. bigemma Wat- son are very small, bearing in mind that although the latter species was de- scribed as having only two rows of beads, the figure and the type clearly show three on the final whorls. How- ever, this is a juvenile, and to make a de- cisión it would be necessary to examine the protoconchs and to compare several samples of both species. The other shell studied (Figs. 15C, 15D) has a profile slightly different from the lectotype, with the whorls expand- ing more rapidly and with 4-5 spirals on the final whorls. This could represent a different species, but it is rather coinci- dent in some characters with the shell figured by Dall (1889) (Fig. 36H) and for this reason it is considered to be an example of the variability of this taxon, which has differences between the first and the last whorls. Inella colon (Dall, 1881) (Figs. 16, 361) Triforis colon Dall, 1881. Bull. Mus. Comp. Zool., 9: 86. [Type locality: off Havana, Cuba 450 fms (823 m); off Cape San Antonio, 1171 and 1834 mi. Triforis (Inella) colon Dall, 1881. In Dall, 1889. Bull. Mus. Comp. Zool., 18: 249, pl. 20, fig. 12. Type material: One syntype (Fig. 16A) (f) here designated as lectotype, 12.7 mm, with the label "off Havana, 823 m" (USNM 87315), and another syntype (Fig. 16B) (f) 7.8 mm, now paralectotype, from the same lot. A syntype (Fig. 16C) (f) 8.6 mm, now paralectotype, from Cape San Antonio, 1171- 1834 m (MCZ 7387). Other material examined: 2 f of 16.2 and 5.6 mm, Egmont Key, Florida, Gulf of México, (J. Moor e/Cav- alier, 1968), 180 m (CHL); 1 s (Fig. 16E), Yucatán, 21° 60' N, 86° 80' W, ENW of Contoy Fight, off Cape Catoche (Riley Black, Apr. 1967) (McGinty coll.) 366-412 m (FLMNH 170221); 1 s (Fig. 16F), 24° 15' 28"N, 81° 57' 89"W, Monroe Co., Straits of Florida, 236-255 m (FLMNH 26660). Type locality: With the designation of the lectotype, the type locality is restricted to off Havana, Cuba 823 m. Description: See Dall (1881). In the original description the protoconch is mentioned as follows: nucleus and first nuclear whorls white, polished, smooth, flattened on top. The most important char- acters of the teleoconch are: shell almost cylindrical (Figs. 16A-16C), white, with two main very narrow spiral beaded cords and two smaller, smooth, slightly wavy cords located near the suture; one of them above and the other below the suture, this being almost imperceptible. The nodules are small and slightly prominent; the uppermost a little larger. The axial ribs almost inapparent, a little prosocline; whorls very short and fíat. Dimensions: The lectotype measures 12.7 mm. Distribution: Known from deep water of Florida, Yucatán and Cuba. Remarks: The lectotype (Fig. 16 A) is in accordance with the original description and it seems to be the shell figured in Dall (1889: pl. 20, fig. 12) (herein Fig. 361), having the same number of whorls and almost the same dimensions. A paralec- totype in the same lot as the lectotype (Fig. 16B) seems to be a different species, because this shell has three spirals instead of two. The paralectotype in MCZ (Fig. 16C) is a little different from the lectotype, but it has the typical distribution of the cords (Fig. 16D) and is probably the same species. Some shells from the Gulf of México (Figs. 16E and 16F) have been included in this taxon but they could rep- 1 14 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 14. A, B. Inella aff. harryleei, shell, 28.8 mm and protoconch (FLMNH). C-E. Inella triseri- alis (Dalí, 1881). C: lectotype, 6.4 mm (USNM); D: paralectotype, 5.6 mm (USNM); E: proto- conch of the paralectotype. F. Triphora aspera, lectotype, 13.0 mm (BMNH) ( =Strobiligera brychia). G-J. Inella intermedia (Dalí, 1881). G: lectotype, 7.9 mm, off Cuba, Yucatán Strait and Cape San Antonio, 1171 m (syntype of T. triserialis, MCZ 7384). H: paralectotype, 7.2 mm, (Havana, off Morro Light, 1473 m (syntype of T. triserialis , MZS 7382); I: detail of the sculpture of the paralec- totype. J: shell photographed by Kaicher (ANSP, unknown size and other data). K-M. Inella ibex (Dalí, 1881). K: lectotype, 10.4 mm, off Cuba, Yucatán Strait and Cape San Antonio, 1 171 m (MCZ 7391); L: paralectotype, 9.2 mm (USNM 87313); M: detail of the spire of the lectotype. Figura 14. A, B. Inella aff. harryleei, concha, 28,8 mmy protoconcha (FLMNH). C-E. Inella triserialis (Dalí, 1881). C: lectotipo, 6,4 mm (USNM); D: paralectotipo, 5,6 mm (USNM); E: protoconcha del paralectotipo. F. Triphora aspera, lectotipo, 13, 0 mm (BMNH) ( =Strobiligera brychia). G-J. Inella inter- media (Dalí, 1881). G: lectotipo, 7,9 mm, Juera de Cuba, Estrecho de Yucatán y Cabo de San Antonio, 1171 m (sintipo de Y. triserialis, MCZ 7384). H: paralectotipo, 7,2 mm, (La Habana, Juera de Morro Light, 1473 m (sintipo de T. triserialis, MZS 7382); I: detalle de la escultura del paralectotipo. J: concha fotograjiada por Kaicher (ANSP, tamaño desconocido así como otros datos). K-M. Inella ibex (Dalí, 1881). K: lectotipo, 10,4 mm, fuera de Cuba, Estrecho de Yucatán y Cabo de San Antonio, 1171 m (MCZ 7391); L: paralectotipo, 9,2 mm (USNM 87313); M: detalle de la espira del lectotipo. 115 IberuSy 26 (1), 2008 resent a different species, because the spirals are more prominent and are more widely separated than on the lectotype. However, the lack of protoconch and the differences of the sculpture are not very important, and so provisionally they ha ve been kept in this species awaiting more material in the fu ture. Dall (1881) referred to the differ- ences between I. íbex and I. colon, proba- bly because he considered both species very similar. Unfortunately at present their protoconchs are not known, but Dall was of the opinión that the proto- conch of I. íbex was twice as large as that of I. colon. In the teleoconch characters, I. íbex has more rapidly expanding whorls. the nodules are larger and are cut at the middle, and fewer whorls occupy the same space. The other Caribbean species may have two spirals at the beginning of the teleoconch, but finally have three or more spirals. This ineludes I. pompona and I. meteora, although spiral 1 is very small and sometimes difficult to see. Inella unicornium Simone, 2006, from Brazil, has a larger shell (up to 50 mm, the biggest known species of Triphori- dae), which according to the original description may be differentiated from 1. colon not only by its larger size, but also by its proportionally longer whorls, taller spiral cords and projecting outer lip. Inella undebermuda spec. nov. Rolán and Lee (Figs. 17A-E) Type material: Holotype (Fig. 17A) in FLMNH; a paratype (Fig. 17B), 12.3 mm, in USNM. Both from type locality (J.R.H. Lightboum and A.T. Guest, 1988; ex J.R.H. Lightbourn 9/8/93; ex CHL). Type locality: S. of Castle Roads, Bermuda, 90 m. Etymology: The specific ñame derives from the Latín word unde "origin" and the ñame of the arch- ipelago where the species was collected. Description : Shell (Figs. 17A, 17B) elon- gate, whitish and solid. Protoconch (Fig. 17C) white, without a clear separation from the teleoconch, with about two whorls, the nucleus smooth, one spiral cord appearing in the middle of the first whorl and becoming two on the second; the diameter of about 380 pm. The first whorls of the teleoconch have two spiral beaded cords (spirals 1 and 3), and a very small smooth and narrow thread very cióse to the suture; between the 6-7* whorls, spiral 2, which is also narrow and smooth, appears between the other two, closer to spiral 1. On the subsequent whorls, spiral 2 increases slowly in size, becoming slightly undulating and with nodules appearing at the end of the spire (Fig. 17D), but always remaining nearer to spiral 1. The nodules of spirals 1 and 3 are larger and cut at the middle. Aperture (Fig. 17E) rhomboid, columella almost straight, siphonal canal very short and curved. Dimensions : The holotype measures 7.8 mm, and the paratype 12.3 mm. Distribution : Only known from the type locality. Remarles : The holotype is the smallest shell, but has a protoconch in good con- dition. The paratype lacks the first proto- conch whorl. The most important differ- ential character is the smaller size of spiral 2 and its location closer to spiral 1. This species must be compared with and separated from the following: Inella longissima is more elongate and cylindrical and has the three spirals similar in size and distribution. In juveniles assumed to belong to this species, the pro- toconch has three whorls, and when spiral 2 appears on the first whorls of the teleo- conch, it is in the middle between spirals 1 and 3 and not closer to the upper one. Inella pseudolongissima spec. nov. has a protoconch with four whorls and the spirals of the teleoconch are different, spiral 3 being larger and more promi- nent than the others. Spiral 2 is only the smaller one at the beginning. Inella harryleei spec. nov. has three whorls in the protoconch, and in the teleoconch spiral 2 becomes nodulous very early (between whorls 5-6) and is not closer to spiral 1, and the suture is 1 16 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 15. Inella torticula (Dalí, 1881). A: lectotype, 6.7 mm (MCZ 7390); B: shell, (labeled as T. bigemma ), 9.7 mm, 9 whorls, Yucatán Strait, 1171 m (MCZ 7388); C, D: shell, 25.0 mm, Straits of Florida, 1000-1200 m (CHL); E: detail of sculpture. F: shell photographed by Kaicher (ANSP, lacking size and data). Figura 15. Inella torticula (Dalí, 1881). A: lectotipo, 6,7 mm (MCZ 7390); B: concha, (etiquetada como T. bigemma,), 9,7 mm, 9 vueltas, Estrecho de Yucatán, 1171 m (MCZ 7388); C, D: concha, 25,0 mm, Estrechos de Florida, 1000-1200 m (CHL); E: detalle de la escultura. F: concha fotografiada por Kaicher (ANSP, careciendo de información sobre su tamaño y otros datos). 1 17 Iberus , 26 (1), 2008 more evident. Furthermore, the colour consists of axial brown flammules and brown siphonal canal, more elongate. Inella intermedia has a shorter shell and spiral 2 is strongly nodulous on the early whorls, being almost the same size as the other two and located at a similar distance between them. Inella ibex and I. colon have only two main spirals. Inella torticula can have up to five spirals on the final whorls. Inella sarissa (Dalí, 1889) Triforis (Inella) sarissa Dalí, 1889. Bull. Mus. Comp. Zoo/., 18: 247. [Type locality: near Barbados, 13° 11.54' N, 59° 38.45' W, 134 m]. Type material: The lot in USNM labeled as "87314 Holotype" of this species is empty (E. Strong, pers. comm.). Description : See Dall (1889).The fol- lowing characters must be pointed out from the original description: three rows of strong tubercles, the uppermost being the largest and spiral 3 the next in size; spi- ral 2 is the smallest and is cióse to spiral 1. It is also mentioned that one specimen had 24 whorls, thus being an elongate shell. Dimensions : 15.5 mm is mentioned in the original description. Distribution: Only know from its type locality. Remarks: Apparently only I. triseri- alis has the uppermost spiral with larger nodules, but the other two spirals have the nodules rather similar in size; none of the other species here described have these characters. Anyway I. triserialis has a shorter shell. Inella compsa (Dall, 1927) (Figs. 17F, G) Triphora ( Strobiligera ) compsa Dall, 1927. Proc. U. S. N. M., 70: 96. [Type locality: Off Georgia]. Type material: Lectotype, here designated (Fig. 17F) 12.0 mm, and one paralectotype (Fig. 17G) 9.4 mm (USNM 333518). Description : See Dall (1927). Distribution : Only known from the type locality. Remarks : The description of the species seems to have been based on the characters of both lectotype and paralec- totype, in spite of the fact that they may be different species. The lectotype (Fig. 17F) is in accordance with the descrip- tion with the exception that it has not two but three prominent beaded cords with almost orthocline ribs, but the dimensions are those indicated in the original description. Ffowever it is a fragment of a shell with a broken aper- ture and lacking a protoconch, and it is therefore difficult to define exactly. The paralectotype (Fig. 17G) has only two spirals, with a third appearing on the final whorls cióse to the undulating thread near the suture, and it could be a different species. Due to the lack of additional material apart from these mentioned types, it is preferable to wait for more suitable material to study in the future. Inella pseudotorticula spec. nov. Rolán and Lee (Figs. 17H-L) Type material: Holotype (Fig. 17H) in FLMNH; a paratype (Fig. 171) in USNM (both vyJ.R.H. Light- boum, 9/8/93; ex CHL). 118 ROLÁN AND FERNÁNDEZ-GARCÉS: New data on the Caribbean Triphoridae Figure 16. Inella colon (Dalí, 1881). A: lectotype, 12.7 mm, off Havana, 823 m (USNM); B: para- lectotype, 7.8 mm, off Havana, 823 m (USNM); C: paralectotype, 8.6 mm, Cape San Antonio, 1171-1834 m (MCZ 7387); D: detail of the sculpture; E: shell, 20.3 mm, Yucatán (FLMNH); F: shell, 29.4 mm, straits of Florida (FLMNH). Figura 16. Inella colon (Dalí, 1881). A: lectotipo, 12,7 mm, fuera de La Habana, 823 m (USNM); B: paralectotipo, 7,8 mm, fuera de La Habana, 823 m (USNM); C: paralectotipo, 8,6 mm, Cabo de San Antonio, 1171-1834 m (MCZ 7387); D: detalle de la escultura; E: concha, 20,3 mm, Yucatán (FLMNH); F: concha, 29,4 mm, estrechos de Florida (FLMNH). 119 Iberus , 26 (1), 2008 Type locality: South of Castle Roads, SE Bermuda, (J.R.H. Lightboum and A.T. Guest, 1988) 100 m. Etymology: The specific ñame alludes to the similarity with I. torticula. Description : Shell (Figs. 17H, 171) very elongate, pointed, with blunt apex, not straight in profile, solid. Protoconch (Fig. 17J) with about 2 l/i whorls, with a wide nucleus and two spiral cords, of which the upper one is smaller and less prominent; diameter of about 400 pm. The teleoconch has about 23 whorls (on the holotype) with spirals 2 and 3 present from the beginning, very similar in size, and with the nodules cut in the middle. On the early whorls spiral 1 is very small and is located cióse to the upper suture, with spiral 4 appearing like a fine thread near the lower suture; on about the ninth whorl, spiral 1 is more clearly defined on the upper part of the whorl, below the suture, but is smaller and less prominent than the other two, while spiral 2 is the most prominent and slightly larger; on about the seventeenth whorl, spiral 1 contin- úes to be closer to spiral 2 (Fig. 17K). On about the twenty-first whorl, the three spirals are almost similar in size. Aper- ture (Fig. 17L) ovoid with a short siphonal canal. The shell has some white whorls alternating with entirely light brown whorls. The cords are often light brown between the nodules. Dimensions : The holotype measures 16.8 mm. Distribution: Only know from the type locality. Remarks : This species must be com- pared with the following ones that have elongate shells: Inella longissima has spirals 1 and 3 from the beginning, with spiral 2 appearing below; at the end of the spire the spirals are of almost equal strength, but spiral 2 is slightly smaller and spiral 3 a little more prominent on most of the shell. I. pseudolongissima spec. nov. has spirals 1 and 3 from the beginning, with spiral 2 smaller; on the final whorls spiral 3 is noticeably larger than the others; the colour is whitish. The proto- conch has four whorls. I. harryleei spec. nov. has spirals 1 and 3 from the beginning; spiral 2 appears after the first few whorls but remains smaller throughout the shell; at the end, spiral 3 is very large in compar- ison with the others; the colour is white or light brown with brown axial flam- mules. I. triserialis (Dalí, 1881) has a shorter shell with three spirals, spiral 1 being larger with more inflated nodules. The apex is more depressed. I. intermedia (Dalí, 1881) has a shorter shell with spirals 1 and 3 present at the beginning; spiral 2 is smaller and appears later. I. ibex (Dalí, 1881) and I. colon have only two prominent beaded cords. I. torticula (Dalí, 1881) has spiral 1 less prominent on the early whorls; sub- sequently the final whorls are wider and have up to 5 spirals. I. undebermuda spec. nov. has spirals 1 and 3 at the beginning of the teleo- conch, with spiral 2 appearing between the other two, always remaining smaller and closer to spiral 1. I. unicornium, described by Simone, 2006 from Brazil, has some general simi- larity with I. pseudotorticula, but the holotype is a shell with 40 whorls that measures almost 50 mm in height; also it is mentioned in the original descrip- tion that the protoconch has only one whorl and the teleoconch whorls have only two beaded spiral cords, with an additional smooth cord appearing on the final whorl. Inella enopla (Dalí, 1927) (Figs. 18A-E) Triphora enopla Dalí, 1927. Proc. U. S. N. M., 70: 95. [Type locality: Off Fernandina, Florida, 294 fms (538 m)]. 120 RoláN AND FERNÁNDEZ-GARCÉS: New data on the Caribbean Triphoridae Figure 17. A-E. Inella undebermuda spec. nov. A: holotype, 7.8 mm, S. of Castle Roads, Bermuda, 90 m (FLMNH); B: paratype, 12.3 mm, same locality (USNM); C: protoconch of the holotype; D: detail of the sculpture, holotype; E: detail of the aperture, holotype. F, G. Inella compsa (Dalí, 1927). F: lectotype, 12.0 mm, off Georgia, 803 m; (USNM); G: paralectotype, 9.4 mm, off Georgia, 805 m (USNM). H-L. Inella pseudotorticula spec. nov. H: holotype, 16.8 mm, SE Bermuda (FLMNH); I: paratype, 8.5 mm, S. of Castle Roads, Bermuda (USNM); J: protoconch, holotype; K: detail of the sculpture, holotype; L: detail of the aperture. Figura 17. A-E. Inella undebermuda spec. nov. A: holotipo, 7,8 mm, S. de Castle Roads, Bermuda, 90 m (FLMNH); B: paratipo, 12,3 mm, la misma localidad (USNM); C: protoconcha del holotipo; D: detalle de la escultura, holotipo; E: detalle de la abertura, holotipo. F, G. Inella compsa (Dalí, 1927). F: lectotipo, 12,0 mm, fuera de Georgia, 805 m; (USNM); G: paralectotipo, 9,4 mm, fuera de Georgia, 805 m (USNM). H-L. Inella pseudotorticula spec. nov. H: holotipo, 16,8 mm, SE Bermuda (FLMNH); I: paratipo, 8,5 mm, S. de Castle Roads, Bermuda (USNM); J: protoconcha, holotipo; K: detalle de la escultura, holotipo; L: detalle de la abertura. 121 Iberus, 26 (1), 2008 Type material: Lectotype (2 f, labeled as "type"), here designated (Figs. 18A, 18B) and 5 paralec- totypes (3 j, 2 f) (USNM 108074). Other material examined: 2 f (with the label " Triforis colon"), off Havana, Cuba 823 m (Blake Expe- dition, Sta. 243) (BMNH acc. n° 2283). Description : Dall (1927). Protoconch (Figs. 18C, 18D) with the apex smooth, pupoid, wide (about 500 pm), followed by about 2 whorls with three very cióse cords, the uppermost smaller and the lowest one the largest and most promi- nent. Teleoconch whorls with two spiral nodulous cords and another two very fine cords that are separated by the suture (Fig. 18E); the subsutural one is very small and has elongate nodules, while the one just above the suture is smooth. The nodules on the main spi- rals are pointed and cut at the middle. Dimensions : The lectotype consists of fragments measuring 7.2 and 6.6 mm. Distribution : Only known from Florida and Cuba. Remarks : The original description men- tions 15 whorls and dimensions of 11.5 x 2 mm. The glass tube labeled "type" con- tained two fragments (7.2 and 6.6 mm respectively), but there were no apparent indication thar they were previously unit- edtherefore it is doubtful, although pos- sible, that they carne from the same shell. However, they give a good representation of the specific characters, and as it is not certain whether or not they are from the same broken shell, it is better to keep both fragments together and consider both as parts of the lectotype. The large pupoid protoconch is the most important differential character. The most similar species are: I. inflata has the protoconch whorls with two scarcely visible spiral cords, the nodules of the teleoconch smaller and sharply pointed, the shell with rapidly enlarging whorls. Inella bigemma has the shell with more rapidly enlarging whorls, spiral 1 more developed and more clearly nodulous. I. ibex is not so cylindrical and has more pointed nodules, with only one small thread near the suture, lacking the small upper spiral on the final whorls. I. harryleei, I. compsa, I. pseudotorticula spec. nov. and I. intermedia have three spiral cords. I. colon has a more cylindrical shell, with spiral cords less prominent, the nodules very small, and the small cords at both sides of the suture almost imper- ceptible. I. undebermuda has three main spirals, spiral 2 being small and cióse to spiral 1; only one small thread near the suture. I. pompona has a much larger space between spiral 2 and spiral 3, the nodules are smaller and the protoconch is wider and more prominent at the apex, and has two spiral cords. Inella meteora (Dalí, 1927) (Figs. 18F-H) "Triphora" meteora Dall, 1927. Proc. U. S. N. M., 70: 95. [Type locality: Off Fernandina, Florida]. Type material: Lectotype (labeled as "type", in three fragments), here designated (Figs. 18F, 18G) (USNM 108081) and 3 paralectotypes. Description : Dall (1927). The proto- conch (Fig. 18G) is pupoid with about 2 whorls and a diameter of about 600 pm; below the nucleus there are two main smooth spiral cords and another smaller subsutural one. The whorls of the teleoconch have three spiral cords from the beginning (Fig. 18H), the lower two more similar in prominence, the nodules slightly larger on spiral 2; the uppermost is smaller from the beginning of the teleo- conch and also is less prominent on the final whorls. The nodules are uniformly rounded on spiral 1 and larger and cut at the middle on the other two. Above 122 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 18. A-E. Inella enopla (Dalí, 1927). A, B: two fragments labeled as “type”, 7.2, 6.6 mm, 538 m (USNM); C, D: protoconch of a paralectotype; E: detail of the sculpture of a paralectotype. F-H. Inella meteora (Dalí, 1927). F, G: two fragments labeled as “type” 4.0 and 5.1 mm, off Fer- nandina, 538 m (USNM); H: detail of the sculpture. Figura 18. A-E. Inella enopla (Dalí, 1927). A, B: dos fragmentos etiquetados como “tipo”, 7,2, 6,6 mm, 538 m (USNM); C, D: protoconcha de un paralectotipo; E: detalle de la escultura de un paralec- totipo. F-H. Inella meteora (Dalí, 1927). F, G: dos fragmentos etiquetados como “tipo” 4,0 y 5,1 mm, fuera de Fernandina, 538 m (USNM); H: detalle de la escultura. 123 Iberus , 26 (1), 2008 the suture there is a small undulating thread. Dimensions : See below. Distribution : Only know from its type locality. Remarks : The original description gives dimensions of 9 x 2 mm; the fragments in the tube labeled "type" measure 6.2, 5.1 and 4.0 mm; they are of the same species but probably not from the same shell. The 3 paralectotypes in the other tube appear to be a different species. The differences with the closest species are: 1. bigemma has two spirals at the beginning of the teleoconch. I. pompona and I. inflata have two spirals at the beginning of the teleo- conch and a wider, pupoid protoconch. I. longissima has two spirals at the beginning, subsequently three regular spirals of similar size, with spherical nodules, and the protoconch has three whorls. I. pseudolongissima spec. nov. has a protoconch with 4 whorls, and spirals 2 and 3 are more evident at the beginning of the teleoconch. I. harryleei has a more depressed apex on the protoconch; on the teleo- conch the lower spiral is larger, with spiral 2 smaller. I. ibex and I. enopla have only two spirals on the protoconch. I. torticula can have up to 5 spirals at the end. I. colon is more cylindrical and has only two main spirals on the teleoconch with small nodules. I. undebermuda spec. nov. has two main spirals at the beginning of the teleoconch and one smaller spiral 2 in- between, located closer to spiral 1. I. compsa is more conical, having more rapidly enlarging whorls; the nodules are less prominent and spheri- cal. I. pseudotorticula spec. nov. has a pro- toconch with a smaller diameter and with 2 well defined cords; only spirals 2 and 3 are present at the beginning of the teleoconch, spiral 1 appearing later. Inella pompona (Dalí, 1927) (Figs. 19A-C) Triphora (Strobiligera) pompona Dalí, 1927. Proc. U. S. N. M., 70: 94. [Type locality: Off Georgia]. Type material: In the material under this ñame from USNM (Figs. 19A-19C) there is a label men- tioning "type", but in the three glass tubes there is no other indication. The shell measuring almost 20 mm with 21 whorls is here designated as lectotype (Fig. 19A); it is referred to in the original description as the "larger but imperfect specimen" because the apex is missing. Numerous para- lectotypes (USNM 108339), most of them fragments (6 in one tube and 48 in another); the largest fragment measures 23.5 mm. Description : Dall (1927). The proto- conch (Fig. 19B) is pupoid and wide, the nucleus and the first whorl wider than the subsequent ones, with the diameter of the first whorl between 650 and 750 pm; in the original description it is described as having 1 1/i whorls, but it can have more than 2 whorls, the second one with two spiral cords. Teleo- conch with two main spirals (spirals 2 and 3); spirals 1 and 4 are very cióse to the suture, which separates them (Fig. 19C). Spiral 1 has very small nodules, while spiral 4 is smooth. The nodules on the main spirals are not very prominent and only a little higher than the cords, and they are slightly cut at the middle. Dimensions : The lectotype measures almost 20 mm, and some shells are somewhat larger. Distribution : Only know from the type locality. Remarks : The species with a pupoid and wide protoconch are: I. enopla has a slightly narrower pro- toconch of three whorls, with three scarcely visible spiral cords. I. ibex and I. colon each have informa- tion on the protoconch in the original description, but shells with a complete 124 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 19. A-C. Inella pompona (Dalí, 1927). A: lectotype, 19.5 mm, 804 m (USNM); B: proto- conch of a paralectotype; C: detail of the sculpture of lectotype. D-I. Inella dinea (Dalí, 1927). D: lectotype, 7.9 mm, 804 m (USNM); E-G: paralectotypes, 6.6, 6.0. 4.7 mm, same locality; H, I: protoconchs, lectotype and paralectotype. Figura 19. A-C. Inella pompona (Dalí, 1927). A: lectotipo, 19,5 mm, 804 m (USNM); B: protocon- cha de un paralectotipo; C: detalle de la escultura del lectotipo. D-I. Inella dinea (Dalí, 1927). D: lec- totipo, 7,9 mm, 804 m (USNM); E-G: paralectotipos, 6,6, 6,0, 4,7 mm, la misma localidad; H, I: protoconchas, lectotipo y paralectotipo. 125 Iberus , 26 (1), 2008 apex are not known; on the teleoconch there are similarly two spiral beaded cords, but on 1. colon the sutural area is narro wer and the nodules are smaller; I. ibex has smooth spirals at the beginning of the teleoconch. I. inflata has a similar protoconch, but clearly narrower; there is a greater increase in the width of the whorls and the suture is more prominent. I. undebermuda spec. nov. has a nar- rower protoconch; in the teleoconch. between the main spirals 1 and 3, there is a small spiral 2 closer to spiral 1. 1. dinea (see below). The species with unknown proto- conch: I. compsa has rapidly enlarging whorls and three almost similar spirals. I. sarissa has spiral 1 larger, with spiral 2 very small and cióse to spiral 1. I. bigemma has small nodules on spiral 1 and fewer whorls, never elon- gated. Inella dinea (Dalí, 1927) (Figs. 19D-I) " Triphora " pompona dinea Dalí, 1927. Proc. U. S. N. M., 70: 94. [Type locality: Off Georgia, 440 fms (805 m)]. Type material: Lectotype (Fig. 19D), here designated, and 11 paralectotypes (Figs. 19E-19G) (USNM 333517). Description : Dall (1927), comparing shells to typical I. pompona, only men- tioned that the beading is less promi- nent and the shells are more slender and compact, measuring 10 mm for 15 whorls, with a máximum diameter of 1.75 mm. Now it can be added that the protoconch (Figs. 19H, 191) has between 2 !A - 2 1 ¡2 smooth whorls and is a little variable in size, but it can reach up to 700 pm and is pupoid at the apex, with a large nucleus and with two or three depressed smooth cords on the second whorl. On the teleoconch, the suture has a smooth fine cord on each side. The axial sculpture is formed by small ribs on some shells; on others the ribs are almost absent; there are small orthocline growth lines. There are three spirals, equal in size on the lectotype but with spiral 1 less prominent on other shells. Dimensions: The lectotype measures 7.9 mm. Distribution : Rosenberg (2005) gives coordinates of 30.73° N, 79.43° W, with a depth of 805 m. Remarks: In the material studied there was no shell measuring 10 mm, but the designated lectotype has 14 whorls including the protoconch, and is almost 8 mm. This shell was chosen because the number of whorls is closest to the number given in the description. Also because it has three similar cords on each whorl, very different from T. pompona. Not all the paralectotypes seem to be the same species. Although the present taxon was pub- lished as a variety of T. pompona, compar- ison of the types of both species allowed us to confirm that they are different species. T. pompona has two nodulous cords per whorl and a very small cord, lacking nodules, on either side of the suture. T. dinea has a similar pupoid pro- toconch but the shell is variable, with some shells having three cords of similar size; other shells have only two spiral nodulous cords and the third, near the suture, is depressed. These could repre- sent characters of two different species but, as the studied material did not inelude enough complete shells, it is better to keep these two morphs within T. dinea. I. sentoma has a fíat profile, lacks nodules and is also a larger shell. I. gaesona has depressed spirals almost without nodules and the proto- conch is narrower, with a depressed apex. I. inflata has a narrower protoconch, with 2 poorly defined cords. The teleo- conch has more prominent spirals and nodules. 126 Rolan AND FernÁNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 20. A, B. Inella sentoma (Dalí, 1927). A: lectotype, 28 mm, off Fernandina, 538 m (USNM): B: paralectotype, 4.4 mm, same locality (USNM). C. Inella gaesona (Dalí, 1927), lecto- type, 9 mm, off Georgia, 805 m (USNM). D-H. Inella differens spec. nov. D: holotype, 7.6 mm, off Egmont Key, Hillsborough Co., Florida (FLMNH); E: paratype, 6.2 mm; F, G: protoconchs of holotype and paratype; H; detail of the sculpture. Figura 20. A, B. Inella sentoma (Dalí, 1927). A: lectotipo, 28 mm, fuera de Fernandina, 538 m (USNM): B: paralectotipo, 4,4 mm, la misma localidad (USNM). C. Inella gaesona (Dalí, 1927), lectotipo, 9 mm, fuera de Georgia, 805 m (USNM). D-H. Inella differens spec. nov. D: holotipo , 7,6 mm, fuera de Egmont Key, Hillsborough Co., Florida (FLMNH); E: paratipo, 6,2 mm; F, G: protocon- chas del holotipo y paratipo; H; detalle de la escultura. 127 Negro Iberus , 26 (1), 2008 Inella sentoma (Dalí, 1927) (Figs. 20A-B) Triphora sentoma Dalí, 1927. Proc. U. S. N. M., 70: 96. [Type locality: Off Fernandina, Florida, 805 m]. Type material: Lectotype, here designated (Fig. 20A), 28 mm and a paralectotype (Fig. 20B), 4.4 mm (USNM 108072). Description : See Dall (1927). The pro- toconch is unknown. Both the lectotype and paralectotype are in very poor con- dition, so the most important characters from the original description are men- tioned here: two low flattened spiral cords, faintly undulated, one at the periphery of the whorl, the other in front of it; the suture linear with a minute threadlet on each side of it, the interspaces shallow. In reality it can be said that there are grooves and orthocline growth lines, but no prominent sculpture. Dimensions: The lectotype measures 28 mm. Distribution : Only know from its type locality. Remarks : This species is very elongate and with depressed sculpture. There is no Information about the protoconch or aper- ture, but in the original description it is mentioned that one of the fragments had a protoconch of a whorl and a half. For this reason and the almost cylindrical shell form it is included in the genus Inella. The paralectotype (Fig. 20B) is more similar to the lectotype of I. gaesona (see below). No other Inella species in the studied area has such depressed sculpture. Inella gaesona (Dall, 1927) (Figs. 20C) Triphora (Strobiligera) gaesona Dall, 1927. Proc. U. S. N. M., 70: 95. [Type locality: Off Georgia, 805 m]. Type material: Lectotype, here designated (Fig. 20C) with 16 whorls, 9 mm (USNM 1088341). Para- lectotypes: 15 s and f (not examined) (USNM). Description: See Dall (1927). Proto- conch paucispiral, pupoid and de- pressed, with the first whorl wider than the following one. The spiral sculpture is formed more by grooves which sepá- rate the whorls into three parts than by prominent spirals. The axial sculpture consists only of orthocline growth lines. Dimensions: The lectotype measures 9 mm. Distribution: Only know from its type locality. Remarks: Species very different from any other due to the pupoid and depressed protoconch and the fíat spiral cords without nodules. It only has some similarity to I. sentoma, but a complete comparison could not be made because of the poor condition of the lectotype of this species and the lack of sufficient material. For this reason they are kept as different valid taxa, awaiting more material for study. Inella differens spec. nov. Rolán and Lee (Figs. 20D-H) Type material: Holotype (Fig. 20D) in FLMNH. Paratype (Fig. 20E) in BMSM (both ex CHL). Other material examined: Off Louisiana: 3 f without protoconch, 28° 05.61' N, 91° 02.245' W by 28° 05.524' N, 91° 02.036' W, 58 m (E. F. Garda/ Pelican 06/09/00) (CHL). Type locality: Off Egmont Key, Hillsborough Co., Florida, USA, (J. Moor e/Caualier, 1962), 135 m. Etymology: The specific ñame alludes to the shell having different characters from others in the area. Description: Shell subcylindrical toconch (Figs. 20F, 20G) pupoid with 1 (Figs. 20D, 20E) solid, light brown. Pro- 3A -2 whorls and a diameter of about 128 ROLÁN and FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 21. A-D. Inella slapcinskyi spec. nov. A, B: holotype, 2.6 mm (FLMNH); C: protoconch of the holotype; D: detail of the fragment (ANSP). E-I. 1: Inella faberi spec. nov. E: holotype, 3.7 mm; F, G: paratypes, 4.4 and 3.0 mm. Louisiana (FLMNH); H, I: protoconch and detail of teleo- conch sculpture, paratype in figure F. Figura 21. A-D. Inella slapcinskyi spec. nov. A, B: holotipo , 2,6 mm (FLMNH); C: protoconcha del holotipo ; D: detalle del fragmento (ANSP). E-I. 1: Inella faberi spec. nov. E: holotipo, 3.7 mm; F, G: paratipos, 4.4 y 3.0 mm. Louisiana (FLMNH); H, I: protoconcha y detalle de la escultura de la telo- concha, paratipo de la figura F. 129 Iberus, 26 (1), 2008 600 pm, the nucleus smooth and the whorls with two smooth spiral cords. The teleoconch begins when the two spiral cords become nodulous; these beaded spirals are equal in size and both are closer to the suture and more widely separated in the middle of the whorl. About the fourth or fifth whorl, spiral 2 appears below and very cióse to spiral 1, beginning as a fine thread undulating beneath the nodules (Fig. 20H). Immediately it becomes nodulous but with a smaller nodular size. About the eleventh whorl the nodules reach almost the size of those on spirals 1 and 3. There are three additional smooth cords on the base. Aperture ovoid elon- gate; siphonal canal short and open. Dimensions : The holotype measures 7.6 mm. Distribution: Only know from its type locality. Remarks : This species is kept in the genus Inella due to the characteristic protoconch. It may be differentiated from most of the other closely similar species by its short protoconch with only 1 3A - 2 whorls and its brown colour: I. dinea has more weakly sculptured whorls and the protoconch has a wider pupoid form. 1. meteora, I. undebermuda spec. nov., I. enopla and I. pompona have a similar number of protoconch whorls but the apex is more inflated and pupoid, and the shells are larger and white; furthermore, I. meteora has three spiral cords on the teleoconch, spiral 1 being smaller; I. undebermuda has spiral 2 beginning on about the tenth whorl (instead of on the fourth in I. differens), and it is very small and closer to spiral 1; I. enopla and I. pompona have two main spiral cords on the teleoconch with an additional cord on each side of the suture, the proto- conch being very much wider. I. triserialis has a rather similar shell but the protoconch has 3 whorls, the apex is more depressed, the nodules of spiral 1 are larger and cut at the middle. I. sarissa is white, spiral 2 is smaller and very cióse to spiral 1. I. compsa has rapidly enlarging whorls, and spiral 2 is not smaller. I. intermedia has a larger shell that is lighter, whitish and more conical, the spirals almost equal in size. See also I.faberi below. Inella slapcinskyi spec. nov. Rolán and Fernández-Garcés (Figs. 21A-D) Type material: Holotype (juvenile, Figs. 21A, 21B) (FLMNH UF350382). Other material studied: 1 f (ANSP). Type locality: Cayman Islands, Little Cayman, Jackson's Bight (P. Watson, Dec/1992), 35 m. Etymology: The species is named after John Slapcinsky, Curator of Molluscs of the FLMNH, for his help in the examination of the material housed in this museum. Description: Shell (Figs. 21A, 21B) small, almost cylindrical, whitish. Proto- conch paucispiral (Fig. 21C) with a diameter of about 400 pm and almost three whorls, with two narrow spiral cords on the first whorl, three on the fol- lowing, well defined, the lower one more prominent; each spiral cord bears a row of small tubercles on its upper surface. The nucleus of the protoconch is very small. The teleoconch has four spirals; spiral 1 has more prominent nodules; spiral 4 is the next in size and spirals 2 and 3 have smaller, elongated nodules (Fig. 21D). The axial ribs are wide and strongly prosocline. On the base there are two more smooth spirals. Aperture rounded with a strong col- umellar callus; siphonal canal short and open. Dimensions: The holotype measures 2.6 mm. Distribution : Only known from its type locality. Remarks : This species has been named in spite of the scarcity of avail- able material because the shell and the protoconch have characters that are absolutely different from any other species of this group in the study 130 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 22. Sagenotriphora osclausum (Rolán and Fernández-Garcés, 1995). A-C: shells, 4.9, 4.9 and 4.8 mm, E Palm Beach, Florida (CHL); D-F: protoconchs; G-H: radula, specimen from off Palm Beach, Florida (CHL); I: operculum. Figura 22. Sagenotriphora osclausum (Rolán y Fernández-Garcés, 1995). A-C: conchas, 4,9, 4,9 y 4.8 mm, E Palm Beach, Florida (CHL); D-F: protoconchas; G-H: rádula, de un ejemplar de fuera de Palm Beach, Florida ( CHL); L: opérculo. 131 Iberus, 26 (1), 2008 area. Only I. torticula has 4 beaded area has a row of tubercles on the pro- spirals, and no species in the study toconch. Inella faberi spec. nov. Rolán and Fernández-Garcés (Figs. 21E-I) Type material: Holotype (Fig. 21E) and 2 paratypes (Figs. 21F, 21G) in FLMNH (UF 291343). Type locality: Louisiana, 28.05731° N, 92.44963° W, 71-74 m. Etymology: The species is named after the Dutch malacologist Marien Faber, who studied many of the species of Caribbean Triphoridae and helped us in some aspects of this work. Description: Shell (Figs. 21E-21G) subcylindrical, solid, brownish in colour. Protoconch (Fig. 21FF) with almost three whorls which have two narrow well defined spirals, the lower of which is a little more prominent; its diameter is about 450 pm. The teleo- conch begins with two nodulous spirals 1 and 3, quite cióse together; spiral 1 is slightly smaller and less prominent than spiral 3. Subsequently spiral 2 appears between them and remains smaller (Fig. 211). The axial ribs which connect the nodules are strongly proso- cline. The nodules are different on the three spirals: on spiral 1 they are spheri- cal and a little ovoid, on spiral 2 they are very elongate and narrow, and on spiral 3 the nodules are larger and cut by the spiral which crosses them. Aper- ture rounded but in poor condition on all the type material. Dimensions: The holotype measures 3.7 mm. One paratype reaches 4.4 mm. Distribution : Only known from the type locality. Remarks: The most similar species are: I. differens spec. nov. has a wider pro- toconch, apically more depressed and with only 2 whorls, the nodules of the teleoconch are all spherical and the axial ribs almost orthocline. I. undebermuda spec. nov. has a proto- conch with only 2 whorls, the nucleus more elevated; spiral 2 on the teleo- conch is closer to spiral 1 and the axial ribs are slightly prosocline. I. apexbilirata spec. nov. has cords of sim- ilar size on the protoconch, the shell is whitish, and on the teleoconch spiral 1 has very small nodules, spirals 2 and 3 have larger nodules which are cut at the middle, and the axial ribs are almost orthocline. I. harryleei spec. nov. has a very elon- gate shell, a protoconch with cords of similar size, and the teleoconch has spiral 3 with much larger and more prominent nodules. Genus Sagenotriphora Marshall, 1983 Type species (by original designa tion): Triphora and northern New Zealand. Description : Protoconch multispiral with reticulate sculpture on the first whorl; radula with a rachidian tooth impulla Hedley, 1903. Recent, Southern Australia which is separated into two parts, and only one lateral and one marginal. Sagenotriphora osclausum (Rolán and Fernández-Garcés, 1995) (Figs. 22A-I) " Triphora " osclausum Rolán and Fernández-Garcés, 1995. Apex, 10(1): 21. Type material: Holotype and paratypes (see Rolán and Fernández-Garcés, 1995) Other material examined: Florida: 3 s, beach drift, Little Torch Key, Monroe Co., (M. Teskey, 5/76) (CHL); 2 sp, 32 miles E St. Augustine, St. Johns Co. (T. Yocius 6/81; ex J. Dawley 12/2/90), 30 m 132 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 23. A-F. Sagenotriphora candidula spec. nov. A, B: holotype, 5.0 mm (FLMNH); C: paratype 3.9 mm, from type locality (BMSM); D: protoconch of the holotype; E: operculum; F: drawing of the radula. G, H. “Triphora” abrupta (Dalí, 1881); G: lectotype, 7.9 mm (MCZ 7389); H: Detail of the spire. I-K. “Triphora caracca Dalí, 1927. I: lectotype, 7.1 mm, off Georgia, 805 m (USNM): J, K: paralectotypes, 6.9, 6.7 mm, same locality (USNM). Figura 23. A-F. Sagenotriphora candidula spec. nov. A, B: holotipo, 5, 0 mm (FLMNH); C: paratipo 3,9 mm, de la localidad tipo (BMSM); D: protoconcha del holotipo; E: opérculo; F: dibujo de la rádula. G, H. “Triphora” abrupta (Dalí, 1881); G: lectotipo, 7,9 mm (MCZ 7389); H: Detalle de la espira. I-K. “Triphora” caracca Dalí, 1927. I: lectotipo, 7,1 mm, fuera de Georgia, 805 m (USNM): J, K: paralectotipos, 6,9, 6,7 mm, la misma localidad (USNM). 133 Iberus, 26 (1), 2008 (CHL); 5 sp (Figs. 22A-22C)(one destroyed by radular study), E Palm Beach (J. Root; ex K. Sunder- land 1991) 30 m (CHL); 5 s, SW Egmont Key, 73-92 m (Steger, 1966) (ANSP 306356); 2 s, Monroe Co. Dry Tortugas area, 25 30' N, 59 m (FLMNH 154901); 3 s, Palm Beach Co., off Singer Id. (McGinty/Jun 1940) 183-274 m (FLMNH 257141); 2 s, Palm Beach Co., off Palm Beach (McGinty/Jan 1950) 92 m (FLMNH 205145); 8 s, Palm Beach Co., reefs (D. Akers/ 1970-1 971) 30 m (FLMNH 127829); 15 sp, Palm Beach Co., off Delray Beach, Manalapan Wreck (McGinty), 10 m (FLMNH 249896); 1 s, Palm Beach Co., off Yamato Rocks (F. B. Lyman/ Apr 1939) 11 m (FLMNH 10242). Bahamas: 2 s. Iridian Cay, Grand Bahama Island 26° 42' 45"N, 78° 39' 15"W (Worsfold) (ANSP 366953). Description : Rolán and Fernández- Garcés (1995). Shell (Figs. 22A-22C) light brown with the protoconch darker. Radula (Figs. 22G, 22H): With the formula 4-1-1-1-4. Central tooth divided in two parts by a deep separation. Each of these two prominences finishes in four short sharp pointed cusps. Lateral teeth elongate and with the form of a half tube, the end curved with 6-7 pointed and short cusps. Four marginal teeth in each side, fíat, elongate, and at their ends are about 15-17 very fine and elongate cusps, which are shorter on one side and more elongate on the other. Dimensions: The holotype measures 4.5 mm. Some shells can reach up to 5.0 mm. Distribution : Known from Florida, Bahamas and Cuba. Remarks : This species was described without a generic assignment, as no soft parts were studied. With the present material it was possible to study the radula from dried animáis retracted into the shell. This radula proved to be very different from all those previously known, and for this reason the creation of a new genus was considered, but it seemed preferable to place the species provisionally in a known genus that had the most similar radula. In this case we found that the genus Sagenotriphora is almost the only one in which the rachid- ian tooth is divided into two parts, although it has only one wide lateral and one marginal, contrary to the four that are found in the present species. Sagenotriphora osclausum had been considered by some malacologists (the label of most of the studied material) to be Triphora dupliniana Olsson, 1916. This taxon is a fossil species and it may be differentiated by the shorter shell and the open siphonal canal. Sagenotriphora candidula spec. nov. Rolán and Lee (Figs. 23A-F) Type material: Holotype (Figs. 23A, 23B) in FLMNH. Paratypes: 1 s (Fig. 23C) in BMSM; 1 s, in USNM; all from the type locality and ex CHL. Other paratypes: 3 s, off Anna Maria Island, Manatee Co., Florida (J. Moore /Cavalier, 1962), 15-30 m (CHL); 4 s, SW of Egmont Key, Florida (Steger, 1996) 40-50 m (ANSP 306346); 10 s, East Tampa Bay, Florida (C. L. Richardson) (ANSP 335494); 5 s, Palm Beach Co., Florida (McGinty, Jul/1941) 55-73 m (FLMNH 249739); 2 s, Hillsborough Co., W of Egmont Key, Florida (J. Moore, 1962) 183 m (FLMNH 249838); 1 s, Palm Beach Co., reefs. Florida (D. Akers, 1970-1971) 30 m (FLMNH 127829); 13 sp, Palm Beach Co., off Delray Beach, Manalapan Wreck, Florida (McGinty), 10 m (FLMNH 249896); 1 s, Palm Beach Co., off Singer Island, Florida (McGinty, Jun/1940) 30 m (FLMNH 178388); 1 s, Palm Beach Co., Breakers Hotel, (McGinty, Jun/1940) 30 m (FLMNH 154860). Other material studied: Florida: 7 s and f, SW of Egmont Key (Steger, 1996) 40-50 m (ANSP); 2 sp (destroyed for radular studies), type locality (CHL); 11 s, in poor condition, East Tampa Bay (C. L. Richardson) (ANSP 335494); 2 s, in poor condition, St. Augustine (FLMNH 286017). Type locality: beach drift. Captiva Island, Florida. Etymology: The specific ñame is the Latín word candidula which means "shining white" (diminu- tive) and alludes to the colour of the shell. Description : Shell (Figs. 23A-23C) elon- the whorls with two spiral threads crossed gate, solid. Protoconch (Fig. 23D) of almost by small undulating axial ribs. The begin- 4 whorls, apex with rounded tubercles and ning of the teleoconch has two spirals, spi- 134 ROLÁN and FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 24. A-E. “ Triphora” georgiana Dalí, 1927. A: lectotype, 7.5 mm, off Georgia, 805 m (USNM); B-E: paralectotypes, 6.0, 4.9, 5.0, 4.1 mm, same locality (USNM). F, G. “Triphora” indígena Dalí, 1927. F: lectotype, 6.6 mm, off Georgia, 805 m (USNM): G: paralectotype, 3.3 mm, same locality (USNM). H. “ Triphora” lilacina, lectotype, 8.5 mm, Turtle Harbor, Florida, 11 m (USNM 83087). I-O. “Triphora” cf. lilacina (Dalí, 1889). I: shell, 7.5 mm, Pinellas Co., Florida (FLMNH); J: shell, 9.0 mm, Monroe Co., Florida (FLMNH); K: shell, 8.0 mm, Palm Beach Co., Florida (FLMNH); L: shell, SEM photo, 4.1 mm, Pickles Reef, off Key Largo, 5 m (CHL); M: apical part with protoconch, shell in figure J; N: protoconch of shell in Figure L; O: detail of the last whorl from shell in Figure I. Figura 24. A-E. “Triphora” georgiana Dalí, 1927. A: lectotipo, 7,5 mm, fuera de Georgia, 805 m (USNM); B-E: paralectotipos, 6,0, 4,9, 5,0, 4,1 mm, de la misma localidad (USNM). F, G. “Triphora” indigena Dalí, 1927. F: lectotipo, 6,6 mm, fuera de Georgia, 805 m (USNM): G: paralec- totipo, 3.3 mm, la misma localidad (USNM). H. “Triphora” lilacina, lectotipo, 8,5 mm, Turtle Harbor, Florida, 11 m (USNM 83087). I-O. “Triphora” cf. lilacina (Dalí, 1889). F. concha, 7,5 mm, Pinellas Co., Florida (FLMNH); J: concha, 9,0 mm, Monroe Co., Florida (FLMNH); K: concha, 8,0 mm, Palm Beach Co., Florida (FLMNH); L: concha, SEM photo, 4,1 mm, Pickles Reef, fuera de Cayo Largo, 5 m ( CHL); M: parte apical con protoconcha, concha de la Figura J; N: protoconcha de la concha de la Figura L; O: detalle de la última vuelta de la concha de la Figura L. 135 Iberus, 26 (1), 2008 ral 1 very cióse to the suture and spiral 3 in the middle of the whorl. About the fourth- fifth whorl spiral 2 appears in the middle of the space between the other two, enlarg- ing quickly and becoming equal to spirals 1 and 3 on the subsequent two whorls. A very fine spiral thread appears at the suture. Aperture oval elongate. Siphonal canal very short, closed by the continua- tion of the outer lip. Colour: protoconch brown and the teleoconch uniformly creamy-white; small areas of cream or light brown may be visible on the base and inside the aperture. Operculum multispiral (Fig. 23E) ovoid, yellowish, translucent. Dimensions : The holoype measures 5.0 mm. Distribution : Known from Florida coasts. Due to the scarcity of available mater- ial, our radular study was limited to only two specimens. Unfortunatly, only a little information could be obtained and the radula could not be photographed (see Fig. 23F). The radula (formula 3-1-1-1-3) has a central tooth with two upward cusps and two more externally and in the opposite direction. The lateral teeth have a sharp border wíth many small cusps. The marginal has only a few cusps. Remarks: This species had been con- siderad by collectors to be conspecific with the previous one (S. osclausum), which is similar but light brown in colour. However, the difference in the radula indi- cated that they were different species. S. candidula spec. nov. must be differ- entiated from the white Cosmotriphora mela- nura (C. B. Adams, 1850) which is larger, with more whorls. Spiral 2 appears earlier(3-4 whorls), the colour of the teleo- conch is milk white and not creamy-white, and the base is white; the apex of the pro- toconch is narro wer; also the protoconch is consistently dark brown and not light brown, has only one spiral on the first and second whorls and is also a little larger. Marshallora ostenta spec. nov. is not totally cream, and some spiral cords (at least the suture) may be brown or light brown; the siphonal canal is short and open and the protoconch is somewhat larger, and has one spiral cord on the first protoconch whorl. Genus Triphora Blainville, 1828 Type species (by monotypy): Triphora gemmatum Blainville, 1828; Recent, Mauritius. Remarks: This genus ñame has is commontly used when generic assig- formerly been employed as the ment is not possible, as " Triphora " sensu nominal for the family. Consequently, it lato. "Triphora" hircus (Dalí, 1881) (Fig. 36J) Triforis hircus Dalí, 1881. Bull. Mus. Comp. Zool., 9: 83. [Type locality: Yucatán Strait, 640 fms (1171 m)]. Triforis (bigemma var.?) hircus Dalí, 1889. Bull. Mus. Comp. Zool., 18: 249, pl. 20, fig. 11. Type material: No type in USNM (E. Strong, pers. corran.); neither in other American museums. Figured in Dall (1889: pl. 20, fig. 11) (Fig. 36f) measuring 12.5 mm. Description: See Dall (1881). With 14 or more inflated whitish whorls. The most important characters in the origi- nal description and in the figure are the two prominent spirals, the upper one more widely separated from the suture. Dimensions: The original description gives a dimensión of 12.5 mm. Distribution: Only known from the type material. Remarks: Dall (1881) described this species as new; later (Dall, 1889) he revised his taxonomic opinión and he considerad it to be probably a variety of T. bigemma Watson, 1880; but that species has only two spirals on the upper part of 1 36 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 25. A. Triforis atlántica E. A. Smith, 1890, lectotype, 6 mm, St. Helena. B-I. “ Triphora atlántica. B: shell, 6.3 mm, W Egmont Key, Hillsborough Co., Florida (CHL); C, D: shells, 7.1, 4.9 mm off Guarapari, Espiritu Santo State, Brazil, 15-20 m (CHL); E, G, H: shells, 6.7, 5.3, 4.2 mm, 42 miles E St. Augustine, St. Johns Co., Florida, 85 m (CHL); F: shell, 4.8 mm, SEM photo, Florida (FLMNH); I: protoconch of the shell in previous figure). J. “Triphora” pyrrha Henderson and Bartsch, 1914, syntype, 2.7 mm, Chincoteague (USNM). Figura 25. A. Triforis atlántica E. A. Smith, 1890, lectotipo, 6 mm, St. Helena. B-I. “Triphora” atlántica. B: concha, 6,3 mm, O Cayo Egmont, Hillsborough Co., Florida (CHL); C, D: conchas, 7,1, 4.9 mm fuera de Guarapari, Estado de Espiritu Santo, Brasil, 15-20 m (CHL); E, G, H: conchas, 6,7, 5,3, 4,2 mm, 42 miles E St. Augustine, St. Johns Co., Florida, 85 m (CHL); F: concha, 4,8 mm, SEM photo, Florida (FLMNH); I: protoconcha de la concha de la figura anterior). J. “Triphora” pyrrha Henderson y Bartsch, 1914, syntype, 2,7 mm, Chincoteague (USNM). 137 Iberus , 26 (1), 2008 the teleoconch, subsequently with three spirals of which spiral 1 is smaller (Watson, 1886, pl. 43, fig. 6). For these reasons both taxa have been considered here to be valid and different species. This species has been kept here in "Triphora" instead of in Inella due to several characters found in the original description: acutely tapered apex proba- bly rather pointed, crowded transverse sculpture (twenty-one slightly oblique ribs, etc.), and also due to the figured shape of the species. Comparison must be made with the species having 2 main spirals, basing the differences on the teleoconch characters: I. etiopia and I. pompona are more cylindrical and have two small spirals besides the two main ones. I. colon and I. ibex have more cylin- drical shells, and the two spirals are located equidistan! between the sutures. "Triphora" cylindrella (Dalí, 1881) (Fig. 36K) Triforis cylindrella Dalí, 1881. Bull. Mus. Comp. Zool, 9: 83. [Type locality: Cape San Antonio, 640 fms (1171 m)]. Triforis (Sychar) cylindrella Dalí, 1881. In Dall, 1889. Bull. Mus. Comp. Zool, 18: 250, pl. 20, fig. 6. Type material: No type in USNM (E. Strong, pers. corran.), AMNH (M. Sidall, pers. corran.), or ANSP. The lectotype, here designated, (Fig. 36K) is the specimen figured by Dall (1889, píate 20, fig. 6). Description : See Dall (1881). Shell with 12-14 white whorls. The most important characters are the presence of three similar spirals and convex whorls, the lo west with more elongate nodules. Dimensions: The original description gives a dimensión of 6.5 mm. Distribution : Only known from Cape San Antonio. Remarks : The placement of this species in the genus "Triphora" s. I is tentative and based on the original description and fig- ure. As the type was not found, the com- parison was made from the characters in the original description with the species which had three equal spirals, as follows: I. intermedia has whorls with a fíat profile, only 2 spirals at the beginning, and spiral 2 is smaller on most of the Shell. I. torticula has convex whorls, but the nodules are larger on spirals 2 and 3, and the shell is more elongate and cylindrical. I. compsa has an unknown protoconch, so comparison is difficult; however, spiral 1 is smaller and the nodules are more prominent. Cosmotriphora melanura is more elon- gate and the profile of the whorls is fíat. Sagenotriphora candidula has only two spirals at the beginning of the teleo- conch and the whorls are rather fíat. "Triphora" rushii (Dall, 1889) Triforis rushii Dall, 1889. Bull. Mus. Comp. Zool., 18: 246. [Type locality: west of North Bimini Island, Bahamas, 200 fms (366 m)]. Type material: Holotype (USNM 61227) missing (Ellen Strong pers. comm.). Description : See Dall (1889: 246). Shell with 12-14 whorls. Four spirals, of which only the lowermost is prominent, with a row of large pointed nodules. Above this are three much finer, lower and undulating threads of equal size. No ribs exist. Shell milky-white in colour. Dimensions : 2.6 mm according to the original description. Remarks : No other species in the studied area shows this kind of sculp- ture with four spirals, the lowermost the most prominent and with the largest nodules. 138 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 26. A-D. “ Triphora ” inaudita spec. nov. A: holotype, 5.4 mm, SW of Dry Tortugas (FLMNH). B: paratype, 4.4 mm (BMSM). C: details of protoconch of the holotype; D: detail of the sculpture. E-I. “Triphora” pseudonovem spec. nov. E: holotype, 7.3 mm (FLMNH); F: detail of the aperture of a paratype (FLMNH); G: protoconch; H: detail of the protoconch; I: detail of the spire of the holotype. Figura 26. A-D. “Triphora” inaudita spec. nov. A: holotipo, 5,4 mm, SO de Dry Tortugas (FLMNH). B: paratipo, 4,4 mm (BMSM). C: detalle de la protoconcha del holotipo; D: detalle de la escultura. E-I. “Triphora” pseudonovem spec. nov. E: holotipo, 7,3 mm (FLMNH); F: detalle de la abertura de un paratipo (FLMNH); G: protoconcha; H: detalle de la protoconcha; I: detalle de la espira del holotipo. 139 Iberus, 26 (1), 2008 "Triphora" abrupta (Dalí, 1881) (Figs. 23G-H, 36M) Triforis (bigemma Watson varj abruptus Dalí, 1881. Bull. Mus. Comp. Zoo/., 18: 84. [Type locality: Yucatán Strait, 640 fms (1171 m)]. Triforis (Sychar) abrupta Dalí, 1881. In Dall, 1889. Bull. Mus. Comp. Zool., 18: 249, pl. 20, fig. 12b. lectotype (Figs. 23G, 23H) (MCZ 7389). The types in Type material: A syntype, here designated as USNM are lost (E. Strong, pers. comm.). Description : See Dall (1881). The lec- totype is a shell without protoconch and aperture. The most important characters are that spiral 1 is formed by small, less prominent nodules; spirals 2 and 3 are more prominent and almost equal, with larger nodules. Below there is a small, smooth spiral 4. Dimensions : according to Dall (1881) the shell measures 7.5 mm. The lecto- type measures 7.9 mm. Distribution : Only known from the type material. Remarks : The lectotype is probably the same shell studied by Dall with the loss of the external lip, with the same size and sculpture. For this reason it is considered as lectotype. No information on protoconch and aperture are avail- able, which makes any kind of compari- son difficult. The placement in "Triphora'' and not in Inella is tentative, based on the rapidly enlarging whorls. There are only a few species with spiral 1 smaller, spirals 2 and 3 larger and similar, and spiral 4 smooth: Inella pompona and I. enopla have shells lighter in colour, more elongated, lacking spiral 1 on the first teleoconch whorls, and with the space between the main spirals more depressed. Inella apexbilirata spec. nov. has a smaller shell, narrower and almost cylindrical in profile. "Triphora" caracca Dall, 1927 (Figs. 23I-K) Triphora caracca Dall, 1927. Proc. U. S. N. M., 70: 93. [Type locality: Off Georgia, 440 fms (805 m)]. Type material: Lectotype, here designated (Fig. 231) and paralectotypes (Fig. 23J, 23K) (USNM 108343). Description : See Dall (1927). Dark brown paucispiral protoconch with at least three whorls with two spiral cords. No more details can be known because of erosión. The most important differen- tial character is that the axial ribs are very fine and strongly prosocline. Spirals 1 and 3 larger and prominent from the beginning of the teleoconch, with spiral 2 appearing about the tenth whorl and remaining very small on the final whorls. Dimensions : The original description gives a dimensión of 7.1 mm. Distribution: Only known from the type material. Remarks: The shell mentioned by Dall (1927) measured 8 mm, but the lectotype examined from USNM only reached 7.1 mm, which could be due to the loss of one whorl. No other triphorid in the Caribbean has spiral 2 beginning on the tenth whorl and remaining small and cióse to spiral 1 down to the end of the spire. "Triphora" georgiana Dall, 1927 (Figs. 24A-E) Triphora (Biforina) georgiana, Dall, 1927. Proc. U. S. N. M.r 70: 93 [Type locality: Off Georgia]. Type material: Lectotype, here designated (Fig. 24A) with 15 whorls and 5 paralectotypes (Figs. 24B-24E) (USNM 333516). 140 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 27. A-F. “Triphora” calva (Faber and Moolenbeek, 1991). A: shell, 3.7 mm, Key Largo, Florida (CMK); B: protoconch of the previous shell; C: shell, 3.7 mm, Key Matias, Cuba (MHNS); D: protoconch, Cienfuegos, Cuba (CFG); E: protoconch, Santa Lucia, Pinar del Río, Cuba (CFG); F: protoconch, Abaco, Bahamas (CCR); G: “ Triphora ” cf. calva, shell, 6.8 mm, Grand Bahama Island (ANSP). H, I. “Triphora” yociusi spec. nov. H: holotype, 3.9 mm, off St. Augustine, St. Johns Co., Florida (FMNH); I: protoconch of the paratype. Figura 27. A-F. “Triphora” calva (Faber and Moolenbeek, 1991). A: concha, 3,7 mm, Cayo Largo, Florida (CMK); B: protoconcha de la concha anterior; C; concha, 3,7 mm. Cayo Matías, Cuba (MHNS); D: protoconcha, Cienfuegos, Cuba ( CFG); E: protoconcha, Santa Lucia, Pinar del Río, Cuba (CFG); F: protoconcha, Abaco, Bahamas (CCR); G: “Triphora” cf. calva, concha, 6,8 mm, Grand Bahama (ANSP). H, L. “Triphora” yociusi spec. nov. H: holotipo, 3,9 mm, fuera de St. Augus- tine, St. Johns Co., Florida (FMNH); L: protoconcha del paratipo. 141 Iberus , 26 (1), 2008 Description: See Dall (1927). It is rele- vant that the protoconch was described as having a smooth nucleus and 2 more whorls sculptured with two spiral threads and numerous "flexuous axial threadlets". Probably all the protoconchs are eroded, so the exact form and sculpture could only be known after the examination of fresh shells. The teleoconch has spirals 2 and 3 from the beginning; later a very weak spiral 1 appears below the suture, and remains the smallest one up to the end. Spiral 2 is the largest and most prominent. Dimensions : The lectotype measures 7.5 mm. Distribution: Only known from the type material. Remarks : Marshall (1983) consid- ered the genus Bif orina to be a synonym of Monophorus Granata-Grillo, 1877. As there is no information on the radula, it seems better to keep it in "Triphora" in sensu lato. The shell designated as lectotype had been separated from the rest of the lot in a different capsule with the indica- tion "type". This shell has the dimen- sions and number of whorls given by the author. Because no holotype was mentioned in the original description, this shell must be designated as lecto- type. This species shows some resem- blance to Strobiligera brychia (Bouchet and Guillemot, 1978) (Fig. 14F) in its teleoconch whorls, but apparently the protoconch of the latter species is more pointed and has more whorls. "Triphora" indígena Dall, 1927 (Figs. 24F-G) Triphora indígena Dall, 1927. Proc. U. S. N. M., 70: 93. [Type locality: Off Georgia, 805 m]. Type material: Lectotype, here designated (Fig. 24F) with 13 whorls, and a paralectotype (Fig. 24G) (USNM 108079). Description : See Dall (1927). Proto- conch multispiral, a little eroded on the paralectotype but apparently brown with two spiral cords. The teleoconch begins with three almost similar spirals, with spiral 1 becoming smaller on subsequent whorls, appearing as a small thread very cióse to the suture. The cords and ribs are rather elevated, and cross to form rectangular inter- spaces. Dimensions : The lectotype measures 6.6 mm. Distribution: Georgia and Fernandina (mentioned in the original description). Remarks : Its most important charac- ter is the formation of rectangular spaces between cords and ribs. Spiral 1 is smaller throughout the spire, the other two being similar to each other. There are no similar species in the study area. "Triphora" lilacina (Dall, 1889) (Fig. 24H) Triforis lilacina Dall, 1889. Bull. M. C. Z., 18: 243. [Type locality: Turtle Harbor, Florida, USA]. Type material: One syntype (Fig. 24H) here designated as lectotype (USNM 83087). Description : See Dall (1889). The pro- toconch is unknown. The most important character is the uniform lilac colour; also, the two spirals, with an additional very narrow spiral only appearing between them on the eleventh whorl. The nodules are spherical and relatively small. Dimensions : The lectotype measures 8.5 mm. Remarks: The lectotype does not have exactly the characters mentioned in the original description (it measures 8.5 mm instead of 9.0 and has 15 whorls instead of 18), but these differences may be the 142 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 28. A-D. “ Triphora ” guadaloupensis spec. nov. A-C: holotype, 2.3 mm, Guadaloupe (ANSP); D: drawing of the protoconch. E-H. “Triphora” amicorum spec. nov. E, F: holotype, 4.4 mm, Florida (FLMNH); G: paratype, 3.8 mm. Florida (FLMNH); H: protoconch of the holotype. Figura 28. A-D. “Triphora” guadaloupensis spec. nov. A-C: holotipo, 2,3 mm, Guadalupe (ANSP); D: dibujo de la protoconcha. E-H. “Triphora” amicorum spec. nov. E, F: holotipo, 4,4 mm, Florida (FLMNH); G: paratipo , 3,8 mm. Florida (FLMNH); H: protoconcha del holotipo. 143 Iberus , 26 (1), 2008 result of a quick examination or due the loss of a small part of the shell (the loss of the first whorls of the protoconch was already mentioned in the original description). In spite of this, the colour is very characteristic and there is no doubt that it is the type mentioned in the original description. Besides, it is necessary to point out that the shell has a defect on the penultimate whorl, which may cause a slight deformation. The shells illustrated by some authors under this ñame (Vokes and Vokes, 1983) seem to be very different. Distribution : Only known from the type locality. Mesophora novem (Nowell-Usticke, 1969) seems to be the closest species, but has a different colour, larger and lighter nodules, and a darker suture; spiral 2 does not appear until the final whorls and is always smaller than the other two, but it has larger nodules than those on spiral 2 of "T." lilacina. No shells resembling the type of this taxon have been collected by any other author in spite of the fact that it comes from shallow water (11 m). Henee, some malacologists (pers. comm.) have con- sidered that this type could be an abnor- mal individual from a population which is usually differently coloured (which we will mention below as "Triphora" atlántica). Another possibility is the one sug- gested by several persons (pers. comm.): if the lectotype is an abnormal shell with an uncommon colour for the species, it may be conspecific with shells from other populations having different morpholog- ical form and colour. One candidate for this is presented below as "T." cf. lilacina. "Triphora" cf. lilacina (Figs. 241-0) Material examined: Florida: 1 s (Fig. 24J), 24° 50' N, Monroe Co., West Coast of Florida (Jun 1961) (McGinty coll.) 61 m (FLMNH 259042); 6 s (Fig. 241) Pinellas Co., SSW John's Pass (Dan Steger) 55 m (FLMNH 238675); 1 s (Fig. 24L), Pickles Reef, off Key Largo, Monroe Co. (L. Hill, May/76), 7-10 m (CHL); 1 s, Palm Beach Co., Breakers Hotel to Biltmore Hotel (R/V Tritón, Apr 1951) (McGinty coll.) 36-55 m (FLMNH 177402); 1 s, Monroe Co., off Dry Tortugas (R. Black, dec. 1990) 960 m (FLMNH 279375); 1 s, Palm Beach Co., Boynton Beach, off Briny Breezes (R/V Tritón, Feb 1951) (McGinty coll.) 50-60 fms (FLMNH 219907); 2 s, Collier Co., SW of John's Pass (Powlus and Steger, May 1958) (McGinty coll.) 55 m (FLMNH 154900); 2 s, Collier Co., 150 miles W (J. Moore, Jun. 1962) (M. Hunter coll.) Cape Romano, 450 m (FLMNH 129846); 1 s (Fig. 24K), Palm Beach Co., Bath and Tennis Club to Palm Beach Pier (R/V Tritón, Apr. 1951) (McGinty coll.) 36-55 m (FLMNH 176649); 1 s, Palm Beach Co.(R/V Tritón, Apr. 1950) 40-50 m (FLMNH 204877). Description: Shell (Figs. 24I-24L) solid, elongated, light brown. Proto- conch (Fig. 24N) multispiral, with about 4 whorls, the apex surface with small tubercles and the subsequent whorls with two spiral cordlets crossed by numerous fine axial ribs that are slightly S-shaped, especially on the last whorl. Teleoconch with about 11-12 whorls, beginning with spirals 1 and 3, crossed by prosocline small axial ribs which form nodules at the intersection points. These two spirals continué on the subse- quent whorls, the upper one with slightly larger nodules; on about the sixth-eighth whorl, a very small spiral 2 appears cióse to spiral 1; on the subse- quent whorls, this spiral 2 enlarges slightly until it is almost similar in size to spiral 3, spiral 1 is always slightly larger with larger nodules. On about the tenth whorl an additional spiral cord appears; this cord is smooth, always smaller, and located just below the lower suture, but on the final whorl it is more obvious and beaded. Below it there are two more spiral cords, the upper one slightly nodulous and the lower one smooth, located on the base of the siphonal canal. Aperture rounded-ovoid, columella curved, siphonal canal elongate and recurved, closed at its base by an extensión of the aperture. 144 ROLÁN AND FernáNDEZ-G ARCES: New data on the Caribbean Triphoridae Figure 29. “ Triphora ” turtleb ay ensis spec. nov. A, B: holotype, 4.1 mm, Bermuda (FLMNH); C, D: paratype, 4.2 mm (BMSM); E: protoconch of the holotype; F: detail of the protoconch of the paratype; G: detail of the aperture and base. Figura 29. “Triphora” turtlebayensis spec. nov. A, B: holotipo, 4, 1 mm, Bermudas (FLMNH); C, D: paratipo, 4,2 mm (BMSM); E: protoconcha del holotipo; F: detalle de la protoconcha del paratipo; G: detalle de la abertura y de la base. 145 Iberus , 26 (1), 2008 The colour is brown on the proto- conch; light brown on the shell, spiral 1 having lighter nodules, more evident on the last whorls. Dimensions : The shells studied measure between 4.0 and 9.0 mm. Distribution: Only known from Florida. Remarles : The shells figured by Abbott (1974) as T. lilacina may belong to this taxon. If the present well defined species is “T." lilacina , the only explana- tion would be that the lectotype of that taxon is an abnormally coloured shell, in which case the synonymy would only be acceptable if intergrades were found. On the other hand, if additional exam- ples of the typical lilac shells of "Triphora" lilacina were collected in the future, showing consistent differences with the shells described here, then the shells referred here as "Triphora" cf. lilacina could be named as a different species. "Triphora" atlántica (E. A. Smith, 1890) (Figs. 25A-I) Triforis atlántica E. A. Smith, 1890. Proc. Zool. Soc. Lond., 18: 292, pl. 21, fig. 16. Triphora lilacina Dalí, 1889. In Abbott, 1974: 112. Type material: Lectotype, here designated, of Triforis atlántica (Fig. 25A) and several paralectotypes (BMNH 89.10.1.1874-93). Other material examined: USA: Florida: 1 s (Fig. 25B), W Egmont Key, Hillsborough Co. (Gulf of México) Qim Moore /Cavalier, 1962), 76 m (CHL); 2 s, off Palm Beach (M. Glickstein, 1980) 82-105 m (CHL); 3 s (Figs. 25E-25G), 42 miles E St. Augustine, St. Johns Co. (T. Yocius, 1979), 85 m (CHL); 2 s (Fig. 25E, G), 42 miles E St. Augustine, St. Johns Co. (T. Yocius, Jul/1980), 36 m (CHL); 1 s, 50 miles S Carrabelle, Franklin Co. (J. Keeler, Feb/1986), 60 m (CHL); 2 s, W Marco, Collier Co. (J. Moore / Cavalier, 1972) 55 m (CHL); 1 s, Monroe Co., West coast, 24° 50' N, 33 fms (FLMNH 259042); 3 s, Palm Beach Co. (D. Akers, 1970-1971) 30 m (FLMNH 127891); 1 s, Palm Beach Co., off Boynton Inlet (D. and H. Akers, Sep. 1970) 36 m (FLMNH 228682). Louisiana: 2 s, off Louisiana, 26° 06.8' 66N 91° 02.418W (E.F. Garcia /Pelican, Jun/2001 57-65 m (CHL); 1 s, off extreme W Louisiana, 29° 45.9' N 13° 02.8' W, (E.F. Garcia /Pelican, Sept/98) 55-65 m (CHL); 1 s, off Louisiana, 28° 05.61N 91° 02.206' W (E.F. Garcia/ Pelican, Jun/2000), 58 m (CHL); Puerto Rico: 2 s. Rincón (L. Germaine, Apr/1961) 30 m (FLMNH 163833). Brazil: 3 sp. (Figs. 25C, 25D) off Guarapari, Espiritu Santo State (R. Bodart, 1994) 15-20 m (CHL). Description: Shell (Figs. 25A-25H) sharp-pointed, elongate, solid, with bands of brown and white. Protoconch multispiral (Fig. 251) with the apex retic- ulated and 4 whorls with narrow axial ribs that cross a single thread on the first whorl and 2 elsewhere, except at the end where there is only one. Teleoconch with about 12-13 whorls, sometimes more on large shells, beginning with spirals 1 and 3, crossed by prosocline axial ribs which form nodules at the intersection points. These two spirals continué on the subsequent whorls, the upper one white and with slightly larger nodules; on about the eighth-ninth whorl, a very small spiral 2 appears cióse to spiral 1; on the subsequent whorls, this spiral 2 enlarges slightly until it is almost similar in size to spiral 3, spiral 1 continuing to have larger, more prominent white nodules. On about the tenth-twelfth whorl an addi- tional spiral cord appears; this cord is only slightly nodulous, always smaller, and located just below the lower suture, but on the final whorl it is more obvious, is located near the periphery and has larger nodules. Below it there are two more spiral cords, the upper one slightly nodulous and the lower one smooth, located on the base of the siphonal canal. Aperture rounded- ovoid, columella curved, siphonal canal elongate and recurved, closed at its base by an extensión of the aperture. The colour is dark brown on the pro- toconch; white on the first two teleo- conch whorls, subsequently with white nodules on spiral 1 and light brown in 146 RoláN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 30. A-F. “Triphora” grenadensis spec. nov. A-D: holotype, 3.2 mm, Grenada (ANSP); E, F: protoconch of the holotype. G-I. “ Triphora ’ sp. 1, 3.3 mm, British Honduras (Belize) (ANSP). Figura 30. A-F. “Triphora” grenadensis spec. nov. A-D: holotipo, 3,2 mm, Granada (ANSP); E, F: protoconcha del holotipo. G-I. “Triphora” sp. 1, 3,5 mm, British Honduras (Belize) (ANSP). 147 Iberus, 26 (1), 2008 the interspaces; the background colour of spiral 2 is brownish, with the nodules somewhat lighter. On some shells the colour is similar but lighter throughout. Dimensions : The lectotype measures 6.0 mm. Some shells from the Caribbean can reach up to 9.0 mm. Distribution: It is known from Florida to Brazil (not collected in Cuba). Other- wise, St. Helena, in the middle of the Atlantic is the type locality of T. atlántica. Remarks: This taxon is very confused, and for many years it was considered by some authors and collectors to be T. lilacina. Yet the lectotype of T. lilacina is uniform pink-lilac, spirals 1 and 3 are well developed and spiral 2 never reaches the size of the other two. The nodules are the same colour as the shell and are not lighter, and there are no dif- ferences between the colour of spiral 1 and spiral 3. No intergrades between these two morphs were found. Further- more, the nodules are smaller and the protoconch more sharply pointed with tubercles on the apex. However, a defin- itive conclusión can not be reached until enough material of true "T." lilacina becomes available for examination, allowing a proper comparison and a study of the animal and radula. On the other hand, the taxon “T." atlántica described by Smith from St. Helena is a shell with constant brown and white colouration and distribution of the bands, and with nodules of similar size. The location of the type locality in the middle of the Atlantic is not a problem for a species which has a multispiral protoconch. The only problem is that the protoconch of the holotype is not known, but the eroded protoconch of a paralectotype showed 2 spiral threads, which is compatible with that of the present population. Therefore we suggest that the shells studied from the Caribbean belong to this taxon, the only doubts being due to the lack of a good protoconch from the St. Helena material. Provisionally this species must therefore be kept under this ñame, awaiting the information that new mate- rial will provide in the future. It may be confused with Monophorus ateralbus Rolán and Fernández-Garcés, 1994, but the latter species is smaller and has a totally different colour pattern, with the lower band white in contrast to that of "T." atlántica, on which the white spiral is always the upper one. Triphora ellyae De Jong and Coomans, 1988 is smaller and does not reach 4 mm. "Triphora" pyrrha Henderson and Bartsch, 1914 (Fig. 25J) Triphoris pyrrha Henderson and Bartsch, 1914. Proc. U.S.N.M. 47: 413, pl. 2, f. 4 [Type locality: Chincoteague, Virginia, USA], Type material: A syntype (Fig. 25J) here designated as lectotype (USNM 252571). Description : Henderson and Bartsch (1914). The lectotype is a shell in poor condition, white or cream in colour, or possibly faded brown. The protoconch is multispiral and has about 4 whorls but is very eroded, and further details are impossible to discern. The next 3 whorls have 2 spirals cióse together; on the following whorls, these cords 1-3 are more sepárate; subse- quently a narrow spiral 2 appears in the middle of the whorls; on the final whorl spiral 2 is similar in size to the other two. The siphonal canal is short and open. Dimensions : The lectotype measures 2.7 mm. Distribution: Only known from its type locality. Remarks: Due to the lack of informa- tion on the protoconch, the validity of this species must be confirmed by the examination of more material from the type locality. It is somewhat similar to Marshallora modesta or M. nigrocincta, but is probably a valid species. 148 RoláN AND FERNÁNDEZ-GARCÉS: New data on the Caribbean Triphoridae Figure 31. “Triphora” abacoensis spec. nov. A, B: holotype, 2.3 mm, Abaco, Bahamas (BMSM); C: paratype, 2.25 mm, Abaco, Bahamas (CCR); D: protoconch of paratype, Abaco, Bahamas (MHNS); E: protoconch of paratype, Abaco, Bahamas (CCR); F: protoconch, Cienfuegos, Cuba (MHNS); G: detail of the microsculpture of Figure E. Figura 31. “Triphora” abacoensis spec. nov. A, B: holotipo, 2,5 mm, Abaco, Bahamas (BMSM); C: paratipo, 2.25 mm, Abaco, Bahamas (CCR); D: protoconcha de un paratipo. Abaco, Bahamas (MHNS); E: protoconcha de un paratipo, Abaco, Bahamas ( CCR); F: protoconcha, Cienfuegos, Cuba (MHNS); G: detalle de la microescultura de la Figura E. 149 Iberus , 26 (1), 2008 "Triphora" inaudita spec. nov. Rolán and Lee (Figs. 26A-D) Type material: Holotype (Fig. 26A) in FLMNH; paratype (Fig. 26B) in BMSM from 28° 05.61' N, 91° 02.205' W to 29° 05.524' N, 91° 02.086' W (E.F. García /Pelican 5/27/00) 58 m (both ex CHL). Type locality: SW Dry Tortugas, Florida, USA 26° 42.9' N, 83° 43.20' W, 73.3-78.5 m. Etymology: The specific ñame derives from the Latín word inauditus, which means "uncommon, unheard-of", referring to some curious characters of the shell. Description : Shell (Figs. 26A, 26B)) very distinctive, solid, elongate. Proto- conch (Fig. 26C) with 3 and 1 /i whorls with two spiral cords crossed by axial ribs. Teleoconch (Fig. 26D) with three spirals from the beginning, crossed by slightly prosocline axial ribs and forming nodules at the intersections. Spiral 1 is very small on the first whorls and is located very near to spiral 2; spirals 2 and 3 are similar in size. On the subsequent whorls, spiral 1 increases in size until it reaches a size similar to spiral 2 on about the tenth whorl, but is still less prominent. Throughout the shell spirals 1 and 2 are closer than spirals 2 and 3. On the last whorl three additional spirals appear towards the base. Aperture ovoid, almost rhomboid, with a short and open siphonal canal. The colour is very characteristic: spiral 1 is mainly brown, with 1-2 white nodules alternating with 1-3 brown ones; spiral 2 is mainly white, with only a small area of brown on the lower part of some nodules; on spiral 3 the brown colour predominates, but white nodules appear between two or more brown ones. Dimensions : The holotype measures 5.4 mm and the paratype 4.4 mm. Distribution : Only known from the type locality. Remarks : Some of the characters of "T. " inaudita are totally different from any other species from the study area and therefore it has been named in spite of the scarcity of available material. The colour pattern, the smaller spiral 1 and the proximity of spirals 1 and 2 are the most important differential characters. The most similar species are: "Triphora" cf. atlántica has two spirals on the first whorls, the nodules of the upper spiral are white and larger, while the rest are brown. Cosmotriphora arnoldoi Faber and Moolenbeek, 1991 has uniform blotches of brown on a white background, spiral 2 does not exist on the first whorls and is smaller on the final ones. Monophorus olivaceus has the lower- most spiral totally white; this spiral is larger on the first whorls, while spiral 2 is the smallest when it appear s. Spirals 1 and 2 ha ve only a few white nodules. Nototriphora decorata (C. B. Adams, 1850) has three spirals similar and equidistant throughout the shell, the brown blotches are isolated on the whorls against a white background. "Triphora" pseudonovem spec. nov. Rolán and Femández-Garcés (Figs. 26E-I) Type material: Holotype (Fig. 26E) and 2 paratypes in FLMNH (193355). Type locality: Barnegat Bay, Massachusetts, USA. Etymology: The specific ñame alludes to the similarity with Mesophora novem. Description : Shell (Fig. 26E) subconi- cal elongate, solid. Protoconch (Figs. 26G, 16H) with 3 l/i whorls and a diam- eter of about 300 pm. The nucleus is tuberculated and is followed by one spiral thread crossed by numerous slightly opisthocline axial ribs. Teleo- conch with about 14 whorls, spirals 1 and 3 appearing beaded at the begin- ning, with the ribs clearly prosocline; on the subsequent whorls the spirals slowly become more widely separated. The nodules are large and spherical, of similar size on both spirals, the upper nodules connected to the lower ones by well defined and slightly curved proso- 150 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae Figure 32. “ Triphora portoricensis spec. nov. A, B: holotype, 4.4 mm, Puerto Rico (FLMNH); C: protoconch of the holotype; D: paratype, 1.8 mm, Abaco, Bahamas (CCR); E: protoconch of the paratype. Figura 32. “Triphora” portoricensis spec. nov. A, B: holotipo, 4,4 mm, Puerto Rico (FLMNH); C: protoconcha del holotipo; D: paratipo, 1,8 mm, Abaco, Bahamas (CCR); E: protoconcha del paratipo. cline axial ribs (Fig. 261). Spiral 2 appears between the other two spirals on the final two or three whorls; it is very fine and is located very cióse to spiral 1, not increasing immediately in size and remaining clearly smaller than the other two on the final whorl. Towards the base there are three more nodulous spirals. Aperture rounded, small, opened to the siphonal canal, which is curved and a little smaller in size than the height of the aperture (Fig. 26F). The entire shell is brown, with the nodules a little lighter. Dimensions : The holotype measures 7.3 mm. Distribution: Only known from the type locality. Remarks : This species has some of the characters of Mesophora novem, such as spirals 1 and 3 separated throughout most of the shell, but this new species is brownish, whereas M. novem is tinged with violet and has the first teleoconch whorls white; another difference is the slowly increasing size of spiral 2. The protoconch has only one spiral thread and it is shorter than that of M. novem. 151 Iberus , 26 (1), 2008 "Triphora" calva Faber and Moolenbeek, 1991 (Figs. 27A-G) Triphora calva Faber and Moolenbeek, 1991. Apex, 6(3/4): 82, figs. 3, 4. [Type locality: Key Bis- cayne. Florida, USA]. Type material: Holotype not examined (ZMA 391003). Represented in the original description. Material examined: Bahamas: 1 j (Fig. 27F), Treasure Cay, Abaco 26 40.12N 77 18.19W (CCR); 5 j, North Sound, Bimini, mangrove (R. Robertson, 1957) (ANSP 329747); 3 s, Grand Bahama Island, 26° 31' 00"N, 78° 46' 30"W (J. Worsfold) (ANSP 373944); 1 s (Fig. 27G) Bootle Bay, Grand Bahama Island, 26° 39' 30"N, 078° 57' 00" (J. Worsfold) (ANSP 371845); 6 sp, North Hawksbill Creek, Grand Bahama Island 26° 32' 00"N, 78° 45' 00"W (J. Worsfold) (ANSP 370520); 1 sp, 7 s. Hotel, West End, Grand Bahama Island 26° 42' 15"N, 78° 59' 50"W (J. Worsfold, Dec. 1981) (ANSP 368674); 1 s, McLeans Town, Grand Bahama Island, 26° 38' 45"N, 77° 57' 30"W (J. Worsfold) (ANSP 368484); 1 s, Hope Town Harbour, Abaco (R. Robertson, Jun. 1953) (ANSP 299409). Grand Cavman Island: 1 s, W of Prospect, SW Sound (A. J. Ostheiner) (ANSP 198925). Florida: 1 s, 1 f (Fig. 27A), Key Largo (CMK); 1 sp, 1 s, Bonefish Key (Aug. 1957) (FLMNH UF368198). Cuba: 3 s, Caibarién, (FLMNH UF365100); 1 f (Fig. 27D), Cienfuegos (CFG); 1 f, Jibacoa (MHNS); 1 f (Fig. 27E) Santa Lucia, Pinar del Rio (CFG); 3 s, Batabanó (CFG); 1 s, Cabo Mafias, Canarreos (MHNS). Description: Faber and Moolen- beek (1991). The most important diag- nostic characters are the following: the shell is dark brown; the brown colour may be uniform, sometimes with the nodules lighter (Fig. 27C) and even with spiral 1 on the last whorl with white nodules (Fig. 27A). The proto- conch (Figs. 27B, 27D, 27E, 27F) is described as having only one smooth whorl, but if the protoconch whorls are counted by the Verduin (1977) method, and the end of the protoconch is at the point where the teleoconch begins with two spirals, it actually has 2 whorls (sometimes a little less); the first whorl is smooth and there is a spiral cord at the middle of the second one. In the original description there is no Infor- mation on the diameter of the proto- conch. Based on the photograph of the holotype it could be deduced that the protoconch is about 275 pm in width; in the material examined the diameter is variable between 250 and 380 pm. This and the brown colour are impor- tant characters for differentiation from other species. Teleoconch sculpture begins with spiral 3; spiral 1 and axial ribs appear on the third whorl (the second whorl of the teleoconch). Spiral 2 emerges on the sixth whorl (the fifth of the teleoconch). Also another smooth sutural spiral com- mences on the 3th-4th whorl. Dimensions : The holotype measures 5.0 mm. Some shells atributed to this species can reach 9 mm. Distribution : Known from several areas of the Caribbean: Florida, Bahamas, Cuba. Remarks : In some works (e.g. Rolán and Fernández-Garcés, 2007) this species is assigned to the genus Marshal- lora. In fact there is no information on the radula and it is therefore better to keep it in "Triphora" s. /. Shells from Quintana Roo were referred to in the original description under "Other material studied" to be this species; these shells were later described as Marshallora nichupte by Rolán and Crúz-Ábrego (1996). This species can be differentiated from "T. " calva by its more variable colour, ranging from dark brown to white, larger shell and wider protoconch, with only 1 to 1 l/z whorls. Below, several species are shown to be different from "T." calva although having a short protoconch as a common feature. The protoconch of this species has been studied in shells from the Bahamas, Florida and Cuba (Figs. 27B, 27D, 27E, 27F), all of which have a first whorl with a similar diameter: 250, 270 pm and 260 pm respectively. Other similar characters inelude a smooth apex and one spiral cord on the second whorl. 152 ROLÁN AND FernáNDEZ-GarcÉS: New data on the Caribbean Triphoridae One shell from the Bahamas (Fig. 27G) resembles "T." calva but is larger (6.8 mm) and elongated (11 whorls), and the protoconch has only one spiral whorl instead of two. It may be a differ- ent species or an abnormal individual. "Triphora" yociusi spec. nov. Rolán and Lee (Figs. 27F1, 1) Type material: Holotype (Fig. 27H) deposited in FLMNH {ex CHL). Paratype (Fig. 271) in BMSM from 67 miles E St. Augustine, St. Johns Co., Florida (T. Yocius 1979) 54 m; both ex CHL. Other material studied: 1 s (eroded), Bermuda (C. Abbott Davis) (ANSP 88722). Type locality: 32 miles E St. Augustine, St. Johns Co., Florida (J. Dawley / Scallop 1982), 30 m. Etymology: The species is named after Ted Yocius, the St. Augustine fisherman who dredged so many of the triphorids from off northeast Florida, including the present species. Description : Shell (Fig. 27H) subconi- cal elongate, solid. Protoconch (Fig. 271) paucispiral with 1 3 A whorls and a dia- meter of about 370 pm. The nucleus and most of the first whorl are smooth; near the end of this whorl two spiral cords appear. Teleoconch with about 8 whorls, with spirals 1 and 3 nodulous at the beginning. The nodules are large and rounded, of similar size on both spirals, the upper ones connected to the lower by fine prosocline axial ribs. On about the fifth whorl of the teleoconch spiral 2 appears between the other two, but closer to spiral 1; it is very narro w, sub- sequently increasing in size until the three spirals are similar on the final whorl. Aperture rounded, small and open to the siphonal canal, which is half the size of the aperture. The entire shell is brown, but a darker band covers spiral 1 and reaches to the suture, the nodules on this spiral being lighter than the background colour; the nodules and the background colour of spiral 3 are cream coloured, as is spiral 2 on the last whorl. Dimensions : The holotype measures 3.9 mm. Distribution : Only known from type locality and area cióse by. Remarks : ''Triphora'' yociusi spec. nov. has a wider protoconch than "T". calva, with 1 3/ 4 whorls, and at the end of the first protoconch whorl there are two spirals, while in "T" . calva there is only one. The teleoconc