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Linnaeus, 1758) and O Comparative morphology of Pliocene, Quaternary and Recent shells of Ocenebra erinaceus (Linnaeus, 1758) and O. brevirobusta Houart, 2000 (Mollusca, Muricidae, Ocenebrinae): reflections on the intra- and interspecific variations Véronique BERROU 8 allée des Avocettes, Parc-Quibias, F-56610 Arradon (France) Didier MERLE Département Histoire de la Terre, Muséum national d’Histoire naturelle, UMR 5143 du CNRS, 8 rue Buffon, F-75231 Paris cedex 05 (France) [email protected] Jean-Louis DOMMERGUES Université de Bourgogne, Centre des Sciences de la Terre, 6 boulevard Gabriel, F-21000 Dijon (France) [email protected] Catherine CRÔNIER Université des Sciences et Technologies de Lille, Laboratoire de Paléontologie et Paléogéographie du Paléozoïque, UMR 8014 du CNRS, F-59655 Villeneuve d’Ascq cedex (France) [email protected] Didier NÉRAUDEAU Université de Rennes 1, Campus de Beaulieu, F-35042 Rennes cedex (France) [email protected] Berrou V., Merle D., Dommergues J.-L., Crônier C. & Néraudeau D. 2004. — Comparative morphology of Pliocene, Quaternary and Recent shells of Ocenebra erinaceus (Linnaeus, 1758) and O. brevirobusta Houart, 2000 (Mollusca, Muricidae, Ocenebrinae): reflections on the intra- and interspecific variations. Geodiversitas 26 (2) : 263-295. ABSTRACT The morphological variations of Pliocene, Quaternary and Recent shells of Ocenebra erinaceus (Linnaeus, 1758) and O. brevirobusta Houart, 2000 are analyzed. The study of these oyster drillers is based on various methods (cladistics, biometry, Procrustes and Fourier analyses) and concerns popula- tions from 40 localities of the Atlantic (from Normandy to Morocco) and GEODIVERSITAS • 2004 • 26 (2) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 263 Berrou V. et al. Mediterranean shorelines. The results indicate a morphological distribution of North/South orientation, the more derived southern forms being closely related. In spite of a strong variability, a beginning of differentiation, proba- bly dating from the Plio-Pleistocene, is observed between three groups: 1) Northern Atlantic Ocean (O. erinaceus); 2) Mediterranean Sea (O. eri- naceus); and 3) Moroccan Atlantic (O. brevirobusta). O. brevirobusta is tem- porarily regarded as a geographic subspecies of O. erinaceus, because of its uninterrupted marginal area, and its P2 cord more atrophied than in the KEY WORDS Italian late Pliocene O. erinaceus. Within these groups, the multivariate analy- Mollusca, Gastropoda, sis of variance distinguishes local biogeographic entities. In addition, two Muricidae, unexpected results deserve to be stressed: 1) the population from Étang de Ocenebrinae, Thau, systematically associated to those of the Northern Atlantic Ocean, is Ocenebra, shells, located in a zone of oyster importation from the Oléron island; and 2) the Neogene, population from Algarve (Portugal), closer to those of the Mediterranean Sea, Recent, Europe, lives in an Atlantic area undergoing the influence of Mediterranean waters. variations, Comparisons between O. erinaceus and Ocinebrellus inornatus (Récluz, 1851), populations, an Asian species introduced on the French Atlantic coast, show more simila- cladistics, morphometry, rities with the Northern Atlantic O. erinaceus, than with Mediterranean or evolution. Moroccan Atlantic O. erinaceus. RÉSUMÉ Comparaison morphologique des coquilles pliocènes, quaternaires et actuelles d’Ocenebra erinaceus (Linnaeus, 1758) et d’O. brevirobusta Houart, 2000 (Mollusca, Muricidae, Ocenebrinae) : réflexion sur les variations intra- et inter- spécifiques. La variation morphologique de coquilles pliocènes, quaternaires et actuelles d’Ocenebra erinaceus (Linnaeus, 1758) et d’O. brevirobusta Houart, 2000 est analysée. L’étude de ces perceurs d’huîtres, fondée sur différentes méthodes (analyse cladistique, biométrie, morphométrie géométrique et transformée de Fourier), porte sur des populations de 40 localités des côtes atlantiques (de la Normandie au Maroc) et méditerranéennes. Les résultats indiquent une répartition morphologique d’orientation Nord/Sud, les formes méridionales, plus dérivées, étant étroitement apparentées. Malgré une forte variabilité, un début de différenciation, datant probablement du Plio-Pléistocène, s’observe entre trois groupes de populations : 1) Atlantique septentrional (O. erinaceus) ; 2) Méditerranée (O. erinaceus) ; et 3) Maroc atlantique (O. brevirobusta). O. brevirobusta est provisoirement considérée comme une sous-espèce géographique d’O. erinaceus, en raison d’une aire marginale non disjointe et d’une atrophie du cordon P2 plus marquée chez les O. erinaceus MOTS CLÉS du Pliocène supérieur italien. Au sein des trois groupes, l’analyse multivariée Mollusca, Gastropoda, de variance distingue des entités biogéographiques locales. Deux résultats Muricidae, inattendus méritent d’être soulignés : 1) la population de l’Étang de Thau, Ocenebrinae, systématiquement associée par les analyses à celles de l’Atlantique septentrio- Ocenebra, coquilles, nal, est située dans une zone d’échanges d’huîtres avec l’île d’Oléron ; et 2) la Néogene, population de l’Algarve (Portugal), morphologiquement proche de celles de actuel, Europe, Méditerranée, est située dans une région de l’Atlantique subissant l’influence variations, des eaux méditerranéennes. Les comparaisons entre O. erinaceus et populations, Ocinebrellus inornatus (Récluz, 1851), espèce introduite par l’homme en cladistique, morphométrie, Atlantique, révèlent de plus grandes similitudes avec les O. erinaceus de évolution. l’Atlantique septentrional qu’avec celles de Méditerranée ou du Maroc. 264 GEODIVERSITAS • 2004 • 26 (2) Shell variations in European Ocenebra (Mollusca, Muricidae) INTRODUCTION O. erinaceus, O. brevirobusta Houart, 2000 (Houart 2000a) and O. chavesi Houart, 1996 During the last decades, numerous allochthonous (Houart 1996). Fossils of O. erinaceus have been species of molluscs have been introduced on the reported from the middle Miocene of Poland French Atlantic coast by human way (Gruet et al. (Baluk 1995), but the identification by Baluk 1976; Sauriau 1991; Montaudouin & Sauriau (1995) is probably wrong, the figured specimens 2000). One of the last introductions is that of an being more similar to members of the genus Jaton ocenebrine, Ocinebrellus inornatus (Récluz, Push, 1837 (type species by original designation: 1851). It is dated from 1997 and reported in the Murex decussatus Gmelin, 1791), particularly Marennes-Oléron basin (Charente-Maritime, J. dufreynoi (Grateloup, 1847) or J. sowerbyi France). The species is often assigned to (Michelotti, 1842). However, the occurrence of Ocinebrellus Jousseaume, 1880 (type species by O. erinaceus is well attested in the Pliocene of original designation: Murex eurypteron Reeve, Italy (Malatesta 1974), Spain (Muniz Solis & 1845 [a junior synonym of M. aduncus Sowerby, Guerra-Merchan 1994; Martinell 1979; 1834]), but has been recently transferred (Houart Martinell & Marquina 1981) and in the & Sirenko 2003) in the genus Ocenebra Gray, Pleistocene of the Northern domain (Glibert 1847 (type species: O. erinaceus Linnaeus, 1758 1963). The geographic range of Recent popula- by monotypy). Ocinebrellus inornatus is an oyster tions of O. erinaceus (Fig. 1A) extends from driller. Although it does extend into warm tem- Southern England and Ireland to the perate waters, the species is properly a low-boreal, Mediterranean Sea and the Gibraltar Strait. The cool water temperate form. It is native from east- species is also reported from Madeira (Houart ern, western Japan and mainland Asia, from 33° 2001a), Canary Islands (Garcia-Talavera 1974) to 51° north latitude. O. inornatus has already and from Azores, where it has been recently been introduced by human way on the Pacific introduced (Houart & Abreu 1994; Houart coasts of Oregon, Washington and British 2001a). The fossils of O. brevirobusta are report- Columbia (Radwin & Attilio 1976; Carlton ed from the lower Pleistocene (Moghrebian- 1992; Noda et al. 1993). On the French shore- Messaoudian) of Morocco by Brébion (1978, line, O. inornatus begins dangerously to damage 1979a, b) under the name Murex torosus the oyster farming, and consequently this inva- Lamarck, 1816, which probably corresponds to a sion may have economical repercussions. The junior synonym of the Pliocene species molecular analysis of the French O. inornatus Heteropurpura polymorpha (Brocchi, 1814) (see demonstrates that it has been introduced from Houart 2000a). The geographic range of Recent a stock having closer relationships with the populations of O. brevirobusta (Fig. 1A) is American population than the Asian (Garcia- restricted to the Moroccan Atlantic coast (El Meunier et al. 2001; Martel et al. 2003). Jadida, Rabat, Temara, Essaouira [= Mogador], Moreover, its life environment corresponds to Agadir and Asilah) and northward, it does not that of O. erinaceus (Pigeot et al. 2000; Koster- pass Tanger. Following Houart (2000a, 2001a), Sperling 2002) and a dramatic decrease of the O. brevirobusta and O. erinaceus are sympatric at autochthonous oyster drill is reported when Asilah. O. chavesi is an endemic species from the O. inornatus shows a high density (Pigeot et al. Azores and is only known from San Miguel 2000). Therefore, some populations of O. eri- Island. No fossil of Ocenebra are reported from naceus might be locally extirpated and a study of Miocene and Quaternary terraces of Santa Maria its morphological variation also answers to
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