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Potamides Et Pirenella (Gastropoda, Prosobranchia) À Partir Des Espèces Actuelles Et Fossiles : Implications Phylétiques Et Biogéographiques

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Potamides Et Pirenella (Gastropoda, Prosobranchia) À Partir Des Espèces Actuelles Et Fossiles : Implications Phylétiques Et Biogéographiques Annales de Paléontologie (Vert.-lnvert.). © Masson, Paris, 1986. 1986, vol. 72, fase. 3, pp. 163-210. REDÉFINITION DES GENRES POTAMIDES ET PIRENELLA (GASTROPODA, PROSOBRANCHIA) À PARTIR DES ESPÈCES ACTUELLES ET FOSSILES : IMPLICATIONS PHYLÉTIQUES ET BIOGÉOGRAPHIQUES PAR Pierre LOZOUET * Mots-clés: Gastropodes. Prosobranches. Potamides. Pirenella. Taxonomie. Phylogénie. Key-words : Gastropoda. Prosobranchia. Potamides. Pirenella. Taxonomy. Phylogeny. Résumé. - Il est montré que Pirenella est synonyme de Potamides et que les « Pirenella » de la littérature paléontologique doivent être attribuées au genre Granulolabium. La reconstitution de la phylogénie de Potamides montre que P. conicus (= cailliaudi) peut être considéré comme une relique mésogéenne. Probablement, le genre Potamides, d'origine européenne, a pénétré tardivement dans la province Indo-Pacifique (Pliocène voire même Quaternaire). La phylogénie du genre Granulolabium est aussi reconstituée et un représentant actuel, très proche de l'espèce oligocène d'Europe G. plicatum est reconnu: « Zeacumantus » diemenensis Quoy et Gaimard. Granulolabium diemenensis est actuellement confiné au sud-est de l'Australie et à la Tasmanie où il est commun dans les habitats vaseux du médiolittoral. Deux sous-genres sont reconnus: Granulolabium (s.s.) et Granulolabium (Tiarace­ rithium); le genre néo-zélandais Zeacumantus apparaît étroitement apparenté. * Lab. de Biologie des Invertébrés Marins, 55, rue de Buffon, 75005 Paris et Lab. du Quaternaire de l'E.P.H.E., Inst. des Sciences de la Terre, 6, Bld Gabriel, 211oo Dijon, France. - 163- Annales de Paléontologie (Vert.-!nvert.), 1986, vol. 72, fascicule 3. 13 2 PIERRE LOZOUET Abstract. - It is shown that Pirenella is synonymous with the genus Potamides and that the « Pirenella » of the palaeontologicalliterature should be assigned to Granulolabium. The reconstitution of the phylogeny of the Potamides proves that P. conicus (= cailliaudi) can be considered as a mesogean relict. Probably, the genus Potamides (of European origin) has entered, only very late, into the Indo-Pacific province (Pliocene and even Quatemary). The phylogeny of the Granulolabium is also reconstituted and a recent representative, quite similar to the European üligocene species G, plicatum has been identified : « Zeacumantus » diemenensis (Quoy et Gaimard). Granulolabium diemenensis is now confined to the South East of Australia and Tasmania where it is easy to find in the muddy habitats of the Midlittoral. Two subgenera have been recognized : Granulolabium (s.s.) and Granulolabium (Tiaracerithium); the New-Zealander genus Zeacumantus seems very weil connected. INTRODUCTION Les Potamides et les Pirenella pullulent souvent dans les terrains cénozoïques européens; par exemple, des comptages précis (Gitton et al., à paraître) ont montré que «Pirenella » plicata était le mollusque le plus abondant de l'Oligocène du bassin de Paris. Il Y a déjà plusieurs années, j'ai remarqué la similitude des espèces types des genres Pirenella (P. conica, actuel) et Potamides (P. lamarcki, Stampien). Or, aucune publication jusqu'au récent article de Kadolsky (1984) ne faisait allusion à la synonymie de Pirenella et de Potamides que je souhaite démontrer ici. La mauvaise interprétation des rapports entre les formes actuelles et fossiles de ces deux genres est à l'origine d'une confusion extrême que l'on retrouve dans la classification de Bouniol (1981) où les Pirenella sont intégrés aux Tiaracerithiinae et les Potamides aux Potamidinae. Cossmann (1906) subdivisait les Cerithiidae en Cerithiinae, Potamidinae et Bittiinae, Thiele (1929) élève les Potamidinae de Cossmann au rang familial et introduit la sous-famille des Batillariinae. Les Potamididae se composent alors des Potamidinae et des Batillariinae. Wenz (1938-44) reprendra cette classification qui est encore celle adoptée actuellement par les malacologues. Cependant Bouniol (1981) a proposé de scinder les Cerithiidae en quatre sous-familles (Cerithiinae, Potamidinae, Batillariinae et sa nouvelle sous-famille des Tiaracerithiinae) qui constitueraient un ensemble polyphylétique. Ce classement ne tient pas compte des caractéristiques anatomiques et notamment du fait bien connu que tous les Potamididae de Thiele possèdent un opercule corné, circulaire, de type spiral polygyré, alors que l'opercule corné des Cerithiidae est de type spiral oligogyré. D'autre part, si la dichotomie classique entre Potamides saumâtres et cérithes marins souffre, ainsi que l'a rappelé Plaziat (1984, p. 295), quelques exceptions, elle n'en reste pas moins dans la grande majorité des cas exacte, et le mode de vie des Potamides n'est pas une convergence fortuite comme le croit Bouniol (1981). Houbrick (1984) prend soin de - 164- POTAMIDES ET PIRENELLA 3 réaffirmer la classification de Thiele sans toutefois mentionner l'hypothèse de Bouniol et la sous-famille des Tiaracerithiinae. Pour ma part, je considère que les Potamididae doivent être compris au sens de Thiele en y intégrant les Tiaracerithiinae de Bouniol qu'il sera nécessaire de redéfinir. Cette rectification de nomenclature m'a obligé à traiter le problème dans sa totalité, c'est-à-dire en retraçant l'histoire des groupes concernés. Le texte a été ainsi divisé en trois parties: une première partie est d'ordre systématique et taxinomique, une seconde retrace l'histoire du genre Potamides de l'Oligocène à l'actuel et la troisième est consacrée aux genres Granulolabium et Zeacumantus. Dans les seconde et troisième parties, figure un relevé des principales « espèces» du cénozoïque européen, témoins du lourd héritage typologique chez les Potamididae, et l'espèce Granulolabium plicatum, espèce type du genre qui doit remplacer les ex-Pirenella fossiles, est l'objet d'une étude biométrique. SYSTÉMATIQUE DES POTAMIDIDAE APERÇU NOMENCLATURAL ET ÉCOLOGIQUE Les différents genres de Potamididae évoqués au cours de ce travail sont présentés dans leur sous-famille respective, en privilégiant les taxons fossiles. Tiaracerithinae Bouniol, 19891. - Bouniol (1981) a créé cette sous-famille pour les genres Pirenella, Terebralia, Tiaracerithium et son sous-genre Gravesice­ rithium. Dans un premier temps, nous y adjoindrons les genres considérés jusqu'à présent comme synonymes de Pirenella : Tiarapirenella et Granulolabium. Pirenella Gray, 1847 (espèce-type: P. conica Blainville, actuel, Méditer­ ranée). Outre Pirenella conica (= mamillata PhiL), on relève en Méditerranée des variants considérés par certains auteurs comme des espèces distinctes : P. decorata, peloritana et cinerascens. Dans la Mer Rouge et l'Océan Indien il existe une autre forme généralement considérée comme une espèce distincte : P. cailliaudi (Poitiez et Michaud). Pirenella conica est une espèce abondante dans les lagunes de Méditerranée orientale. L'écologie et la répartition des Pirenella sont traitées en détail dans la deuxième partie. Terebralia Swainson, 1840 (espèce-type: T. palustris Bruguière; Indo­ Pacifique). Ce genre paraît actuellement inféodé aux mangroves du domaine indo-pacifique. - 165- 4 PIERRE LOZOUET Tiaracerithium Sacco, 1895 (espèce-type: T. pseudotiarella d'Orbigny, Burdigalien d'Aquitaine). Ce genre fossile est particulièrement abondant dans les milieux calmes, évoluant vers une dessalure, de l'Auversien du bassin de Paris. Gravesicerithium Charpiat, 1923 (espèce-type: G. tiara Lamarck, Lutétien du bassin de Paris). Gravesicerithium Bouniol (1981) est rigoureusement synonyme. Granulolabium Cossmann, 1889 (espèce-type: G. plicata Bruguière, Aquita­ nien de la Caunette près de Montpellier). L'espèce G. plicata, qui existe de l'Oligocène inférieur au Miocène inférieur, a reçu un grand nombre de noms en rapport avec une variabilité étendue. Elle est considérée comme à l'origine des Pirenella actuelles. Tiarapirenella Sacco, 1895 (espèce-type: T. bicincta Brocchi, Miocène supérieur d'Italie). Pour Sacco (1895 : 61) «Cerithium» pictum (Basterot) et « Cerithium » mitrale (Eichwald) respectivement du Miocène inférieur du Bordelais et du Miocène moyen de la Paratéthys appartiennent à ce sous-genre. Les genres fossiles Tiaracerithium, Granulolabium et Tiarapirenella sont présents dans le même type de dépôts laguno-marins. Avec les Tiaracerithium de l'Auversien du Guépelle (bassin de Paris) comme avec les Granulolabium de l'Oligocène belge et du bassin de Paris, on trouve le même type de faunule accompagnatrice: Hydrobies, Stenothyrioides, Nystia. Quant à Tiarapirenella, il ne fait que se substituer à Granulolabium au Miocène inférieur. L'aggravation des conditions saumâtres ne semble pas favoriser le groupe des Granulolabium et Tiaracerithium alors concurrencé par les Potamides à plus large valence euryhaline. Dans l'ultime formation marine parisienne (Sables d'Ormoy), avec le retrait de la mer stampienne, on suit la régression de Granulolabium parallèlement au développement de Potamides lamarcki. Potamidinae H. et A. Adams, 1854. - Potamides Brongniart, 1810 (espèce­ type: P. lamarcki Brongniart, Oligocène du bassin de Paris). P. lamarcki caractérise les dépôts saumâtres de l'Oligocène. Les figures d'Alimen (1936, pl. 6) donnent une bonne idée de la variabilité de cette espèce. Dans le bassin parisien, P. lamarcki connaît une explosion démographique à la partie sommitale du Stampien, avec la généralisation des biotopes lagunaires. De nombreux points autour d'Etampes livrent, au sein de lits de calcaires marneux, une communauté composée exclusivement de P. lamarc·ki et d'Hydrobia. Dans le bassin du Rhône, on recueille dans des dépôts de
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