Bollettino Malacologico, Roma, 39 (5-8): 111-112, [2004] - Società Italiana di Malacologia

Djeddilia and Escoffieria, abnormal specimens of (: : ,) Pierre Lozouet

KEY WORDS: , Djeddilia, Escoffieria, Potamides, Tympanotonos, .

ABSTRACT The relationships of the enigmatic modern Red Sea Djeddilia with the Pliocene genus Escoffieria are discussed. Both Djeddilia and Escoffieria (a junior synonym of Potamides) were based on abnormal, partially uncoiled specimens. Djeddilia djeddilia is probably a synonym of Potamides conicus and in consequence Djeddilia would be another junior synonym of Potamides.

RIASSUNTO Jousseaume, nel corso della sua lunga carriera di malacologo, ha descritto molti taxa problematici di difficile interpretazione. Nel corso del presente lavoro vengono discusse le relazioni fra un enigmatico genere del Mar Rosso, Djeddilia, e il genere pliocenico Escoffieria. L’olotipo di Djeddilia djeddi- lia Jousseaume, 1894, fu rinvenuto a Djedda o Jiddah (Coste del Mar Rosso, Arabia), ed è caratterizzato da giri disgiunti e da una conchiglia par- zialmente svolta. La stessa morfologia caratterizza anche il genere Escoffieria Fontannes, 1880 (specie tipo: Escoffieria fischeri Fontannes, 1880), descritto per i depositi pliocenici della Francia sud orientale. Sia Djeddilia che Escoffieria (che è sinonimo più giovane di Potamides) furono istituiti su esemplari anormali, a conchiglia parzialmente svolta. Djeddilia djeddilia è probabilmente un sinonimo di Potamides conicus e, di conseguenza, Djeddi- lia sarebbe un altro sinonimo di Potamides. Altre specie di Potamididae, a partire dal genere miocenico Tympanotonos fino ai recenti Potamides e Batilla- ria possono presentare spettacolari deformazioni della conchiglia probabilmente causate, durante la crescita della stessa, da fenomeni di stress ambientale (ad esempio crisi anossiche) molto frequenti in un ambiente variabile quale quello occupato da questa famiglia di gasteropodi.

Muséum national d’Histoire naturelle, Lab. B.I.M.M., 55 rue de Buffon, 75005 Paris, France E-mail: [email protected]

INTRODUCTION deformed specimens, among which some specimens have partially Various problematic genera or have been described by uncoiled whorls. COSSMANN & PEYROT (1922) assigned these speci- Jousseaume. However KASE & VALDÉS (1997) have shown that mens to the genus Escoffieria (as Escoffieria degrangei Cossmann & Bertinia bertinia Jousseaume, 1883 is based on the spatula shell Peyrot, 1922). We collected specimens very similar to E. degrangei layer of the patellogastropod Cellana nigrolineata (Reeve, 1854). (Fig. 8) that belong without doubt to the species Tympanotonos VAN AARTSEN (2001) focused our attention on an other enigmat- tournoueri (Mayer, 1878) (see LOZOUET et al. 2001, pl. 9, figs 7-9). ic gastropod described by JOUSSEAUME (1894) from Djedda or The genus Escoffieria Fontannes, 1880 (type species: Escoffieria fis- Jiddah (Red Sea coast, Arabia). The specimen is the holotype of cheri Fontannes, 1880) has been described from Pliocene deposits Djeddilia djeddilia Jousseaume, 1894, which is characterised by of southeastern France. As indicated by Fontannes the unique(?) disjointed whorls and a partially uncoiled shell. In the original specimen of Escofferia fischeri (Figs 1-2, 7) was collected from a bed description the systematic position of Djeddilia djeddilia was not that contains principally Potamides species [supposed to be P. gra- indicated. However, JOUSSEAUME (1894) compared it with the nosus (Borson, 1821) and P. graecus (Deshayes, 1832)]. In the opin- genus Velainella (Trochidae) and with Columna Perry, ion of FONTANNES (1879-1882), Escoffieria is a “section” of 1811 (Achatinidae). LAMY (1932) evoked a distant similarity Potamides characterised by an uncoiled shell. Escoffieria degrangei is with Seguenzia laxa Jeffreys, 1885 (which has the shape of very similar to E. fischeri (see Figs 7-8) and both are based on species of Skeneidae or Haloceratidae) or with Pyrgulina (?) abnormal specimens of Potamididae (Potamides and Tympanotonos). (Pyramidellidae). VAN AARTSEN (2001) excluded a position in In turn, Djeddilia djeddilia closely resembles a broken specimen of the Pyramidellidae and suggested a relationship with the family Escoffieria (Figs 5-6, 7, 8). As indicated by LAMY (1932), the col- Vermetidae. In fact, that author did not realizes that, as already umellar part is “divisé en deux lèvres par un sillon longitudinal” indicated by Lamy, this species was probably based on an abnor- (divided in two lips by a longitudinal groove), it is clearly not at mal specimen. In this paper, the systematic position of Djeddilia all a columellar fold but the consequence of the shell defomation. is revaluated after comparison with fossil Potamididae. One Recent species of Potamididae, generally identified as Pirenel- la conica (Blainville, 1826), is extremely abundant on the Red Sea RESULTS shores. LOZOUET (1986) showed that Gray, 1847 (type Recently, in our monograph of the fauna of the Aquitanian Stage species: Cerithium conicus Blainville, 1826) is a junior synonym of (LOZOUET et al. 2001), we discussed the position of various species Potamides Brongniart, 1810, and that the Mediterranean and Red of Potamididae. We noted that the especially large collections of Sea species Potamides conicus (= Cerithium cailliaudi Potiez & Miocene Tympanotonos specimens, generally include a few very Michaud, 1833) is very similar to the Pliocene species P. graecus

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(Deshayes, 1832). A close relationship with the type species of CONCLUSION Potamides (P. lamarcki Brongniart, 1810 from the Oligocene of These examples reveal that different species of the genera Tympan- France) was also pointed. Potamides conicus is well known for this otonos and Potamides may have spectacular, recurrent deformations remarkable tolerance of temperature and salinity fluctuations and in their shell growth that are probably a consequence of stress in a exhibits a high variability of the shell sculpture (PLAZIAT 1989). fluctuating environment. Thus, FUROTA et al. (2002) have shown Potamides conicus occurs at Djedda, and in the Jousseaume Collec- similar deformations of the shells of potamids snail mul- tion (in MNHN) there is at least one sample of shells with a small tiformis (Lischke, 1869) in severe hypoxia habitats. size, similar to that of the holotype of Djeddilia djeddilia. Another Morphs with disjointed whorls and partially uncoiled shells are numerous sample of Potamides conicus from Djibouti, also similar in certainly the most fascinating monsters, and it is not surprising size to the Djedda specimens, is present in the Jousseaume Collec- that they received different generic names. The unique specimen tion. Some specimens from Djibouti (Fig. 3) show a slight defor- of Djeddilia djeddilia has been collected on a beach and is very mation leading to a more slender and distorted apex. abraded. We may therefore infer that Djeddilia djeddilia is likely to be founded on an abnormal, incomplete, partially uncoiled speci- men of Potamides conicus (Blainville). We may accordingly conclude that Djeddilia Jousseaume, 1894 and Escoffieria Fontannes, 1880 are simply synonyms of Potamides Brongniart, 1810.

ACKNOWLEDGEMENTS I am grateful to Alan Beu (Institute of Geological and Nuclear Sciences, New Zealand) and Jean-Claude Plaziat (Université Paris- Sud) for improving the text and useful suggestions. Thanks also to Raphael La Perna (Università di Bari) for comments.

REFERENCES COSSMANN M. & PEYROT A., 1922. Conchologie Néogénique de l'Aquitaine. Actes de la Société Linnéenne de Bordeaux, 4(1) : 1-322, pls 1-7.

FUROTA T., SUNOBE T. & ARITA S., 2002. Contrasting populations status between the planktonic and direct-developing batillariid snails Batillaria multiformis (Lis- chke) and B. cumingi (Crosse) on an isolated tidal flat in Tokyo Bay. Venus, 61(1-2) : 15-23.

FONTANNES F., 1879-1882. Les mollusques pliocènes de la vallée du Rhône et du Roussillon. Tome premier. Gastéropodes des formations marines et saumâtres. Georg, Figs 1-8, Escoffieria, Potamides, Tympanotonos and Djeddilia species. Lyon., 277 pp., 12 pls.

Figg. 1-8, specie di Escoffieria, Potamides, Tympanotonos e Djeddilia. JOUSSEAUME F., 1894. Diagnose des coquilles de nouveaux mollusques. Bulletin de la Figs 1-2, 7, Escoffieria fischeri Fontannes [= ?Potamides granosus (Borson)], Pliocene (France); by Société Philomatique de Paris, 6(8): 98-105. Fontannes (1880, pl. 10, fig. 19), height: 16 mm. Figg. 1-2, 7, Escoffieria fischeri Fontannes [= ?Potamides granosus (Borson)], Pliocene (Francia); KASE T. & VALDÉS A., 1997. The enigma of Bertinia bertinia Jousseaume, 1883 solved. da Fontannes (1880, tav. 10, fig. 19), altezza: 16 mm. Venus, 56(3): 233-240. Fig. 3, Potamides conicus (Blainville), Red Sea (Djibouti), coll. Jousseaume (MNHN-Paris), spe- LAMY E., 1932. Note sur le Djeddilia djeddilia Jousseaume (Moll. Gastéropode). Bulletin cimen with a slightly distorted early spire, height: 3.2 mm. Fig.3, Potamides conicus (Blainville), Mar Rosso (Gibuti), coll. Jousseaume (MNHN-Parigi), du Muséum national d’Histoire naturelle, (1931), (2)3(8) : 740-743, 1 fig. esemplare con le spire iniziali leggermente distorte, altezza: 3,2 mm. LOZOUET P., 1986. Redéfinition des genres Potamides et Pirenella (Gastropoda : Proso- Fig. 4, Tympanotonos tournoueri (Mayer), Aquitanian Stage, Saucats (Lariey) (France), coll. branchia) à partir des espèces actuelles et fossiles. Implications phylétiques et MNHN-Paris, height: 11 mm. biogéographiques. Annales de Paléontologie, 72(3) : 163-210, 15 figs, 3 pls. Fig. 4, Tympanotonos tournoueri (Mayer), Aquitaniano, Saucats (Lariey) (Francia), coll. MNHN- Parigi, altezza: 11 mm. LOZOUET P., LESPORT J.F. & RENARD P. , 2001. Révision des Gastropoda (Mollusca) du Figs 5-6, Djeddilia djeddilia Jousseaume, 1894, holotype (6 by Lamy, 1932) [= ?Potamides Stratotype de l'Aquitanien (Miocène inf.): site de Saucats "Lariey", Gironde, conicus (Blainville)], Red Sea (Djedda, Arabia), coll. Jousseaume (MNHN-Paris), height: 3.3 France. Cossmanniana, Hors-série n°3: 189 pp., 1 tab. , 19 figs, 37 pls. mm. Figg. 5-6, Djeddilia djeddilia Jousseaume, 1894, olotipo (6 in Lamy, 1932) [= ?Potamides coni- PLAZIAT J.C., 1989. Signification écologique et paléogéographique des peuplements cus (Blainville)], Mar Rosso (Gidda, Arabia Saudita), coll. Jousseaume (MNHN-Parigi), altez- oligotypiques de Potamides gastéropodes thalassiques. In: Atti 3° Simposio di za: 3,3 mm. Ecologia e Paleoecologia delle communità benthoniche, Catania - Taormina 12-16 Fig. 8, Escoffieria degrangei Cossmann & Peyrot, 1922 [= Tympanotonos tournoueri (Mayer)], Ottobre 1985. 25-52, 11 figs, 1 tab. Lower Miocene (Aquitanian), Saucats (Lariey), France, coll. MNHN-Paris, height: 6.5 mm. VAN AARTSEN J.J., 2001. On the enigmatic Djeddilia djeddilia Jousseaume, 1894 (Gas- Fig. 8, Escoffieria degrangei Cossmann & Peyrot, 1922 [= Tympanotonos tournoueri (Mayer)], tropoda, Caenogastropoda, ?Vermetidae). Basteria, 65(4-6): 14-146, 2 figs. Miocene Inferiore (Aquitaniano), Saucats (Lariey), Francia, coll. MNHN-Parigi, altezza: 6,5 mm.

Lavoro accettato il 15 Gennaio 2003

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