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Emmericia lucana n. sp. () from the Middle Pleistocene of the Mercure Basin (southern Italy)

Daniela Esu, Odoardo Girotti & Emanuela Pisegna Cerone

D. Esu, Dipartimento di Scienze della Terra, Università “Sapienza”, P.le A. Moro 5, I-00185 Roma, Italy; [email protected] O. Girotti, Dipartimento di Scienze della Terra, Università “Sapienza”, P.le A. Moro 5, I-00185 Roma, Italy; [email protected] E. Pisegna Cerone, Via Paolo Rembrandt 22, I-20148 Milano, Italy; [email protected]

KEY WORDS - Freshwater gastropods, Emmericiidae, Systematics, Southern Italy, Middle Pleistocene.

ABSTRACT - The intramontane Mercure Basin occupies a broad area within the Basilicata region (southern Apennines, Italy) located between Mt. Pollino to the SE and the Lauria Mts to the NW. It comprises non-marine deposits varying from coarse fluvial gravels and sands at the base, to fine, lacustrine, calcareous muds at the top. Large mammals, freshwater molluscs and tephrostratigraphical analyses indicate a Middle Pleistocene age for these deposits. A new extinct and endemic , Emmericia lucana n. sp., representative of the Family Emmericiidae (Caenogastropoda, ), was found among the rich malacological assemblages of the lacustrine layers and is herein described. It is the first record of the genusEmmericia Brusina, 1870 in the Middle Pleistocene of Italy.

RIASSUNTO - [Emmericia lucana n. sp. del Pleistocene Medio del Bacino del Mercure (Italia meridionale)] - Il bacino intramontano del Mercure occupa in Basilicata una vasta area compresa tra il massiccio del Pollino a SE ed i monti di Lauria a NW. Si tratta del riempimento pleistocenico di una depressione di origine tettonica, i cui sedimenti passano da ghiaie grossolane e sabbie, alla base, a limi calcarei lacustri fini e laminati al tetto. La loro età medio-pleistocenica è indicata dalla presenza di grandi mammiferi e molluschi dulcicoli, oltreché da analisi tefrostratigrafiche. Una nuova specie estinta ed endemica della Famiglia Emmericiidae (Caenogastropoda, Truncatelloidea), Emmericia lucana n. sp., è stata rinvenuta nelle ricche associazioni malacologiche dei limi lacustri. Si tratta del primo rinvenimento del genere Emmericia Brusina, 1870 nel Pleistocene Medio italiano.

INTRODUCTION freshwater molluscan assemblages (Cavinato et al., 2001; Mancini et al., 2004; Esu & Pisegna Cerone, 2008). More The Mercure Basin, located NW of the Pollino Mt. recently, the biochronological datum was confirmed by and SE of the Lauria Mts close to the Calabria-Basilicata tephrostratigraphical analyses (40Ar/39Ar dating, 87Sr/86Sr boundary, is a southern Apennine intramontane basin and 143Nd/144Nd isotope ratios) performed on the tephra of tectono-sedimentary origin, wide about 80 km2., at layers, allowing to refer the study succession to the late around 300-500 m a.s.l. It is incised by the Mercure River Marine Isotope Stage (MIS) 15-12 interval (560-440 ka) and filled with a non-marine Pleistocene stratigraphic (Giaccio et al., 2014; Petrosino et al., 2014; Robustelli et succession, not less than 300 m-thick and subdivided into al., 2014), confirming the Middle Pleistocene age. Field two main stratigraphic sub-units (Mancini, 2004; Mancini investigations for palaeontological purposes, carried out et al., 2004; Giaccio et al., 2014). Mainly coarse gravel by the authors in the whole Mercure Basin during the years deposits passing to fine sands, silts and lignite make up 1996-2003, led to record a noteworthy array of freshwater the lower sub-unit, mostly cropping out in the central gastropods and bivalves from several stratigraphical part of the basin (Calorie, Fig. 1). This sub-unit yielded sections of the upper sub-unit (Pisegna Cerone, 2007; a well-preserved middle Galerian mammal assemblage Esu & Pisegna Cerone, 2008). The recorded assemblages (Cavinato et al., 2001), and rare fluvio-lacustrine molluscs are dominated by aquatic prosobranchs, and locally by of low-energy environment, such as few remains of bivalves. leachii (Sheppard, 1823) with opercula, Valvata piscinalis (Müller, 1774) and Pisidium sp. The alluvial deposits of the lower sub-unit pass upward, in the whole MATERIALS AND METHODS basin, to whitish-yellowish finely laminated calcareous lacustrine mud (varve-like) of the upper sub-unit, up to The new species described in this paper was recovered 100 m thick. Thin tephra layers and abundant freshwater from different sites and distinct layers of the following lacustrine molluscs characterize it. These deposits of open stratigraphical sections of the upper lacustrine sub-unit: to marginal lacustrine environment extensively crop out CIC, VA, S1, S4, LS in the western sector of the basin, and in the basin (Mancini et al., 2004; Pisegna Cerone, 2007; S2, MES1, MES2, MES3 in the eastern sector (Fig. 1). A Esu & Pisegna Cerone, 2008). The sedimentary infilling of detailed sampling was carried out for each section. All the the basin was referred to the Middle Pleistocene by means beds yielded numerous fossil remains, such as molluscs of the record of well-preserved large mammal assemblage (shells and opercula), ostracods and terrestrial plant related to the middle Galerian Large Mammal Age remains. A total of 361 specimens of Emmericia lucana (700-550 ka), and of the presence of Middle Pleistocene n. sp. was collected. The malacological material studied

ISSN 0375-7633 doi:10.4435/BSPI.2016.03 ii Bollettino della Società Paleontologica Italiana, 55 (1), 2016

Fig. 1 - Geological map of the Mercure Basin and location of the investigated stratigraphic sections. 1) Alluvial fan gravels and sands (Middle Pleistocene); 2) lacustrine whitish calcareous muds with tephra layers and molluscs (upper sub-unit, Middle Pleistocene); 3) fluvial gravels and sands with large mammal remains (lower sub-unit, late Lower-Middle Pleistocene); 4) Sant’Arcangelo Basin (Middle-Upper Pliocene); 5) Frido Unit (Cretaceous-Eocene); 6) Pollino Unit (Triassic-Miocene); 7) normal and transtensional faults; 8) CIC, VA, S1, S4, LS, S2, MES1, MES2, MES3, stratigraphic sections (from Cavinato et al., 2001 and Giaccio et al., 2014, modified). in this paper is stored at the Museo di Paleontologia Etymology - From Lucania, an ancient district of dell’Università “Sapienza” di Roma (collection Esu- southern Italy, extending from the Tyrrhenian Sea to Girotti, MPUR7). the Gulf of Taranto, comprising the modern region of Gastropod follows that of Bouchet & Rocroi Basilicata. (2005), and the FreshGen database (Neubauer et al., 2014). Locus typicus and stratum typicum - Castelluccio inferiore (Basilicata), 40°00’03,44”N, 15°58’53,05”E; SYSTEMATIC PALAEONTOLOGY calcareous silt and marls of the upper sub-unit of the Mercure Basin, Middle Pleistocene. Class Cuvier, 1795 Subclass Caenogastropoda Cox, 1960 Type material - Holotype, MPUR7-3291 (Section Order Golikov & Starobogatov, 1975 CIC); Paratypes, MPUR7-3287/2. Superfamily Truncatelloidea Gray, 1840 Family Emmericiidae Brusina, 1870 Additional material - CIC, MPUR7-2111/77; VA (Vallecola), MPUR7-3268/1; S1, MPUR7-3283/25; S4, Emmericia Brusina, 1870 4 lost specimens; LS (Laino strada), MPUR7-4114/4; S2, MPUR7-2287/5, MPUR7-2303/1; MES1, MPUR7- Type species Paludina patula Brumati, 1838, Recent, 2109/9; MES2, MPUR7-4115/2; MES3, MPUR7- Monfalcone (Gorizia). 3231/227, MPUR7-3115/3.

Emmericia lucana n. sp. Age - The new species is known only from the Middle (Pl. 1, figs 1-7) Pleistocene of southern Italy (Mercure Basin, Basilicata).

2008 Emmericia cf. narentana Bourguignat - Esu & Pisegna Diagnosis - A species of the genus Emmericia, with Cerone, p. 152. conical-elongated and high spired shell; apex large, flat, D. Esu et alii - Emmericia lucana n. sp. from the Middle Pleistocene of Italy iii obliquely truncated; protoconch smooth, with 1.5 whorls; E. expansilabris Bourguignat, 1880 and E. ventricosa five whorls nearly flattened, axially striated, provided Brusina, 1870, which appear smooth and without varix, or not with a carina; ovate, peristome weakly although both can show a very weak carina on the last expanded, slightly reflected; last provided or not whorl (Radoman, 1967). E. lucana n. sp. differs from E. with an axial swelling (varix) behind outer lip. Umbilicus expansilabris in having tapered shell, less convex whorls, small, nearly closed. slightly reflected peristome margins, and in some cases an evident carina on all the whorls; from E. ventricosa Description - Shell small-sized, conical-elongated, in being more slender and elongated, higher-spired, with high spired, tapered, with five whorls slightly convex less convex whorls and a proportionally smaller last to nearly flattened, provided or not with a thin carina whorl, and in having an evident carina on all the whorls running in the lower part of the whorls, nearly close in some cases. Among fossil species, the only described to the , and in the middle of the last whorl. Last carinate form is Emmericia multicarinata Rust, 1997, whorl large, convex, high about 2/3 of total height, from the Pliocene of Greece (Rust, 1997), which differs somewhat descending in some specimens, mainly in the from E. lucana n. sp. in being smaller and bearing up carinate form. Suture weakly incised and linear. Apex to five carinae. Both E. krizanici Brusina, 1902 and E. large, flat, slightly oblique and protruding laterally. zivkovici Brusina, 1902, from the Pliocene of Slavonia Protoconch smooth, with 1.5 whorls, diameter of first and Serbia, respectively, show an angulation on the last whorl 1 mm, height 0.6 mm, width of initial cap 0.26 whorl (Brusina, 1902, pl. 11, figs 33-39). They differ from mm, teleoconch with regular, dense and slightly raised the Mercure new species in having a wider conical shape, axial striae. Aperture more or less wide, ovate, slightly flatter whorls, and a proportionally higher last whorl. oblique and weakly concave in the adapical part, peristome Pyrgula (Pyrgidium) nodotiana Tournouër, 1866 from the weakly expanded, nearly reflected, partially covering the Pliocene of Bligny-les-Beaune (Côte d’Or) (Tournouër, umbilicus. A weak axial swelling behind the outer lip is 1869, pl. 3, fig. 2) is very similar in shape toE. lucana n. visible in non-carinate shells, or missing in the carinate sp. Comparisons with specimens of P. nodotiana from the form. Umbilicus small, nearly closed. Dimensions (mean): type area, made at the Senckenberg Museum of Frankfurt H = 5 mm, W = 4 mm; max observed: H = 7 mm, W = 4.0 a. M., showed remarkable differences in detail, despite mm. Dimensions of the holotype: H = 6.2 mm, W = 4 mm; the general aspect. The apex of P. nodotiana looks like Aperture: H = 3 mm, W = 2.2 mm (including peristome). that of Emmericia but it is smaller and more elevated than that of E. lucana n. sp.; the whorls are rather flattened and Comparisons and discussion - The numerous bear a sharper and more prominent carina; the aperture specimens, recorded from distinct stratigraphical sections is angulated adapically and, laterally, in correspondence at different sites of the basin, are almost all incomplete, with the carina. lacking aperture or part of the apex, which is corroded The whole of the molluscan fauna from the upper and sometimes reconstructed. Nevertheless, the typical sub-unit of the Mercure Basin is highly diversified, being characters of the genus Emmericia, as flat apex (Pl. 1, figs composed of many species with generally abundant 1, 4, 6) and varix behind the aperture (Pl. 1, figs 1, 3), are and well-preserved specimens of freshwater gastropods preserved in some cases. The recorded populations show a and bivalves, such as Theodoxus fluviatilis (Linnaeus, high degree of intraspecific variability due to the presence 1758), Viviparus contectus (Millet, 1813), B. leachii, or lack of a carina, and to the presence or absence of the Belgrandia latina (Settepassi in Settepassi & Verdel, axial swelling (varix) behind the outer lip. Most of the 1965), E. lucana n. sp., Tanousia subovata (Settepassi specimens lack a swelling, which is recognizable only in Settepassi & Verdel, 1965), Valvata cristata Müller, in non-carinate specimens. Intermediate forms show an 1774, V. piscinalis, Radix auricularia (Linnaeus, 1758), angulated last whorl (Pl. 1, Fig. 2). The presence of the R. balthica (Linnaeus, 1758), Gyraulus albus (Müller, carina in the Mercure specimens suggested (Esu & Pisegna 1774), Dreissena polymorpha (Pallas, 1771) and Cerone, 2008) a possible identity with one of the four Pisidium amnicum (Müller, 1774), the most of which known extant species of Emmericia from the Dinaride are characteristic of clear lacustrine or slowly running region, E. narentana Bourguignat, 1880, which is strongly freshwater (Ložek, 1964; Pisegna Cerone, 2007). The carinate (Radoman, 1967, 1983). Nevertheless, the new very common Middle Pleistocene lacustrine T. subovata species differs in having a more slender and tapered accompanies E. lucana n. sp. in several layers, together shell, less reflected and less robust apertural margins, a with B. latina, V. piscinalis, D. polymorpha, and locally weaker carina (when present), and a weaker to missing T. fluviatilis, indicating oxygenated conditions of the varix behind the outer lip. Likewise, the smooth or only water-body and a temperate climate (Esu & Pisegna angulated fossil Mercure specimens are distinguished Cerone, 2008). The pollen record from the Mercure from the non-carinate E. patula (Brumati, 1838), which is lacustrine sediments, attributed to MIS 13 by Petrosino et provided, in rare cases, with a weak keel on the last whorl al. (2014), supports the assumed temperate palaeoclimatic and bears a prominent varix (Radoman, 1967). The new indication given by the molluscan assemblages. The species is more slender, with higher spired and tapered co-occurrence of E. lucana n. sp. with species of well shell, with less reflected and less robust aperture margins, oxygenated and hard-waters confirms the preference of a weaker (when present) to missing varix behind the outer this species for similar environment, like that of its four lip, minor size and smaller aperture (shell dimensions max known congeners inhabiting the Balkan fluvial system observed: E. patula, H = 9.5 mm, W = 6.5 mm [Radoman, (Radoman, 1967, 1983). The considerable variability of 1967]; E. lucana n. sp., H = 7 mm, W = 4.0 mm). The the new species is probably due to different physical and new species differs also from other two living species, chemical conditions occurred in the water-body during the iv Bollettino della Società Paleontologica Italiana, 55 (1), 2016 sediment deposition. Indeed, carinate specimens prevail Treatise on Invertebrate Paleontology. Part I, 1. The in the whitish-yellowish, finely laminated, calcareous Geological Society of America and University of Kansas Press, lacustrine mud (varve-like) of the stratigraphical sections Lawrence: I310-I324. (MES1, MES2, MES3), and less carinate or smooth are Cuvier G. (1795). Second Mémoire sur l’organisation et les rapports des animaux à sang blanc, dans lequel on traite de la from marl and calcareous clayey silt (for ex., section structure des Mollusques et de leur division en ordre. Magasin CIC). Anyway, nearly smooth, intermediate and more Encyclopédique ou Journal des Sciences, des Lettres et des or less carinate forms can be recorded in the same layer, Arts, 2: 433-449. indicating to belong to the same species. The new species Esu D., Girotti O. & Truc G. (2001). New data on fossil here described displays so far an endemic character. It is Emmericiinae from Italy and France (Gastropoda: Prosobranchia: the only recorded Emmericia from the Middle Pleistocene Emmericiidae). Archiv für Molluskenkunde, 129: 123-143. of Italy; no records are known for the Late Pleistocene Esu D. & Pisegna Cerone E. (2008). Tanousia subovata (Settepassi) and Holocene of Italy so far, whilst some species are (), a Middle Pleistocene correlation element for the spread from the Late Miocene to the Early Pleistocene central-southern Italian lacustrine basins. Suppl. Atti Museo Civico di Storia Naturale di Trieste, 53: 147-155. of the northern and central Italian Peninsula (Esu et al., Giaccio B., Galli P., Peronace E., Arienzo I., Nomade S., Cavinato 2001). G.P., Mancini M., Messina P. & Sottili G. (2014). A 560-440 ka tephra record from the Mercure Basin, southern Italy: volcanological and tephrostratigraphic implications. Journal ACKNOWLEDGEMENTS of Quaternary Science, 29: 232-248. Golikov A.N. & Starobogatov Y.I. (1975). Systematics of Research fund of Ateneo Sapienza of Rome (D. Esu-Project prosobranch gastropods. Malacologia, 15: 185-232. “Biostratigraphy of Middle Pleistocene lacustrine basins of Italy”) Gray J.E. (1840). Shells of molluscous . In Synopsis of the supported this work. Many thanks are due to L. Di Pietro and M. contents of the British Museum. G. Woodfall, London: 105-152. Salvati (Laboratorio cartografico, Sapienza Università di Roma) for Linnaeus C. (1758). Systema naturae per regna tria naturae, help in arrangement of figures. We are grateful to the two reviewers secundum classes, ordines, genera, species, cum characteribus, Mathias Harzhauser and Thorsten Kowalke for their constructive differentiis, synonymis locis. 1. 824 pp. Laurentius Salvius, comments. Holmiae. Ložek V. (1964). Quartärmollusken der Tschechoslowakei. Rozpravy Ustredniho Ustavu Geologického, 31: 1-368. REFERENCES Mancini M. (2004). Stratigrafia integrata delle successioni plio- pleistoceniche dei Bacini di Sant’Arcangelo e del Mercure Bouchet P. & Rocroi J.-P. (2005). Classification and Nomenclator (Basilicata). Relazione finale per l’assegno di ricerca, settore of Gastropod Families. Malacologia, 47: 1-397. scientifico-disciplinare GEO/01, Dipartimento di Scienze della Bourguignat J.-R. (1880). Monographie du genre Emmericia. 87 Terra, Università di Roma “La Sapienza”. 12 pp. pp. Lachèse et Dolbeau, Angers. Mancini M., Cavinato G.P., Esu D., Girotti O. & Zanchetta G. Brumati L. (1838). Catalogo sistematico delle conchiglie terrestri (2004). Pleistocene alluvial and lacustrine successions in the e fluviatili della provincia di Monfalcone. 56 pp. Patermolli, intramontane Mercure and Sant’Arcangelo Basins (southern Gorizia. Appenines, Italy): sedimentary evolution and biostratigraphic Brusina S. (1870). Monographie der Gattungen Emmericia und constraints. Abstract (part 1), 32nd International Geological Fossarulus. Verhandlungen der kaiserlichen und königlichen Congress, Florence 2004: 388. zoologisch-botanischen Gesellschaft in Wien, 20: 925-938. 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EXPLANATION OF PLATE 1 Emmericia lucana n. sp. Fig. 1 - Holotype MPUR7-3291; frontal (a), lateral (b) and posterior (c) view; Section CIC. Fig. 2 - Paratype MPUR7-3287/2; frontal view. Fig. 3 - Paratype MPUR7-3287/2; frontal (a), lateral (b) and postero-lateral (c) view. Fig. 4 - MPUR7-3115/3; frontal view; Section MES3. Fig. 5 - MPUR7-3115/3; frontal (a), lateral (b) and posterior (c) view; Section MES3. Fig. 6 - MPUR7-3115/3; a) first whorls, b) protoconch; Section MES3. Fig. 7 - MPUR7-2303/1; frontal (a), lateral (b) and posterior (c) view; Section S2. All scale bars correspond to 1 mm. D. Esu et alii - Emmericia lucana n. sp. from the Middle Pleistocene of Italy Pl. 1v vi Bollettino della Società Paleontologica Italiana, 55 (1), 2016

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