Microsoft Word - 00_indice_LM04.docx Il Quaternario IT ISSN 039-3356 Italian Journal of Quaternary Sciences 24 (1), 2011 - 67-74 JAMINIA (JAMINIA) MALATESTAE ESU, 1988 (MOLLUSCA, GASTROPODA, ENIDAE) FROM THE MIDDLE AND LATE PLEISTOCENE OF CENTRAL-SOUTHERN ITALY. PALAEOECOLOGICAL IMPLICATIONS Carmine D’Amico1 & Daniela Esu2 1Dipartimento STAT, Università degli Studi del Molise, Pesche (Isernia) 2Dipartimento di Scienze della Terra, Sapienza Università di Roma Sapienza, IGAG - CNR, Roma Corresponding author: D. Esu ABSTRACT: D’Amico C. & Esu D., Jaminia (Jaminia) malatestae ESU, 1988 (Mollusca, Gastropoda, Enidae) from the Middle and Late Pleistocene of Central-Southern Italy. Palaeoecological implications. Jaminia (Jaminia) malatestae is a fossil land gastropod of the family Enidae. The species described by ESU (1988) was dedicated to the Quaternarist geologist and palaeontologist Alberto Malatesta who recognized, at the base of the Ponte Galeria formation (Rome), the “Blue-grey Helicella bearing clays”, stratum typicum of the species, dated to the early Middle Pleistocene. J. (J.) malatestae is an endem- ic species of central-southern Italy ranging from the early Middle Pleistocene to the Late Pleistocene. It was recovered from deposits with oligotypical assemblages of terrestrial gastropods and small mammals of cold-arid climate. During the Middle Pleistocene the species occurred in Lazio, at Fontignano and San Cosimato (Ponte Galeria and Aurelia formations, Rome) in an oligotypic molluscan assem- blage of cold climate and reworked in a politypic assemblage of temperate-warm climate respectively, and in Abruzzo, at Case Picconet- to (Pescara) in a palaeosol of an alluvial deposit, associated with non-marine mollusc species typical of open environments and cold climate. In the Middle and/or Late Pleistocene the species occurred in Marche, at Gola della Rossa (Ancona) in a cave-fill deposit with oligotypic molluscs of mountain grassland, and at Polesio (Ascoli Piceno) in a sandy layer interbedded to gravels of the last glacial pe- riod, associated with gastropods of open environment and cold climate. In the Late Pleistocene (last gacial period) the species occurred in Campania with oligotypic molluscan assemblage of cold climate from a loess deposit at Telese (Calore River valley) and from a sandy layer interbedded to gravels at Tufino (Naples). Both the structure of the molluscan assemblages and the autoecology of the accompa- nying species suggest that J. (J.) malatestae populated open-dry palaeoenvironments in the central-southern Italian peninsula during cold climatic periods of the Middle and Late Pleistocene. Its record in non-marine Quaternary deposits of Italy is an useful tool to infer reliable palaeoenvironmental and palaeoclimatical information. RIASSUNTO: D’Amico C. & Esu D., Jaminia (Jaminia) malatestae ESU, 1988 (Mollusca, Gastropoda, Enidae) del Pleistocene medio e superiore dell’Italia centro-meridionale. Implicazioni paleoecologiche. Jaminia (Jaminia) malatestae è una specie di gasteropode terrestre della Famiglia Enidae conosciuta solo allo stato fossile. La specie, descritta da ESU nel 1988 su materiale fossile proveniente dalle “argille ad Helicella” del Pleistocene medio-inferiore affioranti a Ponte Galeria (Roma), è stata dedicata dall’autrice ad Alberto Malatesta, paleontologo e geologo del Quaternario, che per primo individuò le “argille ad Helicella”, stratum typicum della specie, alla base della formazione di Ponte Galeria. La distribuzione cronostratigrafica di J. (J.) malatestae, che risulta essere una specie endemica dell’Italia centro-meridionale, va dal Plei- stocene medio inferiore al Pleistocene superiore. Generalmente si rinviene in associazioni malacofaunistiche e a micromammiferi indica- trici di ambienti aperti e condizioni climatiche fredde ed aride. Nel Pleistocene medio è nota nel Lazio, a Ponte Galeria e, frammentaria, nella formazione Aurelia (Roma) come elemento non autoctono di un’associazione a molluschi terrestri e dulcicoli di clima temperato- caldo, in Abruzzo (Case Picconetto, Pescara) in un paleosuolo di un deposito alluvionale (età ca 0.48 Ma) associata a molluschi terrestri di ambiente aperto e clima freddo. Nelle Marche è nota a Gola della Rossa (Fabriano, Ancona) in un’ associazione oligotipica a mollu- schi terrestri di prateria montana in depositi di riempimento di una grotta riferiti al “Riss” finale o “Würm” iniziale, e a Polesio (Ascoli Pice- no), in un deposito sabbioso intercalato a conglomerati riferiti all’ultimo glaciale, associata a specie terrestri di ambiente aperto e clima freddo. In Campania è stata rinvenuta in associazioni oligotipiche indicatrici di clima freddo-arido in un deposito di loess appartenente al “terrazzo inferiore” del fiume Calore (Benevento), attribuito al Pleistocene superiore (“Würm”), e in sabbie intercalate a ghiaie riferite allo stadio isotopico 2 nei pressi di Tufino (Napoli) (valle del torrente Clanio-Acqualonga). Il carattere generalmente oligotipico delle associazioni in cui J. (J.) malatestae è presente e l’autoecologia delle singole specie che l’accompagnano suggeriscono che questa specie popolava i paleoambienti privi di vegetazione arborea e relativamente aridi nell’Italia centro-meridionale durante le fasi climatiche fredde del Pleistocene medio e superiore, e ne fanno un elemento utile per significative indicazioni paleoambientali e paleoclimatiche. Key words: Jaminia (Jaminia) malatestae, Enidae, Palaeoecology, Palaeoclimate, Pleistocene, Italy. Parole-chiave: Jaminia (Jaminia) malatestae, Enidae, Paleoecologia, Paleoclima, Pleistocene, Italia. 1. INTRODUCTION The distribution and abundance of terrestrial mol- luscan species are controlled by a number of interacting factors (BARKER, 2001; PFENNINGER & POSADA, 2002; DAVIES, 2008). Among these both temperature and rela- tive humidity constraining the physiology and life cycle of terrestrial molluscs influence the differing geographi- cal distribution of the species. Each species has a ther- mal interval controlling its own physiological functions C. D’Amico & D. Esu 68 and outside the ranges the species cannot survive (MOINE et al., 2002 and references therein). These au- thors demonstrated that the mean temperature of the coldest month and annual thermal magnitude are the most important forcing parameters for the distribution of modern as well as Quaternary land mollusc species. Modern terrestrial molluscs are indices both of vegeta- tion type and climate, permitting them to be grouped in ecological classes (BOYCOTT, 1934). Quaternary terre- strial molluscan assemblages are mainly constituted by extant species which do not show significative changes in their ecological requirements at least from the Middle Pleistocene (GLIOZZI et al., 1997; ROUSSEAU et al., 2007). They can be gathered in ecological groups, as well as the living species, and used to infer general var- iations of environmental conditions related to the climat- ic cyclicity of Quaternary glacial and interglacial periods. Among land molluscan assemblages, woodland and steppe/grassland communities provide the major useful data on climatic fluctuations, indicating respectively warm intervals characterized by woodland expansion, and cold phases with decreased arboreal covering (SPARKS, 1961; ID, 1964; LOŽEK, 1964; ID, 2000). A large number of studies that utilized the ecologi- cal patterns of terrestrial molluscs to infer Quaternary palaeoclimatic signatures was published in past dec- ades (for example, LOŽEK, 1964; ROUSSEAU & WU, 1999; ROUSSEAU et al., 2000; WU et al., 2000). Land molluscan assemblages were intensively studied in Qu- aternary loess deposits of Central and Eastern Europe, as well as in North America and in China, where their palaeoecological framework and related palaeoclimatic attributes have been used as useful biostratigraphical tools (ROUSSEAU, 2001). In the Italian peninsula, at the beginning of the Mid- dle Pleistocene, consequently to a general renewal prob- ably related to the onset of the 100 ka orbital cyclicity, the non-marine molluscan assemblages became quite mod- ern (ESU & GIROTTI, 1991; GLIOZZI et al., 1997) and strongly influenced by climatic change: the politypic as- semblages are related to temperate and temperate-warm climatic phase (interglacial), while the oligotypic ones oc- curred during cold oscillations (glacial) (ESU et al., 1989). In this work we give an overview of the geographi- cal (Fig. 1) and stratigraphical distribution of the terre- strial fossil species Jaminia (Jaminia) malatestae ESU based on literature data published over the last 30 years, and highlight its palaeoecological meaning. The palaeoecology of each molluscan assemblage in which the species was recorded is analyzed. Furthermore we report the unpublished occurrence of J. (J.) malatestae in an Upper Pleistocene deposit of the Marche region. 2. SYSTEMATIC REMARKS Phylum Mollusca CUVIER, 1797 Class Gastropoda CUVIER, 1797 Order Stylommatophora SCHMITT, 1855 Family Enidae WOODWARD, 1903 Genus Jaminia RISSO, 1826 Subgenus Jaminia RISSO, 1826 Jaminia (Jaminia) malatestae ESU, 1988 Fig. 2 1980 Chondrula (Jaminia) reversalis - CONATO et al., p. 143 (non BIELZ). 1988 Jaminia (Jaminia) malatestae ESU, p. 228, Fig. 1. 1989 Chondrula reversalis - ESU et al., p. 288 (non BIELZ). 1991 Jaminia (Jaminia) malatestae - ESU & GIROTTI, p. 144. 1992 Jaminia (Jaminia) malatestae - KOTSAKIS et al., p. 336. 1997 Jaminia malatestai - GLIOZZI et al., p. 378. 1998 Jaminia malatestae - DI VITO et al., p. 281, Fig. 9c. 2003 Jaminia malatestai - MARCOLINI et al., p. 548, Fig. 3a. Description (ESU, 1988). Shell cylindro-conical, si- nistral, of moderate size (height 10÷11.5 mm, diameter 4÷4.5 mm, occasionally 5 mm), with 7-71/2 rather flat- sided whorls. Apex large. Sutures weak, faintly slanting. Shell surface with weak, irregular, growth-lines well visi- ble at SEM. Shell rather thick continuing with the same thickness into the mouth-edge. Peristome neither ex- panded nor reflected, with a characteristic sinuous outer profile. Mouth oval, always furnished with only one blunt parietal tooth more or less developed; sometimes a weak lengthened swelling is present along the columel- lar side. Mouth-edge continuous bearing a little callus across the outer parietal area, always present in the specimens with preserved parietal peristome. The type material of J. (J.) malatestae was col- lected from the “Blue-grey Helicella bearing clays” (ear- ly Middle Pleistocene) at Fontignano (Rome) (Figs. 1 and 3). At first it was ascribed to the species “Chondru- la (Jaminia) reversalis (BIELZ, 1853)” by CONATO et al. (1980), then ESU (1988), after direct comparison with specimens of the BIELZ’s species from Eastern Euro- pean Quaternary non-marine deposits, stored in the Geological Institute of Hungary (Magiar Allami Földtani Intézet) at Budapest, formally classified the Fontignano fossil record as J. (J.) malatestae dedicating it to the Fig. 1 - Geographical distribution of J. (J.) malatestae. A) Fonti- gnano and San Cosimato (Rome); B) Case Picconetto (Pesca- ra); C) Gola della Rossa (Ancona); D) Polesio (Ascoli Piceno); E) Telese (Calore River valley, Benevento); F) Tufino (Naples). Distribuzione geografica di J. (J.) malatestae. A) Fontignano e San Cosimato (Roma); B) Case Picconetto (Pescara); C) Gola della Rossa (Ancona); D) Polesio (Ascoli Piceno); E) Telese (valle del fiume Calore, Benevento); F) Tufino (Napoli). Jaminia (Jaminia) malatestae Esu, 1988 (Mollusca, Gastropoda, Enidae) … 69 Quaternarist geologist and palaeontologist Alberto Malatesta, who first described the “Blue-grey Helicella bearing clays” in CONATO et al. (1980). GLIOZZI et al. (1997) reported the species as J. malates- tai. Nevertheless, according to the Art. 31.1.1 of the International Code of Zoo- logical Nomenclature (ICZN) (INTERNA- TIONAL COMMISSION ON ZOOLOGICAL NO- MENCLATURE, 1999), the correct specific name is J. (J.) malatestae as firstly re- ported by ESU (1988). 3. STRATIGRAPHICAL DISTRIBUTION AND ECOLOGICAL FRAMEWORK J. (J.) malatestae ranges from the base of the early Middle Pleistocene1 to the Late Pleistocene. It is endemic of the Italian peninsula and is recorded from few locali- ties of central and southern Italy (Fig. 1). As the fossil status of the species, its ecological requirements were inferred from the autoecology of similar extant species, such as the congeneric Jaminia (J.) quadridens (MÜLLER) to which it is often associate, and from the characteristics of the accompanying species. Generally J. (J.) malates- tae occurs in Quaternary molluscan assemblages indi- cating grassland dried environments and cold climatic conditions. Below we give an overview of the J. (J.) malates- tae occurrences, focusing on the composition and the ecological framework of each related molluscan assem- blage, according to LOŽEK’s (1964) ecological classifica- tion of the Central and Eastern European Quaternary non-marine molluscs (Tab. 1). 3.1 Lazio 3.1.1 Fontignano (Rome) The oldest occurrence of J. (J.) malatestae is registered in the lowermost Middle Pleistocene at Fontignano (Rome), in the “Blue-grey Helicella bearing clays” (Fig. 3), a lower member of the Ponte Galeria formation in which CONATO et al. (1980) distinguished eight mem- bers, from the bottom: 1) river pebble and cobble con- glomerates, 2) blue-grey Helicella bearing clays, 3) beach conglomerates and bright yellow A. islandica sands,4) pebble gravels and sands with frequent cross- laminations, 5) Venerupis senescens clays, 6) aeolian salmon sands, 7) lacustrine and marshy deposits, 8) “tufi grigi inferiori” (earthy and pisolitic tuffs). This strati- graphical succession registers the transitions from non- marine to fully marine and non-marine conditions. More recently MILLI (1997) utilized a sequence-stratigraphy approach in refining the stratigraphy of the Pleisto- cene/Holocene deposits of the Roman area, distinguish- ing two composite third-order sequences, the Monte Mario Sequence (MMS) (Early Pleistocene) and the Ponte Galeria Sequence (PGS) (Middle-Late Pleisto- cene/Holocene). The “Blue-grey Helicella bearing clays”, falling in the Matuyama reversed palaeomagnetic epoch (MARRA et al., 1998), correspond to a lowstand system tract of the lower fourth-order sequence (PG1) of the PGS; it formed the inner portion of a coastal-barrier de- positional system, laterally associated to a braid delta of a fluvial system evolving in a coastal plain with low- sinuosity braided channels (MILLI & PALOMBO, 2005). This environmental interpretation is consistent with the terrestrial oligotypical molluscan assemblage rec- Tab. 1 - Ecological classes (LOŽEK, 1964) of the mollusc spe- cies associated with Jaminia (J.) malatestae. Gruppi ecologici (LOŽEK, 1964) cui appartengono le specie di molluschi associate a Jaminia (J.) malatestae. 1 HEAD et al. (2008; see also OGG et al., 2008) proposed to correlate the Early-Middle Pleistocene boundary with the Matuyama-Brunhes palaeomagnetic Chron boundary (about 773 ka) (MIS 19). At present this proposal is informally accepted by the Subcommission on Qua- ternary Stratigraphy (SQS) of the International Commission on Stratigraphy (ICS), as reported in the “Global chronostratigraphical correla- tion table for the last 2.7 Ma v. 2009” (http://www.quaternary.stratigraphy.org.uk/ charts/; GIBBARD & COHEN, 2008). Nevertheless, in the present work, according to MILLI & PALOMBO (2005), we utilize the ancient boundary of Early-Middle Pleistocene, at transition MIS 25 to MIS 24. C. D’Amico & D. Esu 70 Fig. 3 - A) The “blue-grey Helicella bearing clays” at Fontignano (stratum typicum of J. (J.) malatestae); B) beach conglomerates and bright yellow A. islandica sands (photo: DANIELA ESU). A) “argille grigio-azzurre ad Helicella” a Fontignano (stratum typicum di J. (J.) malatestae); B) conglomerati di spiaggia e sabbie giallo vivo ad A. islandica (foto: DANIELA ESU). orded in the “Helicella clays” (Tab. 2). Granaria fru- mentum (DRAPARNAUD), Helicella Itala (LINNÉ), and very probably J. (J.) malatestae, belong to the ecologic group 4S, including species that live in dry grassland (Tabs 1 and 2). Pupilla muscorum (LINNÉ) (5O) lives in dry exposed places or grassland, Vallonia pulchella (MÜLLER) (5O) prefers slightly humid open grounds, Trochulus hispidus (LINNÉ) (7M) is a widespread mesic species. G. frumentum, P. muscorum and T. hispidus are species more or less frequent in the loess mollus- can assemblages of Central Europe (LOŽEK, 1964). The oligotypy and the dominance of the species re- ferred to the group 4S (Fig. 4) are indicative of a steppe environment and cold-dry climatic conditions (ESU, 1988; KOTSAKIS et al., 1992). This is also con- firmed by the record in the same deposit of two spe- cies of boreal arvicolids (rodents) of Eastern Europe, Prolagurus pannonicus (KORMOS) and Predicrostonix sp., belonging to the steppe lemming and collared lemming respectively (KOTSAKIS et al., 1992). Both the characteristics of the faunal assemblage and the fact that the “Blue-grey Helicella bearing clays” fall in reversed palaeomagnetic epoch led KOTSAKIS et al. (1992) to attribute the inset of the sedimentary cycle of the Ponte Galeria formation immediately after the coldest peak of marine isotope stage (MIS) 22. Never- theless, MARRA et al. (1998) referred the base of the Ponte Galeria formation to MIS 20-19. 3.1.2 San Cosimato (Rome) Only one fragmentary specimen (the last whorl with mouth), very probably reworked, was collected from the diatomitic muds of the Aurelia formation (CONATO et al., 1980; PG6 of MILLI, 1997), in a rich politypic mollus- can assemblage of about forty land and freshwater spe- cies characteristic of temperate-warm climate (ESU, 1988; ESU & GIROTTI, 1991). Based on stratigraphic evidences, radiometric dat- ing, and vertebrate fossil record the Aurelia formation was dated to Middle Pleistocene (MIS 8-10) and the mollusc-bearing diatomitic layers can be referred to MIS 9 likelihood (CONATO et al., 1980; GLIOZZI et al., 1997; MILLI, 1997; MILLI & PALOMBO, 2005). 3.2 Abruzzo 3.2.1 Case Picconetto (Pescara) MARCOLINI et al. (2003) reported J. (J.) malatestae in a silty-clayey palaeosol of a sedimentary succession, Fig. 4 - Palaeoecological features of the analyzed molluscan assemblages. The ecological classes (EC, see Tab. 1) and their relative frequency are reported for each assemblage (Tab. 2). The percentages of the ecological classes for Gola della Rossa and Telese assemblages are not available. Carattere paleoecologico delle associazioni malacologiche ana- lizzate. Sono riportate le classi ecologiche (EC, vedi Tab. 1) e le relative frequenze per ciascuna associazione (Tab. 2). Le per- centuali delle classi ecologiche per le associazioni di Gola della Rossa e Telese non sono disponibili. Jaminia (Jaminia) malatestae Esu, 1988 (Mollusca, Gastropoda, Enidae) … 71 cropping out at Case Picconetto (Pescara), interpreted as a distal part of an old alluvial fan deposit. The two palaeosols recorded in the succession, one of which, the uppermost, holding J. (J.) malatestae, are related to a local stasis in the alluvial fan accretion. A tephra layer dated 0.48 ± 0.04 Ma (apatite fission track dating) lies above the palaeosol containing J. (J.) malatestae. The molluscan assemblage consists mainly of terre- strial taxa (Tab. 2). Only two freshwater species, Gyraulus (Armiger) crista (LINNÉ) and G. (G.) albus (MÜLLER), occur reaching less than 0.3% of the assemblage. The land spe- cies typical of open ground (5O) are dominant, followed by open dry grassland taxa (4S) (Fig. 4). The mesic species (7M) are represented by a discrete number of specimens of Vitrea contracta (WESTERLUND). The strictly forest (1W) and hygrophilous taxa (8H) are very scarce, represented by Sphyradium cf. S. doliolum (BRUGUIÈRE) and Vertigo angustior JEFFREYS respectively, which, according to LOŽEK (1964), are warm-climate species. The molluscs are associated to an oligotypical mi- cromammal assemblage with dominant arvicolids, such as Microtus cf. M. arvalis (PALLAS), M. (Terricola) gr. savii (DE SELYS LONGSCHAMPS), followed by murids, such as Apodemus gr. sylvaticus-flavicollis, and insecti- vores (Crocidura sp.), indicative of open environment and Mediterranean-type climate (MARCOLINI et al., 2003). Together, molluscs and small mammals indicate open ground environment and cold, but not severe, cli- matic conditions. The radiometric dating, the results of chemical analysis carried out on the tephra layer, and the pa- laeoenvironmental indications suggested by molluscs and small mammals, led MAR- COLINI et al. (2003) to correlate the Case Picconetto succession with MIS 14. 3.3 Marche 3.3.1 Gola della Rossa (Ancona) ESU et al. (1990) recorded J. (J.) malatestae from a cave-fill deposit (Grotta del Vento, Gola della Rossa, Ancona) in an oligotypical molluscan assemblage (Tab. 2). Candidula spadae (CALCARA) and Clausilia (Clausilia) rugosa pinii WES- TERLUND, prefer open woodland, but lo- cally can live in open or semi-open habi- tats [2W(S)]. At present, they have Apen- nine distribution being common in the high mountain zone (MANGANELLI et al., 1995; CIANFANELLI, 2009. J. (J.) quadri- dens (4S), Chondrina avenacea (BRU- GUIÈRE) (4Sf) and P. muscorum (5O) po- pulate dry grassland or open grounds. Small mammals and one reptile are asso- ciated with the land molluscs: Sorex anti- norii BONAPARTE, S. minutus LINNÉ, Rhi- nolophus ferrumequinum (SCHREBER), Myotis dasycneme (BOIE), Pliomys lenki cfr. relictus CHALINE, M. arvalis, Chio- nomys nivalis (MARTINS), Apodemus cfr. sylvaticus (LINNÉ), and Hierophis viridifla- vus (LACÉPÈDE) (revised by T. Kotsakis). Both molluscs and mammals indi- cate open and rather dry mountain pa- laeoenvironment, and cold climatic conditions. Based on small mammals the cave-fill deposit of Gola della Rossa can be attributed to the late Middle Pleistocene (MIS 6) or early Late Pleistocene (MIS 4) (ESU et al., 1990). 3.3.2 Polesio (Ascoli Piceno) We report the unpublished occurrence of J. (J.) malatestae in Upper Pleistocene non-marine deposits at Polesio (Ascoli Piceno). In a recent work BUCCOLINI et al. (2010) (see also GENTILI et al., 1998), analizing the geomorphological evolution of the last 20 ka in the high hill sector (600- 1100 m a.s.l.) of Mt. Ascensione area (southern Mar- che), described Upper Pleistocene stratified slope de- posits made up of alternating sandy and gravel layers. The type stratigraphic sequence, about 30 m thick, crops out at Porchiano and Polesio (Fig. 5). In the se- quence, several palaeosols 20-50 cm thick, located at different heights with respect to the bottom of the depo- sits, occur. BUCCOLINI et al. (2010) constrain chronologi- cally these deposits with four 14C AMS dating of char- coal fragments present in 4 palaeosols, two located 30 cm and 1 m above the bottom of the Porchiano se- quence, and two 3 m below the top of the Polesio se- quence. At Porchiano the palaeosols are dated > 51000 yr BP and 41640 ± 1260 yr BP, respectively, at Polesio 23230 ± 170 yr BP and 22680 ± 170 yr BP. In a sandy layer of the Polesio sequence, close to the dated palaeosols, an oligotypical assemblage of land molluscs rich in specimens was recovered. It is constituted by dominating (about 43%) taxa of steppe C. D’Amico & D. Esu 72 Fig. 5 - Upper Pleistocene stratified slope deposits of the Mt. Ascensione area (southern Marche). Schematic type strati- graphic sequences: Porchiano (A) and Polesio (B); C) strati- graphic section at Polesio (from BUCCOLINI et al., 2010). Depositi stratificati di versante del Pleistocene superiore nell'a- rea del Monte dell'Ascensione (Marche meridionali). Sequenze stratigrafiche tipo: Porchiano (A) e Polesio (B); C) sezione stra- tigrafica presso Polesio (da BUCCOLINI et al., 2010). environment (4S) (Fig. 4) represented by Pupilla triplica- ta (STUDER), Truncatellina callicratis (SCACCHI), J. (J.) quadridens and J. (J.) malatestae (Tab. 2), followed (about 29%) by open land species (group 5O). C. spa- dae, fairly abundant (about 24%), prefers open wood- land, but locally can live in semi-open to open habitats. The ecological framework of the molluscan as- semblage leads us to infer open palaeoenvironment and cold climatic conditions. 3.4 Campania 3.4.1 Calore River valley (Benevento) MALATESTA (1959) reported the presence of a land molluscan assemblage in a loess layer interbed- ded to tufitic and detrital fluvial deposits of a sedimenta- ry succession located near the mineral source named ‘A Uolla on the left side of the Calore River (Telese, Bene- vento). At present, this deposits, related by the author to the “Würm”, are no more cropping out. The molluscan assemblage is constituted only by four species, J. (J.) malatestae (4S), V. pulchella (5O), P. muscorum (5O) and Helicella sp. (Tab. 2), indicating open dry environ- ment and cold climate (ESU, 1988; ESU & GIROTTI, 1991). 3.4.2 Tufino (Naples) DI VITO et al. (1998) reported J. (J.) malatestae in the Clanio-Acqualonga valley (Naples). Non-marine de- posits mainly constituted by carbonate gravels crop out in a quarry near Tufino town. A sandy palaeosol, 80 cm thick, interbedded with gravels yielded a rich, but oligo- typical land molluscan assemblage (Tab. 2). The spe- cies of dry grassland environment (4S) dominate (Fig. 4). Cernuella neglecta (DRAPARNAUD) represented by more than 500 specimens, P. muscorum of dry ex- posed places and V. pulchella of more or less damp open habitats (5O) are well represented. The ecological requirements of the molluscan fauna indicate steppe environment and cold climatic conditions. The deposit containing the molluscs was referred to the last glacial period (MIS 2). 4. CONCLUSION So far J. (J.) malatestae was found only in the seven Quaternary non-marine deposits of central and southern Italy listed above, ranging from the early Mid- dle Pleistocene to Late Pleistocene. In all the deposits J. (J.) malatestae is accompanied by the dominance of steppe species (4S) and open land species (5O) (Tab. 2 and Fig. 4), except at San Cosimato where it was reworked. The strictly forest species (1W) and the hy- grophilous ones (8H), represented only at Case Picco- netto by Sphyradium cf. S. doliolum and V. angustior respectively, are very scarce (Tab 2 and Fig. 4). At Fontignano, Case Picconetto and Gola della Rossa, small mammals were recovered with the land mol- luscs. The ecological and climatic requirements of the mammal species confirm the palaeoenvironmental in- terpretation of open-dry habitats and cold climate in- ferred by the molluscs. The palaeoecological interpretation of the mollus- can assemblages containing J. (J.) malatestae is in agreement with the chronological attribution of the re- lated deposits, correlated with the Middle and Late Pleistocene marine isotope stages MIS 22, 14, 6 (or 4) and 2, which, except MIS 14, correspond to major or minor glacial events occurred at global scale from the estabilishing of the 100 ka cyclicity, at about 900 ka BP (EHLERS & GIBBARD, 2007) (Fig. 6). ACKNOWLEDGEMENTS Many thanks are due to Dr. G. Cilla for the help in the field-research at Polesio (Marche), and G. Manga- nelli and G. Zanchetta for the useful advices. Jaminia (Jaminia) malatestae Esu, 1988 (Mollusca, Gastropoda, Enidae) … 73 Fig. 6 - Relations among Middle-Late Pleistocene glacial events, Marine Isotope Stages and the occurrences of Jaminia (J.) malatestae in Middle-Upper Pleistocene non-marine deposits of central and southern Italy. 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