Rivista Italiana di Paleontologia e Stratigrafia volume luj numero 3 pagrne 369-388 Dtcembre 7997

BIOCHRONOLOGY OF SELECTED MAMMALS, MOLLUSCS AND OSTRACODS FROM THE MIDDLE PLIOCENE TO THE LATE PLEISTOCENE IN ITALY. THE STATE OF THE ART

E. GLTOZZII, L. ABBAZZI2,P. ARGENTI3, A. AZZAROLI2,L. CALOI4, L. CAPASSO BARBATO4, G. DI STEFANO4, D. ESU4, G. FICCARELLI2, O. GIROTTI4, T. KOTSAKISS, F. MASINI6, P. MAZZA2, c. MEZZABOTTA2, M. R. PALOMBO4, C. PETRONIO4, L. ROOK2, B. SALA/, R. SARDELLA4, E. ZANALDAO & D. TORRE'

Key-uords: Continental biochronology, Plio-Pleistocene, large the mammal ages used for the biochronological scale of and micro mammals, molluscs, ostracods. the large mammals are aiso used (mainly for historical reasons) biochronology molluscs and Riassunto, Gli Autori hanno elaborato quattro tavole di distri for the of the buzione cronologica delle faune a mammiferi (macro e micrQ, a mol- ostracods. luschi e ad ostracodi di acqua dolce e salmastra, prendendo in consi- The species distribution charts refer to the Faunal derazione i più significativi giacimenti del Plio-Pleistocene della peni- Units (F.U.) succession (Azzaroli, 1977; Azzaroli et al., sola italiana. In questo lavoro viene istituita una nuova Età a Mammi- feri (Aureliano) correlabile con la pane superiore del Pleistocene Me- 7982, 1988). This succession constitutes the biochrono- dio e con il Pieistocene Superiore: di questa età sono state definite logical scale for the continental succession in the Italian due Unità Faunistiche (Torre in Pietra e Vitinia) per l'Aureliano infe- peninsula, and is calibrated with the geochronological riore e medio, mentre nessun giacimento è stato considerato rappre- scale by means of radiometric and palaeomagnetic da- sentativo per I'Aureliano superiore. k unità biocronologiche sono state calibrate con la magnetostratigrafia, con le scale isotopiche ting. Successive faunal units are not separated by a dell' ossi geno e attraverso éataziom radiometriche. boundary because of the impossibility of defining them with precision. The unreiiability of the boundaries deri- Abstract. The Authors have elaborated four range chans of ves from two causes of a different order: mammalian (large and small), molluscs and fresh-water and brackish continental successions are largely disconti- ostracodes faunas, for the selected Plio-Pleistocene fossiliferous locali- ") ties of the Italy. A new Mamnal Age (Aurelian) correlatable to late nuous and mammal remains afe usualiy concentrated in Middle and Late Pleistocene has been defined. Inside this age two horizons with a punctual distribution in stratigraphic Faunal Units (Torre in Pietra and Vitinia) have been defined as char- successions. Therefore only exceptionally can the transi- acteristic for early and middle Aurelian, while no gisements have been chosen for the late Aurelian. Biochronological units are calibra- tion to the next association be recognised in strati- ted on magnetostratigraphic and isotopic scales and by radiometric graphic succession. datings. b) the definition of F.U., which is based on all the species from local faunas selected as typical associations. This latter point merits a brief discussion. Faunal lntroduction. Units in biochronology correspond to what in biostrati- Present day knowledge about the terrestrial mam- graphy are coenozones and, in the same way, their li- mal faunas, non marine molluscs and freshwater brac- mits are not definable with precision. The distinction kish ostracods of the Italian Plio-Pleistocene enables us between successive coenozones is not based on single to attempt to construct an integrated biochronology. faxa bioevents, but on diffe- ^ppearance,/disappearance This has been achieved by means of constructing four rent associations of characteristic taxa. In fact bioevents distribution charts of selected taxa from the most signi- characterising successive F.lJ. occur during the transition ficant Italian localities. It is worthwhile underlining that between the two F.U., and their calibration is conse-

1l C..r' St".li^ ^.r il Quaternario e I'Evoluzione ambientale CNR c/o Dipanimento di Scienze della Terra, Università degli Studi "La Sapienza" di Roma p.le A. Moro 5, 00185 Roma, Italy. 2) Dipanimento di Scienze della Terra, Università degli Studi di Firenze. 3) Dipanimen- to di Scienze della Terra, Università degli Studi di Perugia. 4) Dipanimento di Scienze della Terra, Università degli Studi "La Sapienza" di Roma. 5) Dipanimento di Scienze Geologiche, Università degli Studi di Roma Tre. 6) Dipanimento di Geologia e Geodesia, Università degli Studi di Palermo. 7) Dipanimento di Geologia e Paieontologia, Università degli Studi di Ferrara. 8) Dipanimento di Scienze della Terra, Università degli Studi di Miiano. 374 E. Gliozzi et al.

quently affected by some inaccuracy (for a wider discus- few cases is it possible to reconstruct the relative chro- sion on this subject we refer the reader to De Giuli et. nological position of each local fauna within a faunal aL, 1986; Lindsay, 1990). Clearly this facror generates a unit. For this reason local faunas in a faunal unit are certain "background noise" in the contents of the distri- conventionally reported in alphabetical order and bution charts. In addition an even more important bias first/last occurrences of taxa are conventionally placed is caused by the fact that different Faunal IJnits are nor at the base of each faunal unit, represented by a dotted homogeneous from the point of view of the diversity of line. The synchrony of events observable on the distri occurring species. bution chart is attributable to this graphic convenrions. An example of this is given by the faunal renewal For the reasons discussed above, the temporal gap that occurred at the beginning of the Middle Pleistoce- between appearances is sometimes emphasised by the ne, between the Colle Curti and the Slivia Faunal Units. scarcity of local faunas, especially in the older part. In fact the few species occurring at Colle Curti - which Furthermore, the scale of the faunal renewal (measured in any case are indicative of a renewal - does not make it by the number of new appearances) is biased by the possible to establish whether this faunai change was al- gîeafer or lesser richness of the local faunas which make ready under way previous to this faunal unit. up each faunal unit. Despite these limitations, the picture which re- sults from this work gives reasonably acceptable infor- mation about the timing of the faunal renewals during Large mammals. the Plio-Pleistocene. Villafranchian Mammal Age. The distribution charts. The oldest fauna we are dealing with is the Triver- The unreliability of biochronologies based on sa F.IJ., which dates back to the Middle Pliocene. Early mammal assemblages mentioned above forced us to use Pliocene mammals in Italy are exrremely poorly repre- some graphic conventions and symbols in the distribu- sented. A few fossils were collected in the Val di Pugna tion charts which need some explanation. (Siena), assigned a Stepbanorhinus megarhinus, Sus mi- On the left in Fig. 1 and 2 are the composire cur- nor, Alephis liryx and Felsinotherium gerztaisi (Azzaroli et ve of oxygen isotopes modified from Shackleton (1995), al., 1988). A further find of this age is a fragmentary the chronologically calibrated magnetostratigraphic scale metapodial of Alephis liryx {rom the fluviolacustrine ba- (Baksi, 1993) and the geochronological scale. The sin of Montecarlo near Lucca (Dallan Nardi, 1988). Ear- boundary between the middle and the late Pliocene is ly Pliocene mammais are not sufficiently well repre- placed at isotopic stage 103, just before the beginning of sented in Italy to be included in the distribution chart. the marked cooling event which took place at about 2.5 The fauna of the Triversa F.U. (the first faunal Ma (R.io et al., 1994). The boundary between the eady unit of the early Villafranchian) show a marked renewal and the Middle Pleistocene is placed close to isotopic with respect to previous associations which are well stage 25, following the proposal made by Cita 6c Casrra- characterised in localities of Spain and southern France dori (1994). The boundary between the Middle and the (see Azzaroli et al., 1988, with references) and which are Late Pleistocene is placed at isotopic stage 5e (Aguirre & composed of taxa which are typical of humid and dense Pasini, 1985). forested environments. Faunal units calibrated by radiometric and/or pa- The Triversa F.U. is characterised by the assembla- laeomagnetic analyses are reported in the column "Cali- ge of fossils from various sites in the Villafranca d'Asti brated local faunas". The graphic convention indicating area (S. Paolo, Solbrito, Dusino, Fornace R.D.B., Casa the transition between Mammai Ages is a vertical bar Crotino, Casa Cassinotta, Arboschio, Arondelli) and ta- which covers the space between the last and the first kes its name from a small river in the area. The verte- faunal unit of two successive mammal ages. The same brate bearing fluvio-lacustrine succession lies on Early graphic convention is used rc indicate the base of selec- Pliocene (MPL 4) marine clays. The age of this F.U. ted events, which is placed in an intermediate position should be around 3.2};,4a, as suggested by palaeomagne- between two successive faunai units. tic analyses at Fornace R.D.B. (Lindsay et al., 1980), The distribution of the taxa is represenred by a although new analyses by Lindsay et al. (1995, and oral continuous vertical line when the presence of taxa is do- communication) suggest that the Fornace R.D.B. section cumented. \When the presence of a taxon is inferred or should be attributed to the upper parr of the Gilbert is documented by scarce or fragmentary material, it is magnetochronoznîe. The first date has however been represented by a domed line. The precise succession of confirmed by the palaeomagnetic survey recently carri- the appearance,/disappearance of each taxon charac- ed out at Santa Barbara near Casrelnuovo dei Sabbioni terising successive faunal units is not known. Only in a in the upper Valdarno (the first fluviolacusrrine cycle E. Gliozzi et al,

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:6:F ; >1 Biochronology of selected large mammals of Italian ó>ÈÈ Peninsula from Middle Pliocene to Late Pleistocene. f

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' The iaxonom cal posli on ot lhese iofms s disclssed by some Aulhors Biochronology of Plio-Pleistocene of mammak, rnolluscs and ostracods in ltaly -)/ |

of the upper Valdarno), where a faunal assemblage of association also includes Stephanorhinus cÍ. S. etruscus the Triversa lJnit was found (Torre et al,, 1996; Albia- while Nlrctereutes rrrcgarnastoides, Stephanorbinus cf. S. nelli et aI., 1997). In addition to the above mentioned jeanaireti and the smali sized suid are no longer present. localities, local faunas referable to this F.U. are present The beginning of the late Villafranchian (Olivola in different areas of Italy: S. Giusto (Torre, 1987), Ponte F.U.) is marked by the appearance of Paclrycrocwta breoi- a Elsa (Azzarolí, 1992), Case Nuove (Dominici et al. rostris, Panthera gombaszoegensis (: P. toscana), Procamp- 1995; Benvenuti et al., 1995) in the lower Valdarno; Spo- toceras brioatense and new species of Eucladoceros and leto in Umbria (Tuccimei, 1898; Sardella et aI., 1995) Pseudodama (E. dicranios, P nesti). Furrhermore mo- and Arciile near Grosseto in southern Tuscanv (Hùrze- dern dogs (Canis etruscu) become widespread. The ma- ler 6c Engesser, 1976; Masini &.Torre, 1987). jority of the Pliocene species which had survived the In this unit there is the first recorded appearance first crisis linked with the first arctic glaciation (isotopic of the genus Pseudodama, Leptobos stenometopon and Ste- stage 100 and afterwards) are no longer present or are phanorhinus jeanvireti, and several large carnivores such extremely rare. The most important local fauna is that as Acinonyx, Homotherium and Ursus minimus. There of Olivola (val di M"g.") which is probably located in are also forms of Viverridae (Viaerra and Megaviaena) the Olduvai chronozone. In fact, research which is still and an extinct form of panda (Parailuru). Tapirus druer- in progress at Poggio Rosso (upper Valdarno; Torre, un- nensis and Sus minor are still well represented whíle Me- published) suggests that the adjacent site at Marassino, sopithecus monspessulanzs occurs for the last time. which in the past had been attributed to this F.U. and The beginning of the early Villafranchian (Triver- palaeomagnetically calibrated with the B event in the sa F.U.) thus represents a marked faunal turnover. The upper part of the Olduvai chronozone (Torre et al., occurrence new taxa enriches fauna a of the which. as 1993, t996), has a faunal assemblage with characteristics whole, has more advanced character compared a with which are transitional between the Olivola F.U. and the the Ruscinian ones. following Tasso F.U. A marked faunal renewal marks the transition be- The Tasso F.U. is characterised by the arríval of tween this F.U. and the following Montopoli F.U.; taxa Htppopoumus antiquus, Praewibos, a bisonlike kptobos typical of wooded habitats disappear bor- lZygolophodon (L. vallisaml, Equus steblini, Canis arnensis and Canis soni, Thpirus araernensis, Sus minor (: Sus araernensis); (Xenocyon) falconeri, together with the disappearance of Ursus minimusf and in Italy one sees the spread of a Gallogoral menegbinii and Gazellospira torticornis (De primitive elephant (Mammuthus gromwi), and the horse Giuli & Masini, 1987; Azzaroli et al., 1988; Ambrosetti (Equus lioenzwensi). Both f^x^ are indicative of more et aL, I995a). These events determine a marked change open landscapes. At that time Stephanorhinus etruscus in the faunal association, and it acquires a typically also probably appears (R.ustioni et aI., 1,995). The local character. Montopoli fauna, in the Lower Valdarno, was collected Quaternary After the turnover responsible for the formation at the top of a marine succession wìth Globorotalia cras- of the Tasso F,IJ., most of the typically Villafranchian saformis and is calibrated with the Gauss/Matuyama taxa disappear or become extremely rare, and species transition in the magnetostratigraphic scale (Benvenuti which are phylogenetically closer the Pleisto- et al., t995; Lindsay et al., 1980; Torre, t987). to Middle cene species appear. This The small local fauna at Collepardo, located in new situation characterises the Farneta F.U. (AzzaroIí, 1977). kptobos and Eucladoceros the Anagni basin in Latium, is composed of Megante- are present reon cultridens, Nyctereutes meganxdstoides, Stephanorhi- still and, for the first time, a deer of the Megaceroides clade (M. obscwru) nus cf. S. jeanoireti, Szzs sp. (of a small size), Pseudodama and the vole Microtus (Allopbaiorny) appear (Masini Santini, lyra and "Hernitragus cf.. H. stehlinl" (Cassoli & Segre & 1.991; Azzaroli Naldini, 1994). \fith some reservations due to the scar- &Mazza, 1993; Abbazzi, 1995). The richest locality for city of material, this small fauna could be referred to the this F.U. is Pietrafitta in Umbria (Ambrosetti er al., end of the early Villafranchian. 1987; Ambrosetti et aI., 1992 a, b) where there are char- Middle Villafranchian faunas are poorly repre- acteristic small sized rhinoceroses (Mazza et al., 1993). sented in ltaly, and they are represented by small assem- Pirro Nord-Cava dell'Erba, karst fissure fills in blages which have not yet been fully studied. One of the Gargano, (De Giuli et al., 1987; Abbazzi et al., 1,996 these comes from the locality of Costa S. Giacomo, again b), and Capena near Rome (Petronio, 1,979) are two lo- in the Anagni basin. The Costa S. Giacomo local fauna calities where the first occurrence has been recorded of is characterised by the first occurrence of the genus Ca- a primitive bison, Bison (Eobison), and of a Megaceroides nis s.s., Sws cÍ. S. strozzii, Leptobos furtiztus, Hystrix cf.. H. species, probably attributable to rhe M. solilbacus phyle- refossa and Gazellospira torticomis, and the last occurren- tic lineage (Abbazzt & Masini, 1997). The presence of ce of Anancus draernensls (Cassoli & Segre Naldini, ovibovine remains at Pirro Nord is also very significant. 1994; Masiní, 1.989; Rook & Torre, t996). The faunal These novelties characterise the Pirro Nord F.U. 372 E. Gliozzi et al.

Galerian Age. shows a more marked faunal renewal which can already be seen in the Slivia F.U. and which is much more noti- Taxa which are considered to be classically Gale- ceable in the Isernia F.U.. Lion, paleoloxodont elephants rian (such as Equus altidens, Equus gr. bressanus-suessen- and the subgenus Mammuthus, a typical Stepba- bornensis and the genus Megaceroi.de) already occur in form of norhinws bundsheimensls, new megaiocerines new the F.U. of the Late Villafranchian and the faunal re- and bovids such as Bos and bison belonging subgenus .Bl- newal that produced typically Galerian assemblages was to son (Bison) and, among the equids, Equus caballus, all a gradual phenomenon. The Colle Curti local fauna is appear. Some Villafranchian conventionally used to mark the beginning of this large carnivores and equids as well as an advanced form o{. Pseudodamd are pre- Mammal Age. This faunal assemblage (Ficcarelli & Sil- srill sent. The oldest sites middle vestrini, 1991), which unfortunately is not very rich, with Galerian faunas be- Ionging to the Slivia F.U. are Slivia (Ambrosetti et al., was found in sediments with a normal polarity and 1979; Bon et a1., 1992) and Monre Tenda (Pasa, 1947). which are correlated with the base of the Jaramillo sub- Due to the persistence of some Villafranchian elements, chronozone (Torre et al., 1996; Albianelli et al., 1997). the Selva Vecchia local fauna (Bon et I99I) seems ro The fauna is characterised by the presence of the first ^1., be slightly more archaic than that at Slivia. assem- forms of Megaceroides perticornis. The small number of The blages from Isernia (Sala, 1983a, 7996a), Cesi (lr4asini et species recorded in the assemblage does not make it pos- al., 1991; Ficcarelli al., press) Valdemino (Sala, sible to verify how many typical Galerian taxa'were pre- et in 1992) and various sites the sent at the timet fossils of a medium-sized bovine could in surroundings of Ponte Ga- leria (Capasso Barbato Petronio, Pa- be attributed either to a small Bison or, alternatively, to & 1984; Caloi & lombo, 1994a) are attributed to the following Isernia lcptobos. The presence of Pliomls lenki together with a F.U. Microtus (Allopbaiomys), which was probably ancestral The fauna from Visogliano (Cattani 1991) to Microtus ex gr. oeconomus, conltrms the innovative et al., and Venosa-Notarchirico (Belli et a1., 1991) character of this faunal assemblage and strengthens its are probably transitional those late attribution to the Galerian. with of the Galerian. The late Galerian assemblage, to which the Fonta- Subsequently there were successive dispersal na Ranuccio F.U. is referred, characterised by the total events of Asiatic and central European taxa into the Ita- is disappearance of typical Villafranchian raxa with the ex- lian peninsula, probably in relation to phases of climatic ception of Homotherium, whrch is represented by an cooling. At the same time new taxa appear as a result of evolved form (Sardella, 1,994). The red deer subspecies local evolution. \íithin the Galerian complex it is there- Centus elaphus eostepl)anoceros is present (Di Stefano & fore possible to recognise three successive associations Petronio, 1993) and Dama clactoniana, E. altidens and E. characterising the early, middle and late Galerian respec- suessenbornensls continue repre- tively. to occur. The most sentative sites are Fontana Ranuccio (Segre, 1984) and The early Galerian assemblage is characterised by Cava Nera Molinario (Di Stefano Ec Petronio, 1993). the presence of Megaceroides uerticornis and by the per- sistence of forms with Villafranchian affinities such as Aurelian Mammal Age. Pa clry cro cuta breo iro stris, H omoteriurn ex gr, crenatidens- latidens, Mammuthus (Archidiskodon) meridionalis, a A marked renewal occurs at the transition to the small sized Stephanorhinus and an advanced form of following faunal assemblage, with the extinction of Pseudodama. The Colle Curti F.U. is attributed to the some Galerian forms such as the megacerine cervids of early Galerian; it was established on the basis of the fau- the Megaceroides oerticornls group, Megaloceros saoini and nas of the Colle Curti site and those from Monte Peglia the more archaic elaphine deer, while taxa which consti- (van der Meulen, 1.973; Piperno & Segre, 1984). At the tute the core of the modern mammal fauna appear for latter site the iarge mammal assemblage, which has yet the first time. Very rich deposits are mainly located in to be studied analyticallt suggests the persistence of a the Tyrrhenian coastal area oÍ Latium, aiong the S.S. 1 large amount of Villafranchian taxa, while the small Aurelia. For this reason we propose to use the term Au- mammals indicate a more evolved assemblage than the relian age for this faunal assemblage. Aurelian faunas one from Pirro Nord. It appears therefore that the early come from deposits which date to the late Middle Plei- Galerian assemblage in ltaly, even though it is poor in stocene and the Late Pleistocene. species and scarcely represented in terms of number of Bioevents such as the first occurrence of the mo- sites, constitutes a complex which is indicative of the dern large wolf Canis lupus and of Ursus spelaeus have transition between typical late Villafranchian and typi- been chosen to define this faunal assemblage. In the Au- cal Galerian faunas. relian faunal complex there are three different and sub- The middle Galerian assemblage, which is charac- sequent associations (early, middle and late Aurelian) terised by the presence of Centus elaphus dcoronatus, which are related to the progressive occurrences of diffe- Biocbronology of Plio-Pleistocene of mammals, molluscs and ostracods in ltafu J/ 3

rent taxa of Asian origin or of other forms which evol- 1983) and Melpignano (Boiogna et a1., 1994), Gtott" ved in the Mediterranean area. Moscerini (lower levels), Guattari and Fossellone (Cir- The early Aurelian assemblage, which is repre- ceo) (Caloi & Palombo, 1.994b). sented by the Torre in Pietra F.U., is characterised by Subsequently, not only in Europe but also within the first occurrence of the modern wolf and of the cave the Italian peninsula, there was an enlargement of the bear as well as by the first occurrence of Megaloceros gi- distribution area of Capra ibex and the spread of large gdnteus and the diffusion of the most archaic forms of cold environment stenothermic mammals such as Coelo- modern red deer Cerous elapbus riaiensis. Correlations donta antiqwitatis and Mammuthus primigeniws. Several between the type locality of Torre in Pietra (Torre di raxa of large herbivores and carnivores progressively di- Pagliaccetto) and the Middle Pleistocene Tyrrhenian sho- sappeared in correspondence with the following phases relines refer this faunal unit to isotopic stage 9 (Conato of climatic deterioration. et al., 1980). The most representative deposits of this The most recent late Aurelian assemblages come Faunal Unit, besides Torre in Pietra (ower level$ (Caioi from a large number of localities all over ltaly and have & Palombo, 1928; Cassolí, 1978), are Castel di Guido mainly been found in cave deposits. Among these the (Sala 6c Barbi, 1996), Malagrotta and Riano (Caloi Ec sites with the most complete palaeontological, palaeoe- Palombo, 1994a) and la Polledrara di Cecanibbio (Anzi- cological and palaeoclimatic data are Grotta del Broion dei et a1., 1989; Arnoldus-Fluyzendveld & Anzidei, and Riparo Tagliente, (Bon et aI., 1991) in Northern 1993), aII located in Latium. Italy, Grotta Breuil, Grotta Barbara and the upper levels The next faunal assemblages, referable to the mid- of Grotta dei Moscerini, Grotta Fossellone, Grotta dle Aurelian (isotopic stage Z), are characterised by the Guattari and Grotta S. Agostino in Central Italy (Caloi occurrence and diffusion of the most primitive subspe- & Palombo, 1994b), and Grotta del Cavallo (Bartolo- cies of the modern fallow deer Dama dama tiberina (Di mei, 1980), Grotta Paglicci (Sala, 1983 b; Abbazzt et al., Stefano & Petronio, 1997) and Equus lrydruntinus, du- in press b) and Grotta della Serratura (Bertolini et al., in ring a period of climatic amelioration which is also re- press) in southern Italy. cognisabie in the marine environment. The Vitinia F.U. is repre- has been attributed to the middle Aurelian; it Small mammals. sented by the fauna from Vitinia, Bucine, Cerveteri, Se- dia del Diavolo and Torre in Pietra (upper levels) (Caloi In contrast with the large mammals, studies of lta- & Palombo, 1994a). lian fossil small mammals started only much more re- The late Aurelian assemblage, which begins with cently. In fact, with the exception of the research carri- the Eemian and terminates with the end of the last gla- ed out by Bosco and Forsyth Major almost a century rWorld ciation, is difficult to define because it is mainly charac- ago, it was only after the end of the Second War terised by the disappearance of large and medium sized that research was renewed by Pasa and Bartolomei and mammals. In the time span corresponding to this faunal only since the seventies that a greater number of pa- assemblage there was a series of important climatic laeontologists began to work on this topic. events and the different microclimatic and environmen- The Middle and Late Pliocene and the Early Plei- tal conditions have strongly influenced the faunal com- stocene small mammal record is relatively scarce and the number of sites is small, while the Middle and Late Plei- position along both the Tyrrhenian and the Adriatic co- stocene deposits are as numerous as those large asts of the Italian peninsula (Sala 1983b; Masseti et al., for the mammals, although the number sites both 1995; Torre et al., in press). During this period the fau- of with kinds of faunal assemblages is relatively low. nal assemblages become poorer and, in correspondence The biochronological attribution of the assembla- with the beginning of the last Interglacial, are more and ges follows the scheme proposed by Fejfar Heinrich more conditioned by anthropic influence. Even though & (1989). FIowever, since the large mammal data for the the deposits related to this assemblage are both nume- Pliocene and Early Pleistocene in the Italian peninsula rous and well studied, for the reasons mentioned above are considerably more detailed than that of the smail none of them can be considered to be fully repre- mammals, where possible the fauna have been referred sentative of the late Aurelian faunal population, and as a to the biochronological scale of the large mammals. result it is not possible to choose a local reference fauna to define faunal units within this assemblage. Villafranchian Mammal Age. The oldest local faunas (isotopic stage 5) referable to the late Aurelian assemblage are characterised by the A rich fauna with insectivores, rodents and lago- first occurrence of the modern form Dama dama dama. morphs comes from Cascina Arondelli (Villafranca The most important deposits are mainly located in cen- d'Asti, Piedmont) and is typical of the Triversa F.U. tral and southern Italy, such as S. Sidero (De Giuli, The assemblage is characterised by shrews such as Episo- J74 E. Gliozzi. et al.

q ! Calibrated ;l 180 curue Selected tocal Faunal Uniìs (Shacklelon, 1 995) iaunas localities ò a

Grolta deL Broion. R paro :: lezzena, Cava A Veia. Gro ta Gh acc a a. Casielclvìla, '' 5 u Groita Seratura, S. S defo E È S Befnard na [,lagqiore e É tl Itl '.' EI

Loara S. Agoslino (Co i ì s',1'rp c Sl.9e I ll Berict t- Boscochlesanuova, San 0 458 Ma 1l E -i- Giovanni di D!ìno. FI Spessa lL èl:il Él ::-' ='l= Venosa'Nolarchirico ,lIr I I I .i 3 -.::. 1o, ? 0.736 Mqi I (KArddn!-i E .: I O .È '*"it!1j" - : lr I t_'-r- ioB o Sliv a F. U Monle lenda Slìvia i o Fofl gnano. S€lva Vecchia iî io, il rl tl .+ ill :.- n irc 99 a Co le Cuill. Monle ltl t Colle Cudr ill i-=: =:- . ;= ill i,, *î Soave Cava Sud :l t --:- sl _ : - - 6_9 -..-.= l* 1.5 tl --t :'= ;; tl

Case lnjerno (Uppef ;î eZ --..i . ..:: ,- Olvoa F U ilt "l Casle francó d Sòpra - I -=.= rrÌ:

20 . .<- Costa S Giacomo, 9 I Cosla S G It . I tl É ltl 22 .. = 23 li -t-- 24 :.]

1 25 Co lepardo tm I >16 26 "i i& = Monlopo i ol -:.:-,'' 27 ,l tl 2A [-* ll i -l- I tl 29 tl -jt'- ll :t_ I ll 30 -a.' tl a : .,- " I J2 Arcii-. Santr Barbara. Tr;versa (vario!s silesl I :> I 33 lt

gÉ qi;: E íÉ ì si * s: ltj liîi ? E;àlÉ: Ii;:3É e; iar i ;r ÉsÉ: s t' :,Èi: i i; i = a= iÉi gàa;! ÈrÈ lÉÈiiÉiÉÈiÉiiiuiÉi! iiÉ i ";i ;j* ;É,È, j

Frg.2 - Biochronology of selected small mammals of Italian Peninsula from Middle Pliocene to Late Pleistocene. Biochronology of Plio-Pleistocene of marnmals, molluscs and. ostracods in Italy 375

riculus gibberodon (:Asoriculus after Flutterer, 1994), Crocidura and of a modern type of mouse of the subge- Blarinoides mariae, Petenyia bungarica and Beremendia nus Apodernus (Sylvaernus)(De Giuli et a1.,1987; Masini fissidens, by the voles Mimomys hajnackenesis and Mi- & Santini, I99l; Abbazzi et aL, I996b). The faunas rnorlys stehlini and by other rodents and lagomorphs from Palena (LAquila, Abruzzn) (Kotsakis, unpubl. (Mwscardinus cf. M. pliocaenicils, Glirulws pusillus, Prola- data) and Soave Cava Sud (Verona, Veneto) (Pasa, 1947 gus saaagei etc.) (Berzi et aI., 1967; Michaux, 1970; Masi- Bartolomei, 1980) can also be referred to the Pirro F.U. ni 6c Torre, 1987). The fauna from Arcille (Grosseto, Tu- All these micromammals assemblages indicate the early scany) with Mimom.ys hajnackensis and from San Giusto Biharian. (Pisa, lower Valdarno, Tuscany), rype locality of Mi- mom)s stehlini, are also attributed to the Triversa F.U. Galerian Mammal Age. These three faunas are referable to the early Villanyian In the type locality of the first Galerian F.U., Col- (Masini & Torre, 1987). Many European deposits bea- le Curti (lr4arche), together with large mammals there ring the remains of Mimomls hajnackensis have been ca- 'were the remains of Microtus (Allopbaiorrryy' sp., which is librated with the lower part of the Gauss normal ma- probably a primitive form of the Microtus oecono?nus gnetochronozone. group, and Pliom.ys lenki Qorre et al., in press; Abbazzi The following faunal assemblage was found in the Ec Masini, unpublished data). The fauna from the karst karst deposit at Rivoli Veronese (Verona, Veneto). In this deposit of Monte Peglia (Orvieto, Umbria) can be amri- fauna there are several voles which occur for the first buted to the same F.U., aithough it may be slightly ol- time: Mitnomjs pliocaenicus, Mimom.ys pitymyoides, Mi- der. At this site there are two fossiliferous levels, but the rnorn)s tornensis, Villanya cf. V. exilis, cf. Eilobius sp., a faunal composition of each is very similar. The rich as- primitive form the genus Dinarom.ys, allegranzii of D. semblage of small mammals includes the lasr forms and shrew subgenus (Drepanosorex). a of the Sorex The known in Italy of the genus Berentendia, Glirulus, unga- specimens genvs Mimomlts îound of the in the deposit romjs. The subgenus M. (Allophaiomls) occurs with the allow this fauna be attributed to the early to Villanyian more evolved forms, M. (A.) nutiensis (Microtus sp. A, in and the middle Villafranchian (Sala et al., 1994; Sala, van der Meulen, 1973) and lr[. (A.) burgondiae (Microtus 1.996 b; Fanfani 6c Masini, unpublished data). Some re- sp. B in van der Meulen, 1973). Mirnomlts saoini, Mi- mains of Apodemus cf. A. dominans found in Cava Top- mornys blanci and Pliomys episcopalis are also presenr (Todi, petti Umbria) (Abbazzi et aL, 1996a) are roughly (van der Meulen, t973). The Colle Curti and Monte Pe- coeval with those found at Rivoli Veronese. glia faunas indicate rhe final phase of the early Biharian. The isolated remains of Mimornys pliocaenicus An assemblage composed of only two rodents, found at Castelfranco di Sopra (Upper Valdarno, Tusca- Prolagurus pannonicus and (?) Predicrostonyx sp., associa- ny) and Bocchignano (Sabina, Latium) (lr4asini & Torre, ted with a molluscan fauna indicative of cold climatic 1987; Kotsakis, 1988) are referred to fhe late Villanyian conditions, was found at Fontignano (Ponte Galeria, and to the late Viliafranchian (Olivola F.U.) (on the ba- Rome, Latium). This fauna comes from the clays be- sis of the geological context of the deposits and the large neath the conglomerates where the Ponte Galeria large mammals assemblages). mammals were found and corresponds with the final Stiil in the Upper Valdarno (locality Case Inferno, phase of isotope stage 22 (Kotsakis et al., 1992). Tuscany), in deposits containing large mammals of the In the type locality of the second F.U. of the Ga- Tasso F.U. (late Villafranchian), there was an isolated lerian, the Slivia F.U. (Aurisina, Trieste, Yenezía Giulia), find of Mimomys saoini (:Cromerom.ys after Neraudeau some species of micromammals were found as well as et al., 1995), a species that marks the beginning of the numerous large mammal remains. In the breccia at this Biharian (Torre, 1985; Masini &Torre,1,987). site there were Mimomys saaini and Dinarom),s sp., and At Pietrafitta (Perugia, Umbria) the remains of vo- the first appearance of Allocricetus bursae and Microtus les were found associated with large mammals of the (Neodon) gregaloides (Ambrosetti et al., 1979). This as- Farneta F.U. (late Villafranchian): Mimomys pusillus and semblage can be referred to the late Biharian. The fauna the first forms known in Italy of the subgenus Microtus from the karst fill at Monte Tenda (Soave, Verona, Vene- (Allophaiomyt), M. (A.) ex gr. M. (A.) rffii and M. (A.) to) belongs to the same Mammal Age (Late Biharian). It cbalinei. The assemblage includes typical species of the is characterised by the occurrence of Mimomys saaini early Biharian (Gentili et a1.,7996). and Pliomys episcopalis and by the first appearance of At Pirro Nord (Foggia, Apulia), the type localiry the subgenera Microtus (Microtus) and Miuotus (lerricola) of the Pirro F.U. which is the last F.U. of the late Viila- (Pasa, 1947; Bartolomei, 1969). franchian, an abundant small mammal fauna was found At the type locality of the following Galerian associated with large mammals which included M. (A.) F.LJ., Isernia (lr4olise), both large and small mammal re- cf. rffii and the first occurrence in Italy of the shrew mains were found. In the fluviolacustrine deposits 376 E. Gliozzi et al.

Pliomls episcopalis, Pliomys lenki, Miuotus (Miuotus) aff. The fauna from the karst fill deposits at San Side- M. (À4.) araalis and Microtus ([erricola) gr. mwltiplex-swb- ro 3 (lr4aglie, Lecce, Apulia), where Microtus (Terricola) terraneus occur together with a Clethrionomys gr. glareo- savii ís the dominant species associated with Microtus ar- lus, Microtus (Iberomys) brecciensis and a small vole, nor oalis, Lepus europaeus and Oryctolagus cuniculus (De Giu- very hypsodont, with mimomian enamel and at least li, 1983; Nocchi & Sala, in press b), can be referred to one radicated M1, which has been classified as cf. Arui- the last Interglacial. cola cantiana (Coltorti et al., !982; Sala, 1983a; Sala, There is a large number of small mammal faunas I996a). The fauna of Valdemino cave (Borgio Yerezzi, which date to the last Glacial and the sires are distribu- Savona, Liguria), where M. (.) brecciensis is very com- ted all over the Italian peninsula. During this period mon and where a new species of lagomorph was found, Italy represented a refuge area for many species of mam- Oryctolagws burgi (Sala, 1992; Nocchi & Sala, in press a), mals, and for this reason in some localities temperate belongs to the same F.U. but to a different climatic pha- forms survived during the phases of maximum climatic se. Both the Isernia and Valdemino faunas may be refer- deterioration. The geographical and morphological [ea- red to the beginning of the Toringian. tures of the Italian peninsula resulted in a subdivision The fauna of Riparo A di Visogliano (Trieste, Ve- into areas characterised by very different ciimates and nezia Giulia), with its typical glacial features, is proba- landscapes. Therefore the small mammal assemblages bly more recent than the faunas described above, with vary not only as a result of the climatic changes in a Pliom.ys episcopalis, Dinarom.ys bogdanwi, Anticola can- given area but also vary significantly as a resulr of the tiana and Allocricetus bursae and the first appearance of geographical location and altitude. Notable differences Microtus (Stenocranius) gregalis, Sicisa sp. and Ochotona existed between the Adriatic side of the peninsula, which cf. O. pwsilla. The fauna can be referred to the early was characterised by a colder and more arid Mediterra- Toringian and it is probably located between isotopic nean climate, and the Tyrrhenian side which was more stages L4 and 10 (Bartolomei & Tozzí, 1,978; Cattani et temperate, and between the faunal associations from coa- aI., 1.991; Tozzi, 1995). The fauna from the fluviolacu- stal and foothill areas and those for the mountains. (Venosa, strine deposit at Notarchirico Potenza, Basiiica- The Last Glacial small mammal faunas are gene- ta) can be attributed to the same period; bgether v/ith rally characterised by a greàrer abundance of Microtus Microtus afÍ. M. aroalis which is dominant, rhere are also (Microtus) arpalis and Miuotus (Microtus) agrestis compa- the remains of Pliornys episcopalis, Ar"uicola cantidna and red with the other voles and the mice of the genus r4po- Cbionom.ys nioalis (Sala, 1991). demus (e.g. Ingarano, Foggia; Capasso Barbato et a1., Finally, faunas referred two can be by indirect cor- 1.992;Petronio et al., 1996). Dormice are represented by relation the Fontana to Ranuccio F.U., the last F.U. of Myoxus glis, Muscardinus aoellanarius, Eliomys quercinus the Galerian: that found ossiferous in the breccia at Bo- and DrTomys nitedula (which were also found at Grotta scochiesanuova (Verona, Veneto) vzith as voles such Breuil - Monte Circeo, Latium; Kotsakis, 1991; Alhaique Pliomls episcopalis, Pliomys lenki, Dinarornys bogdanwi et al., !995 - a locality situated between the two areas of and shrews (Sorex runtonensis, Crocidura (Bartolo- zorzi) Italy where it is currently found) and are usually very mei & Pasa, 1970), and that found in the karst deposit rare. Among the voles, Microtus (Tenicola) always occurs at San Giovanni di Duino (Trieste, Yenezía Giulia), whe- in the Southern Italian regions which represented a refu- re Pliomys episcopalis, Dinaromys bogdanwi, Microtus ge , and it became more abundant during the more (terrícola) saztii, Allouicetus bursae and other species oc- ^re temperate phases (e.g. at Grotta di Castelcivita, Salerno, cur (Bartolomei, 1976).In both deposits there is a large Campania, Cioni et a1.,1.979; Masini tt Abbazzi,1997). soricine, probably Episoriculus sp.. The faunas from the- Among the insectivorous, besides the nearly constanr se deposits are referred to the final phase of the late To- presence of Talpa and Erinaceu.s, the shrews of the genus ringian. Sorex are relatively widespread, while Crocidura seems to be absent from Northern Italy. In the North Italian Aurelian Mammal Age. sites and those of the Adriatic coast there are sporadic The fauna from the outer levels at Grotta di San occurrences of species with "steppic" affinities during Bernardino Maggiore (Colli Berici, Yicenza, Veneto) is colder and more arid periods which came from North- characterised by the presence of Marmoa mdrrnota, Cri- ern or Eastern Europe. The main species is Microtus oe- cetus cricetus, Dinaromys bogdanoui and Microtus gr. ared- conotnu.s, numerous populations of which are sometimes lis-agrestis (Bartolomei, 1960). These levels have been re- found in sites in the Po valley (Zanalda, 1994). The oc- ferred to the penultimate glaciation by the radiometric currence of others "steppic" species (Sicista cf. S. betwli method U/Th (Falguères et al., t996) and probably cor- na and Ochotona pusilla) is very rare and so far has been respond with the Vitinia F.U. of the Aurelian Mammal discovered only in the regions of Veneto and Marche Ag.. (Bartolomei, 1980; Sala, 1990). Biocbronology of Plio-Pleistocene of mamrnals, molluscs and ostracods in ltaly )/ /

Among the most important stratigraphic sequen- of deposits referred to the early Villafranchian Mammal ces from the point of view of the abundance of the fos- Age; column 2 shows the late Pliocene species which sil record and the stratigraphical continuity is that at can be correlated with the middle Villafranchian Mam- Grotta del Broion (Colli Berici, Yícenza; Zanalda, 1994). mal Age; species from sites referred to the faunal units The first part of the Last Glaciation is well documented of Olivola, Tasso, Farneta (ate Villafranchian Mammal at Grotta di Veja, Riparo Mezzena, Riparo della Ghiac- Age) are listed in columns 3, 4 and 5 respectively; spe- ciaia (Monti Lessini, Verona) and the lower levels of Ri- cies related to the Galerian Mammal Age are shown in paro Tagliente (Grezzana, Verona) (Bon et al., 1991). column 6 and, finally, column 7 groups the species of The recent part of the inter-Pleniglacial is clearly the Aurelian Mammal Age. documented in the deposits at Grotta della Cala (lr4arina di Camerota, Salerno; Bartolomei eT. al., 1,975) and Grot- Villafranchian Mammal Age. (Monti Salerno; et a1., ta di Castelcivita Alburni, Cioni Early Villafranchian Mammal Age. The non-marine 1,979; Masini Abbazzí, 1997) ín Southern Italy, while & molluscs of this mammal are referred to the Middle the Late Pleniglacial and the Late Glacial are well docu- ^ge Pliocene and to the Triversa F.U. These assemblages are mented many sites: these include the upper levels of in characterised by an abundant stock of species, 35 in the deposit at Riparo Tagliente (Grezzana, Verona), number, belonging mainly to land pulmonates, two Grotta Paglicci (Rignano Garganico, Foggia; Bartolomei, thirds of which do not seem to reach the Middle Villa- 1975), Grotta della Serratura (Marina di Camerota, Saler- franchian. Whilst some species are limited to the no; Bertolini et al., in press), the upper levels at Grotta Piedmont basin (Viaiparus pollonerai, Emntericiz plioce- della Cala di Camerota, Salerno) and Grotta di $4arina nica, Discus pantanellii, Janulus angustiumbilicatus, Reti- Praia a Mare (Cosenza, Calabría; Capasso Barbato & nella (Lyrodiscus) cf. R.(L.) jourdani, Tiiptychia mastodon- Glíozzi, 1995). tophila, etc.), others are also widespread in central Italy (Laminifera (Laminiplicaa) villafranchiana, Carychium (Saraphia) pseudotetrodon, Negulus oillafranchianus, Ga- Molluscs. strocopta (Albinula) acuminata fossanensis, Gastrocopta (Wrtigopsis) debrni, kiostyla gottschicki, Eostrobilops aloi- Key for the distribution chart of the non-marine molluscs. sii, Palaeoglandina lunensls, etc.) (Esu & Girotti, 1991; Ciangherotti et aI., 1996). Sacco (1887, 1892) cites nume- The Plio-Pleistocene assemblages of land and rous other taxa from Pliocene sediments at various sites freshwater molluscs of the Italian peninsula are chrono- in the Piedmont basin (Tassarolo, Ceresole d'Alba, etc.). logically related to the mammal ages shown in the mam- IJnfortunateiy the exact stratigraphic position of these mal distribution chart in this paper. Only extinct spe- sites is unknown and therefore these species are not li- cies and those living species which have some biochro- sted'in the chart. Nevertheless the age may be consi- nological significance in connection with their appea- stent with the Triversa F.U. (see also Esu et aI., 1993).It rance and extinction in Italy have been taken into consi- is impossible to compare these assemblages with older deration. ones (Ruscinian, Early Pliocene) because of the almost In the distribution chart the species of molluscs complete absence of data on molluscs from this period are plotted against their site(s) of provenance which, in resulting from the scarcity of continental sediments in turn, are ordered according to the mammal ages, since italy which date to the Lower Pliocene (Esu, 1982). most of the sites are the same for both the molluscs and From a paleobiogeographic point of view some species the vertebrates. In the case of sites which are important of the Piedmont basin are also present in the basins of from a malacological point of view but lack vertebrates, La Bresse (Rhone, France) and in the Rhine (Germany) the chronostratigraphic position was obtained by means (Esu et a1.,1993). of correlations with marine successions, palaeomagnetic data, isotopic stages or with other continental molluscan Middle Villafranchian Mammal Age. Two Umbrian faunas. sites, Cava Toppetti ("upper beds') and Dunarobba, The first part of the chart, which contains the list have been referred to the Late Pliocene (middle Villa- of coevai sites arranged in order, gives the impression for franchian Mammal Age) for various reasons. At Cava some species of a more extensive stratigraphic range Toppetti the lower leve1s, which are part of the Fosso than is actually the case, while for others their appea- Bianco Formation (Basilici, in Ambrosetti et al., 1995 b) rance is scaled over time. The second part of the chart, and contain Prososthenia sp. n. and Zilcheuchilus sp. n,, which is related to the mammal ages and not to the si- have been referred to the Middle and Late Pliocene on tes, gives a more realistic picture of the stratigraphic di- the basis of palaeomagnetic data (from Gauss to Reu- stribution of the species: column L contains the species nion, see Albianelli & Napoleone, in press); the two 378 E. Gliozzi et al.

MOLLUSC BIOCHRONOLOGY

r - - I i i m m m r - r g t a m I It a o I I r I z z m- o z o I T - r I t g z zt z T I I : l o q z T o o F o 6 -m m f I ]' a I c - t = zt tz c 5 z z

Fig. 3 - Biochronology of the extinct non-marine molluscs of Italian Peninsula from Middle Pliocene to Late Pleistocene. The asterisc indica- tes selected living specres. Biochronology of Plio-Pleistocene of mamrnals, molluscs and ostracods in ltaly 379

Prosobranchia mentioned above come from the levels characterising the late Villafranchian basins of the with Gauss normal magnetic polarity. The "upper Tyrrhenian side, such as T (N) groyanus and Viaiparus beds"(: Ponte Naja Formation, Basilici, 1995) are refer- ampullaceus (Crispino & Esu, 1995; Ciangherotti et al., red to the Late Pliocene (middle Villafranchian Mammal in press, a). Age) by Abbazzi et al.(1996a) on the basis of their The Leffe basin in Northern Italy (Bergamo) has a mammal content. The malacological assemblage in this completely different fauna, with still living species and formation is quite different from that of the "lower only one exrincr species, Val"tata chalinei, which has beds" and the species are mostly common to those of been found outside of Italy in the Late Pliocene and the Dunarobba site where the sediments belong to the Early Pleistocene of the La Bresse basin (France) (Esu et top of the Fosso Bianco Formation (Ambrosetti et al., al., L993). 1995b). The Prosobranchia of the two assemblages form a limited stock of freshwater species which reach the Galerian Mammal Age. early Pleistocene the Tiberino basin, whereas the in The malacofaunal complex referable to the Gale- pulmonates land disappear at the end of the Pliocene rian Mammal Age is characterised by mainly modern apart from dehmi G(V) which has recently been recor- prosobranchs and pulmonates; the Villafranchian species "Villafranchian" ded in Upper sediments near Fighille are completeiy extinct and only a few of the Galerian (Anghiari, Arezzo) (unpubl. data). species. Of these, as far is we know; L .aff. L. jahni and T aÍf. T danubialis are only present in the upper beds of Late Villafranchian Mammal Age. Non-marine as- the Stirone sequence (early Galerian Mammal Age); semblages from several sites, most of which are in cen- Tbeodoxus isseli and Jaminia malatestal extinguish during tral Ital;r, are referred to the late Villafranchian Mammal the Aurelian Mammal Age (Esu & Girotti, 1991). The Age, divided between the various F.U. of the mammals. extinction of several Villafranchian endemic taxa of cen- The rich maiacofaunal complex is constituted by nume- tral Italy is due to the climatic and geological history of rous species of prosobranchs and pulmonates (Esu & Gi- these basins at the end of the Early Pleistocene. At the rotti, 1975; Conti & Esu, 1981; Esu 6r Girotti, 1991; beginning of the Galerian Mammal Age oligotypic as- Ambrosetti et al., 1995a; Ciangherotti et al., in press b). semblages similar to those recorded in the loess deposits Among the pulmonates some Pliocene species survive: of central Europe (Lozek, 1964) appear in central Italy N. villafranchianus, P lwnensis, Eobania oermicwlaria sur- (Kotsakis et al., 1992). They are composed of species vive up to the Olivola F.U. whrle Carychium (Carychiel- which now live in northern countries (as Helicella itala) la) puisseguri and G(V) dehmi suwle up to the Tasso or of species still living in Italy but which are less com- F.U. The latter species was also found in a quarry near mon or widespread in northern Italy (as Tiichia hupida). Fighille associated with remains of a bovid, Leptobos It is clear that the non-marine mollusc assemblages of merlai-furtir.tar (pers. com. F. Masini). The prosobranchs the Middle (and Late) Pleistocene are influenced by cli- are the most characteristic group of the late Villafran- matic oscillations. Generally the politypic assemblages chian Mammal Age: genera (such as Stepbania, Prosostlte- are related to temperate and temperate-warm climatic nia, Tournouerina, Neumayria) and endemic species of phases, whilst during cold periods strictly oligotypic as- central Italy (Tyrrhenian side) which disappear at the semblages are recorded (Esu et al., 1989). end of the Early Pleistocene. A smali group of species is of Pliocene origÀ (Theodoxus (Neritaea) groyanus, Pro- Aurelian Mammal Age. sosthenia opdta, Tournouerina belnensis, Micromelania The transition from the Galerian malacofaunas to (Goniochilus) zitteli, Emmericia umbra, Melanopsis ffi- the Aurelian ones is not detectable. Assemblages of nis). Some species of prosobranchs and pulmonates oc- cold-temperate and warm-temperate character similar to curring in the Tasso F.U. do not seem to be present in those of the Galerian Mammal Age continue to alterna- the Farneta F.U.: 7 (N.) groyanus, Stephania bronni, Pro- te. They are mainly composed of living species. In some sosthenia menegbiniana, Tànousia litboglyphoides, Neu- Aureiian deposits of central-southern Italy various ex- mayria priscillae, M. (G.) zitteli, C.(C) puisseguri, G(V) tinct species are recorded Hydrobia melii, Tanowsia subo- de h rni, A n cy lus parmopb o rus. oata, Belgrandia zilchi (Esu & Girotti, 1991; Esu & Two species, Lithoglyphus aff. L. jahni and Theo- Kotsakis, i996). Two species, T isseli and J. malatestai, doxus aff. T danubialis, occur for the first time in sedi- which were present in the Galerian assemblages, become ments of the Stirone and Crostolo Rivers (northern extinct during this interval. At the beginning of the Au- Italy) of Emilian age; these species survive in the same relian Mammal Age, during a warm climatic oscillation, area vp to the Galerian Mammal Age. The Stirone and a tropical iiving species, Melanoides tuberculaa (see trsu Crostolo Rivers have an Adriatic drainage, but the Emi- 6c Girotti, 1991), was present in the southern part of lian sediments they cut also contain some elements the Itaiian peninsula, probably transported there by 380 E. Gliozzi et al.

birds. In general the Aurelian malacological assemblages ferent ostracod faunas are necessary in order to make are more useful for ecostratigraphic studies than for bio- out a distribution chart which could be of use for bio- stratigraphic or biochronological purposes. chronological purposes.

Ostracods. Villafranchian Mammal Age. There are few sedimentary series bearing ostra- Distribution chart. cods referable to the early and middle Villafranchian Studies on the Italian Plio-Pleistocene continental Mammai Ages: the R.D.B. quarry (Villafranca d'Asti, (fresh water and brackish) ostracod faunas are scarce. Piedmont), Arondelli and Roatto (Piedmont) (Ghozzi, Only in the last few years particular attention has been 7996) referable to the Triversa F.IJ., Cava Toppetti-Fosso paid to their analysis, since they proved to be very use- Bianco Unit (Todi, Umbria) (probably referable to the ful for detailed palaeoecoiogical reconstructions. F{ow- Montopoli F.U.), Cava Toppetti-Ponte Naja Unit (Costa ever, at the moment it is still not clear whether ostracod S. Giacomo F.U.) (Gliozzi, unpubl. data) and, finally, faunas are also a valid tool for Plio-Pleistocene biochro- Dunarobba (Umbria) which is referable, on the basis of nology, mainly for two reasons: 1.) in several cases the the mollusc assemblage (Ambrosetti et al., I995a), to a life of the species is much longer than the time period generic middle Villafranchian (GIíozzì, unpubl. data). under consideration (for example, and only to cite some All these sequences pertain to a limnetic domain, except species which are very frequent in Plio-Quaternary fresh for Cava Toppetti-Ponte Naja Unit which records a water deposits, Cyclocypris laevis is known from the Da- brackish environment. nian to the present day, Daruinula steoensonl from the In this wide time interval, corresponding to the Oligocene to the present day, and Candona neglecta, Po- Middle and Late Pliocene, Candona candida, Stenocypris tamocypris fuloa and Pseudocandona marchica from the sp. Candona fabaeformis, Paralimnorythere compressa, Late Miocene to the present d^ù; Z) ostracods are Pseudocandona parallela, Cyprideis sp. 1 and Cypridopsis benthic organisms which are particularly sensitive to the par"ua seem to appear. All these species characterise the environmental characteristics, and consequently their modern Italian ostracod fauna, except for: 1) Stenoqpris presence,/absence in one particular deposit does not rep- sp., a genus at present wideiy distributed at tropical lati- resent only a biochronological data but must be filtered tudes, which was found as a fossil at the R.D.B. quarry; through an environmental interpretation. In fact it is at present it is recorded as being present in Italy by known that ostracods are used for both the definition of Ghetti & McKenzie (1981) represented by the species S, classical zones inside a conventional biostratigraphy (or fontinalis e S. major and is only found in lakes with a biochronology), taking into account the same environ- hot water input. According to McKenzie & Moroni mental conditions (e.g. littoral marine environment bio- (1986), these forms can be considered "foreign guests", zones), and for the creation of ecological community thar is forms which have been accidentally introduced units (or ecozones) inside an ecostratigraphy (Mar- by man with foreign good transports;2) Cyprideis sp. L, tinsson, 1973).In both cases, in order to avoid the verti- an extinct form found only at Cava Toppetti-Ponte Naja cai distribution of the taxa being altered by ecological Unit, which seems to be limited ro the Late Pliocene. background noise, it is necessary to work with a large This form pertains to the Cyprideis torosa group, but dif- number of sites pertaining to different environments for fers from C. torosa in several morphometric characters each time period. such as its dimensions and general shape (Gliozzi, unpu- Despite the scarcity of sites where ostracods have blished data). been analysed so far in Italy, Fig. 4 shows a tentative Three more species, Limnorythere aff. L. inopina- chronological ordering of Italian ostracod faunas known ta, Metacypris cordata and Pseudocandona compressa, were from freshwater and brackish (oligo-mesohaline) envi- recorded by Di Napoli (1962) in the lignite deposits at ronments. F{ere, the presence of several selected taxa Morgnano, Torgiano, Spina, Ilci, and Collazzone (Tibe- from different Italian deposits is reported. Only those rino Basin), which he referred to the Late Pliocene. At species not recorded prior the Middle Pliocene have present these deposits are no longer visible, and conse- been taken into consideration; almost all these species quently a stratigraphic control using the recent Plio- are still living in the Italian Peninsula, so Fig. 4 is only Pleistocene chronological sequence is impossible; these indicative of their first occurrence. The sites are referred species have therefore not been put into the sbck of to the Mammal Ages and, when possible, also to the species which appeared during the Middle-Late Pliocene. Faunal Units which represent the datum-point for the The deposits referable to the late Villafranchian Plio-Pleistocene biochronology. Mammal Age (Early Pleistocene) are more numerous, It is important to point out that this table repre- but they are mainly located in central Italy: several loca- sents only a starting point and that other studies on dif- lities in the Tiberino Basin (Umbria) [Bastardo I and II, Biocbronology of Plio-Pleistocene of mammals, molluscs and ostracods in laly

Deruta and Collazzane, generically referred to the first Pleistocene (post-Santernian) (Barberi et al., 1995)l and part of the Early Pleistocene; Bastardo III, ry V and VI, Montallegro (Sicily) (Decima, 1963) correlated with the referable to a post-Santernian (Di Napoli, 1962), and Emilian. The majority of these series record brackish, Villa S. Faustino (Ambrosetti et al., I995b) which can oligo-mesohaline environments. During the late Villa- be correiated with the Tasso F.U.], Pietrafitta (Umbria) franchian Mammal Age the present living species of (Gliozzr, unpublished data), several localities of the Rieti Cyprideís torosa, Cytberissa lacustris, Herpetorypris che- Basin (Latium) [Case Strinati, Fosso Filundici, Apoleg- oreuxi, Candona leoanderi, Lirnnoclthere sanctipatricii gia, Vicchiagnone, correlated with the Tasso F.U. and and Poamocypris zschokkei all seem to appear. Moreover Belmonte Sabino, probably referable to a younger Early Limnocythere inopinata, Metacypris cordata and Pseudo-

PLIOCENE MIDDLE LATE Ch ronostratigraphy EARLY PLEISTOCENE PLEISTOCENE )LEISTOCENE MIDDLE I LATE Early Middle Mammal ages Villaf ran. Vilìaf ran. Late Villafranchian Aurelian

lr o È o .ì lr ii N @ t! o o o o a g) .D zl q l- È.. b o q q o tt o o o o o o € î .; (t) tr- () F I o o 6 c lr o o E o u, o l o g 5 E 5 o o c c qa F 'c o -c 'È o o o E ! o o c o c 6 o f F o o € 5 c F .6 '6 @ î F r Ò. f.- 'c N t- c0 tL F ! o c c î z lt U g t g g g qo l a o o f o U Lrl uJ o o '6 O .E 6 z .6 o g c _g '- O z z 6 o o .P o É .9 6 _g) (L l (, c z = 'È .9 = o = o= q= =(, '- F F z z r o 6 z o = o È F U) U) lt) tr F F 6 uJ a o- z z U) z z U) a fL uJ al tr tr F tr d) fL ff) co z co J (, 0- z ul u z uJ z O z z O o o a J a o J o z o u) F tr J o o- J fr o l o- - - É. F z z c0 m z U J LU t! U U É. É. E. ts cc Í. J J u U F F z F !! (I z z J z z z LJ C' J ) f co I g o U g cc g (-) cc (r È À ts F F cc ts F = L J Candona candìda a c{. a o a o -Stenocvpris sp, a Candona fabaeformis a a a a Paralimnoc!,there compressa o cî. cî. o a t uvpnders sp. 1 a Cypridopsis parua a vprideis torosa o o a a a a o o o a Lr m noc!,the re rnopr nala a o a 'Metacvoris cordata o o a a a a a a C)4herissa lacustris a a a a HerDetocvDns chevreuxl a c cf. o a Candona levanderì a a Limnocvthere sanclroatricli a Polarnocvoris zschokkei a cl a o Candona pubescens a a a a Cypridopsis vidua a a a t Leptocythere (Amnicythere) tallax a '"Candona caudata a '"Candona lobipes a 'Scottia browniana a ""tucypns prgra o o a uypncercus relrcuratus a Prionocypris zenkeri a a Cypris pubera o o a araltmnocvînere messanal a a Eucyprìs clavata a Eucvpris serrala cl

Fio 4 Biochronology of selected living and extinct brackish and freshwater ostracods of Italian Peninsula from Middle Pliocene to Late o Pleistocene. Legend: 'f species at present distributed at circumtropicai latitudes; species at present living out of Ita|y at higher latitudes; oo species at present widely distributed at higher latitudes; + estinct species. J82 E. Gliozzi et al.

candona co?rrpressa are now recorded with certainty. Vi- cercus reticulatus, Herpetocypris intermedid, Psychrodro- thin this group, the presence of Cyprideis torosa. needs to mus olioacews, Cypris pubera and Paralimnocythere messa- be explained better: Cyprideis torosa ís recorded in the nai. Actually, this noticeable abundance of forms is literature as being frequent in the European Pliocene, mainly due to the fact that some deposits of the Liri and sometimes it is reported as a species while at other Valley correspond to cold periods (Isotope Stage 6) and times it is reported as C. gr. torosa. It is not possible to many of the forms which occur during this stage are exclude the fact that this species may reaI|y have appea- northern forms, presently found in central and northern red during the Pliocene, although all the Italian deposits Europe (Scottia broaniana, Herpetoqtpris intermedia e in which its presence has been controlled are of Pleisto- Psycbrodromus olioaceu) (which can be considered as cene age. "cold guests") or forms which, although they are still found in Italy (only in scattered localities of Piedmont Aurelian Mammal Age. and Lombardia), show their main distribution arealoca- ted in more northern areas (Candona cawdata, Candona At present no studies on ostracod faunas referable lobipes, Eucypris pigo).Taking inro account the fact that to the Galerian Mammal Age have been carried out, the Liri Valley ostracod faunas are the only cold osrracod while there are numerous studies on Aurelian deposits, assemblages known in Italy, it is not possible to ascertain unfortunately all of which are located in Latium (central whether these cold species entered Italy only from Stage Italy): Torre in Pietra (Moiinari Paganelli, 1978) (correla- 6 onwards, or whether they migrated southwards during ted with Oxygen Isotope Stage 9), Pianura Pontina (lr4i- other earlier cold periods. It is interesting to underline gliara 47 borehole) (Barbieri et a1., in press) referable to the presence during the Aurelian Mammal Age of an ex- Stages 9, ?7, 6 and 5, several localities along the Liri tinct form, Icptocytbere and the first occurrence Valley (Devoto, 1965) referable to Stages 9-8 and, tentati fallax, oÍ Para I imnocythere messa nai. vely, 7, Valle di Castiglione borehole, referable to Stages The beginning of the late Aurelian Mammal Age 7,5 and 3 (Gliozzí &Mazzini, in press), and Lago Lun- (Isotopic Substage 5e) records the appearance of the pre- go borehole, referable to Stage 5 (Calderoni et aI., 1994). sent-day living species Pseudocandona sarsi, Eucypris cla- The species that seem to appear at the beginning of the aata and, perhaps, Eucypris sey'rata, characterised by ha- Aurelian Mammal Age are numerous, and this is partly ving oniy one summer reproductive season (Bronshtein, due to the important taxonomic studies which have 1e47). been carried out on Aurelian ostracod faunas by Moli- nari Paganeili (1978) and Devoto (1965): Candona pube- scens, Herpeto cypris reptans, Cypridopsis vidua, Leptoqttbe- Achnouledgemmts. re fallax, Candona caudata, Pseudocandona lobipes, Scot- This paper was supported by CNR grants, Centro Studio per tia brouniana, Eurypris pigra, Prionorypris zenkeri, Cypri- il Quaternario e l'Evoluzione Ambientale, Roina.

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