Plio-Pleistocene Biogeography of Italian Mainland Micromammals
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Tassos Kotsakis1, Laura Abbazzi2, Chiara Angelone1, Patrizia Argenti3, Giancarlo Barisone1, Flaviano Fanfani2, Federica Marcolini4 & Federico Masini5 1 Università di Roma Tre 2 Università di Firenze 3 Università di Perugia 4 Università di Pisa 5 Università di Palermo Plio-Pleistocene biogeography of Italian mainland micromammals Kotsakis, T., Abbazzi, L., Angelone, C., Argenti, P., Barisone, G., Fanfani, F., Marcolini, F. & Masini, F., 2003 - Plio-Pleistocene biogeography of Italian mainland micromammals - in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 313-342 [ISSN 0923-9308] Published 1 December 2003 The analysis of the distribution of small mammals in the Italian mainland during the Plio- Pleistocene and their immigration in the Peninsula indicates the presence of many species of orien- tal European origin, a few iberoccitanic elements and some emdemic species. The Italian penin- sula belongs to a western Mediterranean bioprovince. The north-eastern region of Italy is a transi- tional biogeographical zone between this province and the central European and Balcanic areas. Correspondence: Tassos Kotsakis, Chiara Angelone & Giancarlo Barisone: Università di Roma Tre, Dipartimento di Scienze Geologiche, Largo San Leonardo Murialdo 1, Roma 00146, Italy, e- mail [email protected]; [email protected]; Laura Abbazzi & Flaviano Fanfani, Università di Firenze, Dipartimento di Scienze della Terra, Via La Pira 4, 50121 Firenze, Italy, e-mail lab- [email protected]; [email protected]; Patrizia Argenti, Università di Perugia, Dipartimento di Scienze della Terra, Piazza Università, 06100 Perugia, Italy e-mail [email protected]; Federica Marcolini, Università di Pisa, Dipartimento di Scienze della Terra, Via Santa Maria 53, 56126 Pisa, Italy e-mail [email protected]; Federico Masini, Università di Palermo, Dipartimento di Geologia e Geodesia, Corso Tukory 131, 90134 Palermo, Italy e-mail [email protected] Keywords: Italy, Plio-Pleistocene, micromammals, paleobiogeography INTRODUCTION only large-sized ‘micromammals’ such as Many Italian fossil mammals of Plio- Marmota, Castor, Hystrix and lagomorphs, Pleistocene age are known and several locali- easier to discover during excavations, have ties of the Italian peninsula are famous been carefully studied. After the World War among vertebrate paleontologists. However, 2, the works of A. Pasa and G. Bartolomei till the Second World War the number of fos- illustrated for the first time the rich micro- sil remains of small mammals was very low. mammalian Quaternary faunas of north-eas- A few paleontologists (C.I. Forsyth Major, A. tern Italy. During the last decades of the 20th De Gregorio, C. Bosco, R. Fabiani & D.M.A. century an increasing number of researchers Bate) illustrated remains of rodents and followed the first pioneers. insectivores (especially from the islands) but Italian micromammalian assemblages are 313 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003 mainly found in Pleistocene deposits. ning faunal lists of small mammals and the Nevertheless, Neogene remains are very bibliographical references are reported in important for paleobiogeographical recon- chronological order in the Appendix. A syste- struction of Italian bioprovinces. During Late matic revision or a study of biochronological Miocene, mammalian evidences point out the succession of several groups of Italian fossil presence of three different sub-provinces: the micromammals is a topic of some general Apulia-Abruzzi, the Sardinia-Tuscany (both papers followed in this publication: Chaline with strong endemic character) and the (1977), Bartolomei (1980), Masini & Torre Calabrian-Sicilian subprovince (with evident (1987), Kotsakis (1988), Bon et al. (1991), African affinities; Torre et al. 2000). Only Brunet-Lecomte et al. (1994), Zanalda since the latest Miocene Italy has firm con- (1994a), Masini et al. (1996, 1998), Maul et tacts with the rest of Europe, as testified by al. (1998), Argenti (1999), Argenti & the presence of several immigrated taxa Kotsakis (1999), Fanfani (2000), Marcolini (most of them from eastern Europe, and a (2002). In the biochronological charts the few elements from western Europe; Kotsakis position of some localities is aproximate and et al. 1997). their chronological attribution is not certain. Italian Pliocene micromammalian fossil assemblages are very few and very unbalan- RUSCINIAN ced from a geochronological point of view. Only a single rodent tooth (M/1), collected in Remains of Ruscinian age are almost un- an ?early Middle Pliocene (Lina Barbera known from the Italian mainland (this also pers. comm.), brackish-marine deposit near applies to macromammals), whilst fossils of Ariano Irpino (Campania, southern Italy) has Villányian age (Fig. 1) are relatively few, but been found. This molar has been attributed to sufficient for a paleobiogeographical analysis. a murid, Centralomys sp. (T. Kotsakis & P. The number of Biharian fossiliferous sites Argenti unpublished data). The genus is bearing small mammals is quite low too and known from the late Turolian site of only Toringian assemblages are abundant, Brisighella (Emilia-Romagna, northern Italy) even if not well distributed in the Peninsula with the type species Centralomys benericet- (Fig. 2). tii (DE GIULI) (De Giuli 1989). The age of the In the present paper we examine micro- brackish-marine sediments of Ariano Irpino mammalian assemblages of the Italian main- ought to correspond to the uppermost part of land, excluding endemic assemblages from MN 15 unit. The more advanced faunas from Sicily, Sardinia, smaller islands and paleois- the Gargano Archipelago (Apulia, southern lands of the Gargano Archipelago because of Italy) fossil islands (where many micromam- their peculiar problems. Among fossil assem- malian endemic lineages evolved) have been blages we chose to examine just the recently assigned to the latest phases of Miocene described or re-examined ones and chose to and/or to the earliest phases of Pliocene ignore isolated remains (if not important for (Abbazzi et al. 1998). A discussion of the paleobiogeography). Fossiliferous sites are affinities of these endemic faunas is not a correlated with MN unites (Mein 1990) for topic of the present work. the Villányian, with zones of Fejfar & A migration from the European mainland to Heinrich (1990) and also with the Faunal Sardinia near the Zanclean-Piacenzian boun- Units (F.U.) succession system proposed by dary, corresponding to the uppermost part of Azzaroli (1977) and emended by Gliozzi et MN 15 unit (Angelone & Kotsakis 2000, al. (1997), a system accepted by all the 2001), is suggested in recent works. Such a Italian paleontologists working on continental migration very probably followed an Italian- Plio-Pleistocene fossil animals, both vertebra- Corsican route but no fossil evidence of the tes and invertebrates. The locality list, contai- ancestors of Sardinian Prolagus and Rhaga- 314 KOTSAKIS et al.: Plio-Pleistocene micromammals of Italy Superzones 1. 9 0 1 2 2. 2. 2. 6 7 8 9 0 1 2 3 Figure 1 Biochronology of Villányian sites of Italy with micromammals. In bold character recently studied localities with relevant micromammalian remains.The chronological position of some localities is approximate. podemus (known in southern France) has Mimomys hajnackensis see Mörs et al. 1998) been found in Italy till now. + Mimomys stehlini zone by Fejfar & Heinrich (1990) and Fejfar et al. (1998), has EARLY VILLÁNYIAN (Fig. 3) been collected in a sequence of mudstones, Cascina Arondelli in the Triversa Valley clays and fine-grained sandstones. The two (Piedmont, north-western Italy) is a famous rodent species collected in the fluvio-lacustri- Italian site of Early Villányian age. An ne sediments of Fornace RDB (Piedmont, assemblage of insectivores, lagomorphs and north-western Italy) belong to taxa also pre- rodents has been described from this site. sent at Arondelli. Azzaroli (1977) formalised This fauna, assigned to MN 16a by Mein the Triversa F.U. as a unique fossil assembla- (1990) or to the Mimomys hassiacus (= ge of small mammals (collected in Cascina 315 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003 Figure 2 Biochronology of Biharian and selected Toringian sites of Italy with micromammals. In bold character recently studied locali- ties with relevant micromammalian remains.The chronological position of some localities is approximate. Arondelli) and large mammals (collected in moment there is no paleontological support to Fornace RDB). After Lindsay et al. (1997) the opinion of Lindsay et al. (1997). the age of macromammals from Fornace Micrommamals of these assemblages belong RDB is older than that of the micromammals to species of large European distribution. from Arondelli. For this reason they assigned Similar taxa of insectivores and also rodents the large mammals of Fornace RDB (the clas- are present at Arcille (small sequence of clays sical Villafranchian fauna) to MN 15. For the and lignite) and San Giusto (marine-brackish 316 KOTSAKIS et al.: Plio-Pleistocene micromammals of Italy Figure 3 Distribution of micromammals in Italy during Early Figure 4 Distribution of micromammals in Italy during Late Villányian. Arondelli - a) Apodemus alsomyoides; b) Mimomys Villányian. Rivoli Veronese - a) Dinaromys allegranzii; b) cf. stehlini; c) Mimomys hassiacus. S. Giusto - d) Mimomys stehlini. Ellobius sp.;