Plio-Quaternary mammal fossiliferous sites of Umbria (Central Italy)

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INTRODUCTION

The fossil mammalian fauna in Umbria is rich. The majority of the Plio-Pleistocene mammal faunas, disco- vered in Umbria, are represented by macromammals.

These are more abundant because in the past only large animal remains were collected, as they are easier to discover and to recover them. Only large sized species of rodents (beaver, porcupine) and some lagomorphs were collected. However the fossil faunas of Umbria are represented by five orders of large mammals and four of micromammals. These fossil mammal remains represent all the orders and the majority of the families collected in the Pleistocene sites of Europe.

Present day knowledge about terrestrial mammal fau- nas of the Plio-Pleistocene of Umbria does not allow attempting a detailed biochronology for the whole region. The chronological distribution of the mammalian fossiliferous layers is clear in the central and southern part of Umbria, because of the abundance of the fossils.

In the northern part of the region there are instead many gaps due to the scarcity of fossiliferous beds. The distribution of the fossiliferous localities is referred to Faunal Unit (F.U.) successions and Mammal Ages fol- lowing the biochronological scale of the continental suc- cession in the Italian peninsula (Gliozzi et al., 1997). The fossil mammal remains found in Umbria are not uniform- ly distributed within the Plio-Pleistocene deposits, they are in fact concentrated in Late Pliocene and Early Pleistocene ones. Not all analysed localities are referred to a F.U. because in many sites only isolated fossils were discoveded or the fossils come from ancient collections, which are impossible to review; the localities are thus referred to Mammal Ages (Fig. 1). Some mammal assem- blages are kept in cave deposits. In recent times two gen- eral works regarding some faunal assemblages of Umbria are published (Azzaroli, 2001; Petronio et al., 2003).

The location of the fossiliferous sites is reported in Fig. 2.

EARLY VILLAFRANCHIAN ASSEMBLAGES (MIDDLE PLIOCENE)

The only mammalian assemblage found in Umbria, referred to the Middle Pliocene was collected in the lig- nite mine of Spoleto. This can be referred to Triversa F.U. (early Villafranchian, Middle Pliocene) (Kotsakis, 1986) for the presence of Mammut borsoni, Anancus arvernensis and Tapirus arvernensis. This assemblage is characteristic of the Early and Middle Pliocene (Ruscinian and Early Villafranchian). The beginning of Triversa F.U. should be around 3.2 M. years b.p. as sug- gested by paleomagnetic analysis at Fornace R.D.B.

(Villafranca d’Asti, Piedmont) (Gliozzi et al., 1997).

Some Mastodon arvernensis remains were found near Montoro (Tuccimei, 1891) and referred to Pliocene age.

MIDDLE VILLAFRANCHIAN ASSEMBLAGES (LATE PLIOCENE)

The assemblage of Cava Toppetti (Todi) is referred to Costa S. Giacomo F.U. (middle Villafranchian, Late Pliocene) (Abbazzi et al., 1997). The Apodemus domi- nans remains of Cava Toppetti are the same age as those found at Rivoli Veronese. Middle Villafranchian faunas are poorly represented in Italy.

LATE VILLAFRANCHIAN ASSEMBLAGES (LATEST PLIOCENE AND EARLY

PLEISTOCENE)

The beginning of the late Villafranchian (Olivola F.U.) is marked by the appearance of Pachycrocuta brevi- rostris, Panthera gombaszoegensis and new species of Eucladoceros while dogs become widespread (Gliozzi et al., 1997). The mammal fossil assemblage collected in Pantalla (Todi) likely refers to the Olivola F.U. (late

ABSTRACT - This work aims at providing the most complete catalogue of fossil mammal localities of Umbria, ranging from Pliocene to Holocene. The catalogue consists of fossil sites, known and reported since the XVIII century, in which faunal assemblages or even just single fossil remains were found. The catalogue allowed to biocronologically allocate the Plio-Pleistocene mammal assemblages of Umbria. The Pliocene assemblages are scarce, but the one found in Cava Toppetti (Todi) that was calibrated with paleomagnetic data is rather important.

A very meaningful and rich fossil assemblage ascribable to the Plio-Pleistocene passage comes from Torre Picchio (Montecastrilli, Terni). Lower Pleistocene faunas are abundant and important since they are distributed on great part of the region’s territory. One example for all is the very rich collection from the Pietrafitta lignites. It is possible to ascribe, more or less in detail, such assemblages to the various Faunal Units. There are several Middle Pleistocene assemblages, but they are often not very rich. Upper Pleistocene faunas end Holocene faunas mostly come from Karst cavities.

KEYWORDS: Fossiliferous sites, Mammals, Plio-Pleistocene, biochronology, Central Italy.

PLIO-QUATERNARY MAMMAL FOSSILIFEROUS SITES OF UMBRIA (CENTRAL ITALY)

Patrizia Argenti

Dipartimento di Scienze della Terra, Università degli studi di Perugia

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Villafranchian, latest Pliocene) (Gentili et al., 1997): car- nivors mainly characterize this assemblage.

Torre Picchio is referred to the Olivola F.U. (or between the Olivola F.U. and Tasso F.U.) for stratigraph- ical correlation with marine strata, despite the presence of Mimomys medasensis, known from Late Pliocene of Ibero-occitan area. This arvicolid survives in Italian peninsula for longer time (Girotti et al., 2003).

As regards the Parrano-Frattaguida fauna, this is quite heterogeneous and its stratigraphic position is unknown, it could be referred to the middle - late Villafranchian,

after Ambrosetti (pers. com.) in this site is present Megantereon cultridens and Stephanorhinus etruscus (Verri, 1886; Clerici, 1896; Gliozzi et al., 1997).

The assemblages found at Villa Spinola and Ponte S.Giovanni may be referred, very probably, to Tasso F.U.

for their geographical and geological location (Argenti

& Sardella, 2003). Fornace Ferrini-Ponte S.Giovanni remains are referred to the late Villafranchian for the correlation with the fauna of Villa Spinola and Piscille, which comes from the same levels (Argenti & Sardella, 2003).

Fig. 1 - Biochronological distribution of the fossiliferous site of Umbria.

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Fig. 2 - Distribution map of the fossiliferous sites: 1) Lignite mine of S. Croce (Spoleto); 2) Cava Toppetti (Todi); 3) Cava est Casigliano and Casale Violino (Casigliano, Acquasparta); 4) Fighille (Citerna); 5) Ellera of Corciano (now Filiale Lancia); 6) Casale Montecchio of Casalina (Deruta); 7) near S. Maria di Ciciliano (Montecastrilli); 8) near Orvieto; 9) Parrano - Frattaguida - S. Faustino; 10) Fontignano (Perugia); 11) Piscille (Perugia);

12) Villa Spinola and near Ponte S. Giovanni (Perugia); 13) Some isolated localities in the neighbourhhods of Perugia (Colombella, Civitella d’Arno, S. Martino Delfico, Castel delle Forme, from S. Egidio to Deruta);14) Vigna Nuova (Piegaro); 15) Pietrafitta (Piegaro); 16) near Sanfatucchio (Vaiano); 17) Cava S. Faustino (Villa S. Faustino - Massa Martana); 18) Colle Pizzuto (Sangemini); 19) near Capitone (Narni); 20) near Colvalenza (Todi); 21) near Chioano (Todi); 22) Due Santi (Todi); 23) Colle Lame (Capitone, Narni); 24) M. Peglia (S. Venanzo); 25) Promano (Città di Castello); 26) near Marsciano; 27) near Lubriano (Corbara); 28) Poderetto (Pietrafitta); 29) M. Cucco cave; 30) Pozzo della Piana (Titignano); 31) Pantalla (Todi); 32) Civitella de’ Pazzi (Orvieto); 33) near Bettona; 34) Colle S. Andrea (S. Faustino); 35) Colle Violino (S. Faustino); 36) Casale le Grotte (S. Faustino); 37) near Pozzuolo (Trasimeno Lake); 38) near Montone (Umbertide); 39) near Paciano; 40) near Montefalco (Foligno); 41) near Montoro, Monte Campano lignite mine (Narni); 42) Rena between S. Fortunato e S. Martino in Colle (Perugia); 43) near M. Dell’Abbate between Perugia and Gubbio; 44) Railway tunnel of S. Costanzo (Perugia); 45) Fornace Ferrini, Ponte S. Giovanni (Perugia); 46) Torre Picchio (Montecastrilli); 47) near Gubbio (Scheggia); 48) Triponzo; 49) Taizzano (Narni); 50) Collelungo- Ravigliano, Tresa Valley; 51) Tane del Diavolo (Parrano); 52) S. Firmina- Fighille (Pistrino); 53) Colleluna (Terni).

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The Tasso F.U. is characterized by a change in faunal assemblage with the appearance of typical Quaternary elements such as Mimomys savini, Canis (Xenocyon) falconeri, Hippopotamus, Praeovibos and Leptobos val- lisarni (Gliozzi et al., 1997). The fossils from S.Fau- stino, Cava est Casigliano, Colle Violino, Casale Le Grotte and Colle S. Andrea are referred to Tasso F.U.

(Ambrosetti et al., 1995; Sardella et al., 1995).

Vigna Nuova assemblage is probably referred to Tasso F.U., for its stratigraphic position with respect to Pietrafitta and its the paleontological assemblage (main- ly for the presence of Megantereon cultridens). Fighille clays are referred to late Villafranchian, Tasso F.U. on the basis of mollusc assemblage (Gastrocopta (Vertigopsis) dehmi Schlimckum & Strauch, Multi-den- tula helenae Esu and Parmacella (Parmacella) cf. P.

(P.) sp.1 sensu Manganelli & Giusti) (Ciangherotti &

Esu, 2000).

Capitone must be probably referred to the end of the late Villafranchian because of the presence of Stepha- norhinus etruscus, represented by an advanced form (Ambrosetti, 1972). This induces the formulation of a hypothesis regarding a stratigraphical correlation between Capitone and Torre Picchio the proof of which is left to further investigation. Anyway the Totte Picchio fauna is clearly older than the more advanced rhino found in Capitone.

The Farneta F.U. is characterized by a new situation:

Leptobos and Eucladoceros are still present, while the Megaceroides clade (M. obscurus) and the voles Microtus (Allophaiomys) chalinei and M. (A.) cf. M. (A.) ruffoi appear (Gliozzi et al., 1997). Ellera remains are referred to the Farneta F.U. for the mammal assemblage (Ambrosetti et al., 1994). The mammal assemblage from Pietrafitta is referred to the Farneta F.U. and it is typical of this faunal unit with Megaceroides obscurus, the voles of subgenus Microtus (Allophaiomys) and Mimomys pusillus. The faunal remains of Sanfatucchio and S. Maria di Ciciliano are referred to the late Villafranchian, probably the upper part, because they were collected in the upper part of the S. Maria di Ciciliano Formation. Casalina sands are referred to the latest Villafranchian because Mammuthus meridionalis collected in this locality belongs to an advanced form.

The mammal remains from Fontignano are referred to the upper part of late Villafranchian, for the advanced form of hippo recorded (Kotsakis, 1992).

GALERIAN ASSEMBLAGES (LATEST EARLY PLEISTOCENE AND MIDDLE PLEISTOCENE)

The Colle Curti F.U. is characterized by the appareance of voles such as Microtus (Allophaiomys) nutiensis and Microtus (Allophaiomys) burgondiae, Pliomys lenki and the first form of Megaceroides verticornis. The mammal assemblages found in the M. Peglia cave are referred to the Colle Curti F.U. (Meulen, 1973; Gliozzi et al., 1997), for the presence of Microtus (Allophaiomys) nutiensis

and Microtus (Allophaiomys) burgondiae. The mammal remains found in the sands of Promano (Città di Castello) (lower assemblage) are referred to the lower part of the early Galerian, probably Colle Curti F.U., mainly for the presence of Mega-ceroides verticornis and an advanced form of Mammuthus meridionalis (Argenti et al., in prep.). The fossil remains found near Marsciano are referred to the early Galerian for the presence of a cervid classified as Megaceroides cfr. M. verticornis. The ele- phant of Fighille is probably referred to early Galerian unfortunatly the sediments where it was recorded do not exist anymore. The elephant’s site is on the Tiber terrace that covers the clay quarry. Another assemblage was found in Promano sands, referred to middle Galerian, but after Colle Curti F.U., for the presence of Stephanorhinus hemitoechus (Argenti et al., in prep.).

AURELIAN ASSEMBLAGES (LATE MIDDLE PLEISTOCENE AND LATE PLEISTOCENE)

A marked faunistic renewal occurs at the beginning of the Aurelian, with the extinction of some Galerian forms such as the archaic megacerine cervids, the more archa- ic elaphine deer and the arrival of Canis lupus and Ursus spelaeus. The sands overlapping Poderetto lignites are referred to the middle Aurelian for their radiometric age (160-170.000 years b.p., Ambrosetti, pers. comm.).

During late Aurelian Dama dama (modern form) appeared, Capra ibex, Coelodonta antiquitatis, Mam- muthus primigenius, Marmota marmota spread over Italy and several taxa of large herbivores and carnivores progressively disappeared in correspondance with the successive phases of climatic deterioration. The mam- mal assemblage from the M. Cucco cave is referred to late Aurelian, for the presence of Dama dama, Capra ibex, Coelodonta antiquitatis. The assemblage from Tane del Diavolo is referred to Aurelian, for the presence of Equus ferus, probably for the presence of abundant archeological remains of upper Paleolithic (Calzoni, 1956).

The assemblage from Pozzo della Piana is referred to the cool or cold phase of late Aurelian for the presence of Capra ibex.

HOLOCENE DEPOSITS

For the third mammal assemblage of Promano an attri- bution to the late Aurelian or to Holocene is uncertain whilst the mammal assemblage found at Triponzo belongs clearly to Holocene.

ACKNOWLEDGEMENTS - Many sincere thanks to Prof.

T. Kotsakis, Prof. P. Ambrosetti and Prof. O. Girotti for the critical revision of manuscript. I am indebted to Dott.ssa M.C.

De Angelis, Soprintendenza Archeologica dell’Umbria and Prof. C. Giovagnotti, Istituto di Pedologia agraria, Università degli Studi of Perugia.

Many thanks also to Mrs. H. Terry and Dott.ssa N. Minciotti for the revision of the english manuscript.

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Serv. Geol. Ital., 71, 51-57.

Paganelli A. (1960) - Primi saggi per uno studio pollinologico del deposito lignitifero di Pietrafitta (Umbria). N. Giorn.

Bot. Ital., 67, 601-605.

Petronio C., Argenti P., Caloi C., Esu D., Girotti O. & Sardella R. (2003) - Updating Villafranchian mollusc and mammal faunas in Umbria and Latium (Central Italy). Geologica Romana, 36, 369-387.

Piperno M., Segre A. G. & Segre Naldini E. (1985) - Monte Peglia (Umbria). In “I primi Abitanti d’Europa”, 115-119.

Principi P. (1922) - Alcune osservazioni sul Quaternario del- l’Umbria centrale. Atti R. Acc. Naz. Lincei, s. 5, 31, 56-59.

Principi P. (1930) - Alcune osservazioni intorno all’età dei travertini di Ellera nei dintorni di Perugia. Atti R. Acc. Naz.

Lincei, s. 6, 11, 493-495.

Rabeder G. (1981) - Die Arvicoliden (Rodentia, Mammalia) aus dem Pliozän und dem älteren Pleistozän von Niederösterreich. Beitr. Paläont. Österr., 8, 1-373.

Ramaccioni G. (1936) - Scheletro di ippopotamo rinvenuto nel Villafranchiano dei dintorni di Perugia. Atti Soc. Tosc. Sc.

Nat. P. V., 45, 31-34.

Rook L. (1995) - Pannonictis nestii (Carnivora, Mammalia) from the late Villafranchian of Pietrafitta (Umbria, Italy).

Preliminary note. Eclogae Geol. Helv., 88, 853-864.

Rustioni M. & Mazza P. (1993) - The Late Villafranchian bear from Pietrafitta (Perugia, Central Italy). Palaeontographia Ital., 80, 51-62.

Sardella R., Di Stefano G. & Petronio C. (1995) - The Villafranchian mammal faunas from the Tiber River Basin (Umbria, Central Italy). Il Quaternario, 8, 509-514.

Simonelli V. (1917) - I mammiferi fossili della Caverna di Monte Cucco. Mem. R. Acc. Sc. Ist. Bologna, s. 7, 3 (1915- 1916), 272-284.

Taliana D., Alessio M., Allegri M., Capasso Barbato L., De Angelis C., Esu D., Girotti O., Gliozzi E., Improta S., Mazzini I. & Sardella R. (1996) - Preliminary results on the

“Grotta del Lago” Holocene deposits (Triponzo, Nera River Valley, Umbria, Central Italy). Il Quaternario, 9,

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745-752.

Tuccimei G. (1891) - Sui mammiferi fossili dell’Umbria e della provincia di Roma. Atti Acc. Pont. Nuovi Lincei, 44, 274-275.

Tuccimei G. (1895) - Sopra i resti fossili di mammiferi trovati alla Villa Spinola presso Perugia. Atti Acc. Pont. Nuovi Lincei, 48, 44-46.

Tuccimei G. (1896) - Resti di Felis arvernensis nel Pliocene della Villa Spinola presso Perugia. Mem. Acc. Pont. Nuovi Lincei, 12, 1-27.

Ugolini R. (1921) - Sopra alcuni resti di rinoceronte del Pliocene lacustre di Pietrafitta in Provincia di Perugia. Riv.

Ital. Paleont., 27, 15-18.

Verri A. (1886) - Azione delle forze nell’assetto delle valli con appendice sulla distribuzione dei fossili nella Valdichiana e nell’Umbria interna settentrionale. Boll. Soc. Geol. Ital., 5, 416-454.

Accettato per la stampa: Marzo 2003

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In the following list the number in haevy type is referred to the number of the site in the map.

Locality: Montoro, lacustrine clay (41).

Mammal assemblage: Anancus arvernensis (Croizet & Jobert) Age: Pliocene.

Collection: University Museum of “La Sapienza”, Roma.

References: Tuccimei, 1891.

Locality: lignite mine of S. Croce (Spoleto) (1).

Mammal assemblage: Mammut borsoni (Hays), Anancus arvernensis, Tapirus arvernensis Croizet & Jobert, Cervidae indet., Castor sp.

Age: Triversa F.U.

Collection: Toni Collection now preserved in the Liceo Scientifico of Spoleto (Perugia).

References: Capellini, 1888; Clerici, 1894, 1895, 1896; Meli, 1895; GE.MI.NA., 1963; Kotsakis, 1986.

Locality: Cava Toppetti (Todi) (2), Ponte Naia Formation.

Mammal assemblage: Stephanorhinus cf. S. etruscus (Falconer), Pseudodama cf. P. rhenana (Dubois), Proca- preolus sp., Leptobos sp., Apodemus dominans Kretzoi.

Age: Costa S. Giacomo F.U.

Collection: Dipartimento di Scienze della Terra, Università degli Studi of Perugia.

References: Ambrosetti et al. 1987a; Basilici, 1992; Abbazzi et al., 1997; Argenti, 1999.

Locality: Pantalla (Todi) (31) in the S. Maria di Ciciliano Formation.

Mammal assemblage: Lynx cf. L. issiodorensis (Croizet &

Jobert), Panthera cf. P. gombaszoegensis (Kretzoi), Canis etr- uscus Major, Equus sp., Sus cf. S. strozzii Meneghini, Pseudo- dama cf. P. nestii (Major), Leptobos sp., Apodemus dominans, Aves.

Age: probably Olivola F.U.

References: Gentili et al., 1997a.

Locality: Torre Picchio (Montecastrilli, TR) (46), in the S.

Maria di Ciciliano Formation.

Mammal assemblage: Canis cf. C. arnensis Del Campana,

?Baranogale sp., ?Homotherium crenatidens Fabrini, Mammuthus meridionalis (Nesti), Equus cf. E. stenonis Coc- chi, Stephanorhinus cf. S. etruscus, Sus strozzii, Eucladoceros dicranios (Nesti) vel ctenoides (Nesti), Axis nestii (Major), Cervidae gen. et sp. indet., Leptobos cf. L. etruscus (Falconer), Bovidae gen. et sp. indet. (medium size antilope), Bovidae gen.

et sp. indet., Mimomys medasensis Michaux, Prolagus sp., Oryctolagus cf. O. lacosti (Pomel).

Age: probably Olivola F.U.

Collection: Soprintendenza per i Beni Archeologici of Umbria.

References: Argenti, 1999; Girotti et al., 2003.

Locality: Cava est Casigliano and Casale Violino (Casigliano, Acquasparta), (3), S. Maria di Ciciliano Formation.

Mammal assemblage: Pachycrocuta brevirostris (Aymard), Rhinocerotidae indet., Pseudodama sp., Castor sp.

Age: Tasso F.U.

Collection: ex-church of S. Tommaso of Terni.

References: Basilici, 1992; Ambrosetti et al., 1995.

Locality: clay quarry of Fighille (Citerna) (4).

Mammal assemblage: Leptobos cf. L. vallisarni Merla and incisors of rodents.

Age: Tasso F.U.

References: Gentili & Masini (pers. comm.); Argenti, 1999.

Locality: Vigna Nuova (Piegaro) (14).

Mammal assemblage: Megantereon cultridens (Cuvier), Equus sp., Rhinocerotidae indet., Sus sp., Cervidae indet. (a big form and a very small one), Bovidae indet.

Age: probably Tasso F.U.

References: this paper.

Locality: Cava S. Faustino (Villa S. Faustino - Massa Marta- na) (17).

Mammal assemblage: Megantereon cultridens, Mammuthus meridionalis, Equus stenonis, Pseudodama nesti s.l, Euclado- ceros sp.

Age: Tasso F. U.

References: Ambrosetti et al., 1995; Sardella et al., 1995.

Locality: Colle S. Andrea (S. Faustino) (34).

Mammal assemblage: Pachycrocuta cf. P. brevirostris, Cervidae indet., Leptobos cf. L. vallisarni, Castor sp.

Age: Tasso F.U.

Locality: Colle Violino (S. Faustino) (35).

Mammal assemblage: Stephanorhinus etruscus.

Age: Tasso F. U.

Locality: Casale le Grotte (S. Faustino) (36).

Mammal assemblage: Eucladoceros sp.

Age: Tasso F. U.

Locality: Ellera of Corciano (now Filiale Lancia of Ellera) (5).

Mammal assemblage: remains referred to “Elephas antiquus Falconer & Cautley, Cervus elaphus Linnaeus, Bos primige- nius Bojanus, Sus scrofa Linnaeus, Ursus spelaeus Rosenmüller & Heinroth” and other rests of bovids were collected in the past and now probably destroyed.

In the collection are preserved: cf. Panthera gombaszoegensis, Stephanorhinus cf. S. etruscus, Hippopotamus cf. H. antiquus Desmarest, cf. Eucladoceros sp. (senezensis type, big form), Pseudodama sp., Leptobos cf. L. vallisarni (big form, enough evolved).

Age: Farneta F. U.

Collection: Museum of Corciano.

References: Principi, 1930; Ambrosetti et al., 1994.

Locality: Pietrafitta (Piegaro) (15), in the lignite.

Mammal assemblage (and other fossil remains): Sorex sp., (size S. minutus Linnaeus), Macaca sylvana florentina Cocchi, APPENDIX

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Panthera gombaszoegensis, Pannonictis nestii (Martelli), Ursus etruscus Cuvier, Mammuthus meridionalis vestinus Azzaroli, Equus sp., Stephanorhinus cf. S. hundsheimensis (Toula), Megaceroides obscurus Azzaroli (= M. boldrinii Azzaroli), Pseudodama farnetensis Azzaroli, Leptobos aff. L.

vallisarni, Castor fiber Linnaeus, Mimomys pusillus Mèheley, Microtus (Allophaiomys) cf. M. (A.) ruffoi (Pasa), M. (A.) cha- linei Alcade, Agustì & Villalta, Oryctolagus cf. O. lacosti.

Abundant fossil remains were also found of: birds (Anatidae, Ardeidae, Podicipedidae and Tetraonidae), reptiles (Emys orbicularis (Linnaeus), Natrix sp., Vipera ammodytes (Linnaeus)) and amphibians (Latonia cf. L. ragei Hossini, Rana gr. R. ridibunda Pallas). The numerous fish remains have not yet been studied. In the lignite some levels bearing a rich molluscan fauna have still not been studied in detail: Ancylus sp., Bithynia tentaculata, Lymnaea bucciniformis, Unio sp..

Finally the macrofloristic fossil remains (leaves, seeds, fruits and stems) and the pollens are referred to: Pinus, Tsuga, Quercus, Pterocarya, Carya, Zelkova, Liquidambar.

Age: Farneta F.U.

Collection: the Thermoelectric Station “Città di Roma” of Pietrafitta of ENEL (the Italian National Electricity Company) of Pietrafitta and in the Boldrini Collection (Pietrafitta).

References: Ugolini, 1921; Moretti, 1947-49; Ambrosetti et al., 1987b; Ambrosetti et al., 1992a, Ambrosetti et al., 1992b;

Argenti, 1990; Gentili, 1991; Azzaroli & Mazza, 1993; Mazza et al., 1993; Rustioni & Mazza, 1993; Rook, 1995; Ikome, 1995; Abbazzi, 1996; Gentili et al., 1996; Gentili et al., 1997b;

Argenti 1999; Gentili et al., 1999; Barisone et al., (in press), Argenti & Kotsakis (in prep.) for mammals. Kotsakis &

Gregori, 1985; Delfino, 2002 for reptiles and amphibians.

Ambrosetti et al., 1979; Conti & Esu, 1981; Esu, (pers. comm.) for molluscs. Paganelli, 1960; Bertoldi, 1988 for pollen and macrofloriristic remains.

Locality: Casale Montecchio of Casalina (Deruta) (6).

Mammal assemblage: Mamuthus meridionalis, cervids, bovids, carnivores and rhinoceros.

Age: Early Pleistocene.

References: Gentili, 1985.

Locality: near S. Maria di Ciciliano (Montecastrilli) (7).

Mammal assemblage: some remains of a cervid.

References: Basilici (pers. comm.).

Locality: near Orvieto (8).

Mammal assemblage: some remains of rhinoceros.

Collection: probably Museo dell’Opera del Duomo of Orvieto, no further information is known.

References: Ambrosetti (pers. comm.).

Locality: Parrano - Frattaguida - S. Faustino, in sands (9).

Mammal assemblage: “Megantereon cultridens, Equus sp., Canis sp., Elephas sp., Stephanorhinus cf. S. etruscus, Equus stenonis, Cervus sp., Bos sp. and an Antilope”.

Collection: Paleontological Museum of Università degli Studi of Firenze, Natural History Museum of London and Museo Capellini of Bologna.

References: Verri, 1886; Clerici, 1896; Ambrosetti (pers.

comm.).

Locality: Fontignano (Perugia) (10).

Mammal assemblage: Mammuthus meridionalis and Hippopotamus antiquus.

Collection: now we do not know where they are preserved, it is possible that they are held in the Verri Collection.

References: Verri, 1886; Ramaccioni, 1936; Kotsakis, 1992.

Locality: Piscille (Perugia) (11).

Mammal assemblage: some remains of a cervid.

Collection: Cicioni Collection, Università degli Studi of Perugia.

Locality: Civitella dè Pazzi (Orvieto) (32).

Mammal assemblage: a tusk of Mammuthus meridionalis.

Collection: it has been destroyed.

References: Ambrosetti, (pers. comm.).

Locality: some isolated localities in the neighbourhood of Perugia (Colombella, Civitella d’Arno, S. Martino Delfico, Castel delle Forme, from S. Egidio to Deruta) (13).

Mammal assemblage: “Elephas meridionalis Nesti, Elephas primigenius Blumenbach, Elephas antiquus Falconer &

Cautley, Rhinoceros merki Jäger”.

Collection: now some of them are preseverd in the Istituto di Pedologia Agraria, Facoltà di Agraria, Università degli Studi of Perugia, previously belonging to the ancient Canali Collection.

References: Bortolotti, 1904; Principi, 1922.

Locality: near Pozzuolo (Trasimeno Lake) (37).

Mammal assemblage: “Elephas meridionalis and Rhinoceros sp.”

Collection: R. Istituto Superiore Agrario now Facoltà di Agraria, Università degli Studi of Perugia.

References: De Angelis d’Ossat, 1918.

Locality: near Paciano (39).

Mammal assemblage: a mandible of “Hyaena striata Zimmerman” (the remains are referred to Pachycrocuta brevi- rostris).

Collection: belonging to the ancient Canali Collection, its present location is unknown.

References: Bortolotti, 1905.

Locality: near Montoro, in the lignite mine of Monte Campano (Narni) (41).

Mammal assemblage: remains of “Rhinoceros cf. etruscus Falconer”.

Collection: their present location is unknown.

Locality: Rena between S. Fortunato and S. Martino in Colle (Perugia) (42).

Mammal assemblage: some remains of rhinoceros.

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Locality: near M. Dell’Abbate, between Perugia and Gubbio (43).

Mammal assemblage: “Rhinoceros etruscus”.

Collection: the ancient Canali Collection, their present location is unknown.

Locality: railway tunnel S. Costanzo (Perugia), in the clay with

“Helix” (44).

Mammal assemblage: “Elephas sp.”, a fragment of a tusk.

Locality: Fornace Ferrini near Ponte S. Giovanni (Perugia), in the clay with “Helix” (45).

Mammal assemblage: “Hyaena (small size) probably Hyaena striata, Rhinoceros ertuscus, Cervus etueriarum Croizet &

Jobert” (the remains are referred to Cervidae indet.).

The ancient Canali Collection, collected by Canali in the XVIII century, belonged to the free University of Perugia until the begining of this century, and preserved in the Geological Museum of the Regio Istituto Superiore Agrario. After this period the collection, together the other fossil remains and the mineralogical collection, was lost and destroyed, the few remains today are preserved in the Istituto di Pedologia of Facoltà di Agraria, University of Perugia.

Locality: Villa Spinola and near Ponte S. Giovanni (Perugia) (12).

Mammal assemblage: “Elephas meridionalis” (the remains are now referred to Mammuthus meridionalis), “Rhinoceros sp., Equus sp., Sus sp., Bos sp, Cervus sp.”, “Felis arvernensis Croizet & Jobert ” (the remains are now referred to Panthera gombaszoegensis), “Elephas sp.”, “Hyaena striata” (the remains are referred to Pachycrocuta brevirostris),

“Rhinoceros etruscus” (the remains are referred to Stephanorhinus etruscus).

Age: late Villafranchian.

Collection: Cicioni Collection and R. Istituto Superiore Agrario now Facoltà di Agraria, Università degli Studi of Perugia.

References: Tuccimei, 1895, 1896; Bortolotti, 1903; De Angelis d’Ossat, 1918.

Locality: near Sanfatucchio (Vaiano) (16).

Mammal assemblage: few remains of elephant (dorsal verte- bra) and of Leptobos aff. L. vallisarni (jugal tooth).

Locality: Colle Pizzuto (Sangemini) (18).

Mammal assemblage: a tibia referred to Leptobos sp.

Locality: near Capitone (Narni) (19).

Mammal assemblage: a skeleton of Stephanorhinus etruscus.

Collection: Paleontological Museum of Università degli Studi

“La Sapienza”, Roma.

References: Ambrosetti, 1972.

Locality: near Colvalenza (Todi) (20).

Mammal assemblage: some remains of Mammuthus meridion- alis and Stephanorhinus cf. S. etruscus.

References: Ambrosetti (pers. comm.).

Locality: near Chioano (Todi) (21).

Mammal assemblage: Cervidae indet. and Leptobos sp..

Locality: Due Santi (Todi) (22).

Mammal assemblage: remains of a cervid.

Locality: Colle Lame (Capitone, Narni) (23).

Mammal assemblage: an antler of Eucladoceros sp., some remains of Equus stenonis and some other fossils of verte- brates.

Locality: in “yellow sands” at an unknown locality between Trasimeno Lake and Chiusi Lake.

Mammal assemblage: “Equus sp., Canis sp. and Elephas sp.”

Age: Early Pleistocene-Middle Pleistocene.

Collection: their present location is unknown may formerly have been kept in Roma, R. Scuola d’Applicazione per Ingegneri.

References: Meli, 1895.

Locality: the cave of M. Peglia (S. Venanzo) (24). The first dis- covery of the layer was in July 1955. In the first excavation, under the direction of A. C. Blanc of Istituto di Paleontologia Umana of Rome, some remains of micromammals and few of macromammals were found. In 1963, during the second excavation, under the direction of G.H.R. von Koenigswald of Utrecht University, the majority of the micrommals were collected. Therefore the collection of the mammal assemblage of M. Peglia is divided into two parts, the first preserved in Rome and the second in Utrecth.

Mammal assemblage: Also the fossil remains constitute two assemblages. The first assemblage, from red clays and “terra nera”, called Mimomys blanci assemblage, is characterized by the abundance of Mimomys blanci and Microtus (Allophaiomys) nutiensis, and include Talpa cf. T. fossilis Petényi, Sorex runto-

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nensis Hinton, Beremendia fissidens (Petényi), Asoriculus cf. A.

castellarini (Pasa), Macaca sylvana Linnaeus, Homotherium crenatidens Fabrini, Canis cf. C. etruscus, C. cf. C. arnensis Del Campana, Ursus etruscus, Leptobos sp., Glirulus sp., Ungaromys nanus Kormos (= Ungaromys meuleni Rabeder, according to Rabeder (1981) this species is more evolved than U. nanus, but it is possible to consider it as synomym of U.

nanus), Pliomys episcopalis Mèheley, Mimomys savini Hinton, Mimomys blanci Meulen, Microtus (Allophaiomys) nutiensis Chaline, Apodemus cf. A. sylvaticus (Linnaeus), Apodemus sp., Lepus terraerubrae Kretzoi.

The second assemblage, called Allophaiomys sp. B assemblage, was collected in the clays and in the breccias overlaied the cal- cite layer. It is characterized by the abundance of M. (A.) bur- gondiae, the faunal list is: Talpa cf. T. fossilis, Sorex runtonen- sis, Beremendia fissidens, Neomys cf. N. newtoni Hinton, Ungaromys nanus, Mimomys blanci, Microtus (Allophaiomys) burgondiae (Chaline), Apodemus cf. A. sylvaticus, Apodemus cf. A. maastrichtiensis Kolfschoten, Lepus sp.

Age: Colle Curti F.U.

Collection: the Institut of Earth Science of the University of Utrecht, the Museo di Paleontologia Umana of Roma.

References: Blanc, 1955; Meulen, 1973; Piperno et al. 1985;

Basilici et al. 1991.

Locality: Promano (Città di Castello) (25) from sands.

Mammal assemblage: Pachycrocuta brevirostris, Canis ex gr.

arnensis - mosbachensis Soergel, Ursus arctos Linnaeus, Mammuthus meridionalis cf. M.m. vestinus, Equus gr. bres- sanus Viret - suessenbornensis Wüst, Stephanorhinus etr- uscus, Hippopotamus antiquus, Megaceroides verticornis (Dawkins), Pseudodama farnetensis, Bison cf. B. schoetensac- ki Freudenberg, Castor fiber.

Age: Colle Curti F.U.

Collection: Raccolta Civica, Biblioteca Comunale of Città di Castello (PG).

References: Bartolini, 1975; Argenti et al., in prep.

Locality: near Marsciano (26).

Mammal assemblage: some remains of Megaceroides cf. M.

verticornis.

Age: early Galerian.

Collection: its present location is unknown.

References: Ambrosetti et al., 1987a.

Mammal assemblage: Ursus deningeri Reichenau, Stephanorhinus hemitoechus (Falconer), Hippopotamus tiberi- nus Mazza.

Age: middle Galerian.

Locality: near Lubriano (Corbara) (27).

Mammal assemblage: some remains of Elephas antiquus.

Age: Middle Pleistocene or Late Pleistocene.

Locality: Fighille (Citerna) (4), in the quarry between the clay and the overlapped sands.

Mammal assemblage: a skull of Elephas cf. E. antiquus.

Age: Middle Pleistocene.

Collection: no longer exist.

References: Lippi Boncampi, 1963; Bartolomei 1969.

Locality: Taizzano (Narni) (49).

Mammal assemblage: Elephas antiquus.

Collection: ex-church of S. Domenico in Narni (TR).

References: IRRES, 1990.

Locality: S. Firmina- Fighille (Pistrino) (52).

Mammal assemblage: an astragalus of Bison sp.

Collection: Museo Civico of Sansepolcro (AR).

Locality: near Bettona (33).

Mammal assemblage: a fragment of antler of a giant deer.

Age: ?Middle Pleistocene.

Locality: near Montone (Umbertide) (38).

Mammal assemblage: some remains of “Elephas antiquus”.

Locality: near Montefalco (Foligno) (40).

Mammal assemblage: some remains of “Elephas sp.”.

Locality: Collelungo-Ravigliano, Tresa Valley (50).

Mammal assemblage: “Elephas antiquus”, “Rhinoceros lep- torhinus Cuvier” (the remains are referred to Rinocerotidae indet.), “Bison priscus Bojanus” (the remains are referred to Bovidae indet.), “Hyaena sp.”

Collection: probably Museo Capellini of Bologna.

References: Verri, 1886.

Locality: Colleluna (Terni) (53).

Mammal assemblage: tusks of Elephas antiquus.

References: Ambrosetti, Basilici & Girotti, pers. com.

Locality: in the sands with volcanic sediments of Poderetto (Pietrafitta) (28).

Mammal assemblage: Ursus sp., Equus sp., Rinocerotidae indet., Cervidae indet. (a big form and a small one), Bovidae indet., Testudinata indet. and lithic industry.

Age: middle Aurelian (preliminary radiometric data 160 - 170.000 years B.P.).

Collection: Thermoelectric Station “Città di Roma” of Pietrafitta of ENEL (the Italian National Electricity Company) of Pietrafitta.

Locality: the M. Cucco cave (29).

Mammal assemblage: Rhinolophus ferrumequinum (Schreber), Myotis (Myotis) myotis (Borkhausen), Myotis (Myotis) blythi (Tomes), Myotis (Selysius) bechsteini (Kuhl), Myotis (Selysius) emarginatus (E. Geoffroy), Felis silvestris Schreber, “Felis pardus” (Linnaeus) (the remains are referred to Panthera par- dus (Linnaeus)), Vulpes vulpes (Linnaeus), Ursus spelaeus,

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Ursus arctos, Equus cf. E. caballus Linnaeus, Coelodonta antiquitatis (Blumenbach), Dama dama (Linnaeus), Cervus elaphus Linnaeus, Bos sp., Capra ibex Linnaeus, Rupicapra rupicapra (Linnaeus).

Age: late Aurelian

Collection: Museo Capellini of Bologna, Dipartimento di Scienze Geologiche, Università degli Studi “Roma Tre” of Roma, Museo Civico of Orvieto (TR).

References: Bonarelli, 1891; Capellini, 1889, 1890; Simonelli 1917; Capasso Barbato et al. 1985; Capasso Barbato &

Kotsakis 1986.

Locality: Pozzo della Piana (Titignano) (30).

Mammal assemblage: Cervus sp., Capra ibex and human rests.

Age: late Aurelian.

Locality: Tane del Diavolo (Parrano) (51) from cave.

Mammal assemblage: Equus ferus Bodeart.

Age: probably late Aurelian.

Collection: Soprintendenza Archeologica per l’Umbria.

References: Calzoni, 1956.

Locality: near Gubbio (Scheggia) (47).

Mammal assemblage: “Elephas primigenius”.

Age: unknown.

Collection: probably Palazzo dei Consoli, Gubbio (PG).

References: Cuvier, 1834; Bonarelli, 1891; Esu & Girotti, 1991.

Mammal assemblage: Equus ferus, Sus scrofa, Bison cf. B.

priscus Bojanus.

Age: Holocene.

Locality: Grotta del Lago, Triponzo (48).

Mammal assemblage: Sorex araneus Linnaeus, Crocidura sp., Rinolophus ferrumequinum, Myotis bechsteini, Canis familiaris Linnaeus, Meles meles (Linneaus), Sus scrofa, Cervus elaphus, Capreolus capreolus, (Linnaeus), Bos taurus Linnaeus, Apodemus (Sylvaemus) sp., Clethrionomys glareolus (Schreber), Microtus (Terricola) sp. Arvicola terrestris (Linnaeus).

Age: Holocene (6500-3100 anni BP).

Collection: Soprintendenza per Beni Archeologici of Umbria.

References: Taliana et al., 1996.