Bollettinodella

Societn Paleontologic aItalianaPubblicato sotto gli auspicidel Consiglio Nazionale delle Ricerche

Volume 42rr.312003

MTJCCHI MODENA

Bollenino della Societh Paleontohgica ltaliana 42 (3), 2003 rssN 0375-7633315-32r I pt. Modena, Dicembre 2003

Early Pleistocene large mammals from Villa Spinol,(Perugia, Central Italy)

Patrizia ARcT,NTIDipartimento di Scienze della Terra-

Universit) di Perugia

INTRODUCTION

"Sul versante orientale della collina di Perugia a

sud della citt), dove il Tevere uscito dalla stretta di P.S. Giovanni, si allarga nella Valle tra M. Malbe eDeruta, poco a monte della conflu enza con ilChiascio, si trova Villa Spinola nei pressi di S.Martinello

" (Text-fig. 1).tVith these words Tuccimei described the fossiliferouslocaliry, not far from Perugia, where in 1894-95Marquise Virginia Spinola found several mammalbones. He produced the followitg faunal list: Felisaruernensis, Elephas meridionalis, Rhinoceros etrctscr,ls,Bos sp., Equus sp., Sus sp. and Ceruus sp.

The fossi l bones were distr ibuted amongMarquises Spinola themselves, the St. ApollinareMuseum in Roma, the Cicioni Collection

-and the

Bellucci Collection in Perugia.The large fel id bones are part of the CicioniCollection, today stored at the Universiry of Peru gia,the mammoth remains belong to the Bel lucc iCol lect ior, today in the

".urtody of the

Soprintenden za per i Beni Archeologici dell'Umbria.The geological section of S. Martinello, described

byTirccimei (1895), included from the bottom to thetop, blue marls, blue marly sands, yellow sands andnot well cemented gravels; the deposits were subhor-izontal or slightly tilted. The section was exposed onthe south side of the hill, towards the Tiber River

Valley and Ponte S. Giovanni and could be seen fromVilla Spinola.

The fossil bones were probably discovered duringthe construction of the Roma-Perugia railway(Cioci, 2001). For this reason it is possible to delim-it the fossiliferous deposit exposure to the right sideof the Tiber Rivet in the area berween S. Martinelloand Piscille (today included in Perugia) that is part ofthe wide Marquises Spinola estate. Today the area isstrongly urbanised and the stratigraphic succession isnot exposed anymore.

Field observations and data inferred from litera-ture suggest that the fossiliferous deposit can be con-sidered as part of a fluvial sequence correlatable withthe S. Maria di Ciciliano Formation (Basilici, 1995),which widely outcrops in the south-western branchof the Tiber river basin.

Several remains of Mammuthus meridionalis werefound in 1894-'95: fragments of a skull, a tusk and ahemimandible, some ribs and vertebrae, partiallydestroyed during the excavations. All the bones arereferable to an old male (the well restored tusk wasstored at Villa Spinola).

The skeleton elements came from the yellowsands; the top of the tusk reached the overlyitg grav-els (Tirccimei, 1895). Later on fossil remains belong-irg to a young specimen of the same species, togeth-er with a hemimandible of a rhino, referred byTuccimei to Rhinoceros etrusctts, were found and

Raffaele SnnoELLADipartimento di Scienze della TerraUhiversitl di Roma "La Sapienzi'

KEY V/ORDS - Mammalia, Panthera gombaszoegensis, Mammuthus meridionalis, 7Early Pleistocene, Cennal ltaly.

ABSTRACT - At the end of the I gth century, in the surroundinss of Villa Spinol^a (Pboscideans, rhinos, equids, suids, ieruids, bouids arl a pantber were disioueied. Unfirtunately,worh focuses on the siitt auailable fossil bones ofPanthera sombaszoegensis and Mammuthus rin thi Cicioni and Bellucci Collictions. Such' anallsis ,nZbltt to poi-nt out considerations on tstratigraphy of the deposit, not uisible anlmore. fhi faunA cAn be' referred to the Late Vilkfan

fossiEzaiio, iuggett tltat the fossil bones iome fom a" paleosoil that has neuer been recognised be1

RIASSUNTO - [ srandi mammiferi del Pleistocene Inferiore di Villa Spinola (Peru

furono rinuenuti pressi VTlk Spinola (Perugia) resti di mammiferi fossiti riferibili a proboscidatuna pantera. Nel corso degli inni, purtroppo, gran parte di qiesti"reperti toro anditi perduti.Panthera gombaszoegetrr'ir r Mamm.rtlinr rileridionalis (for*a eioluta) ancora disponibili,Bellucci. f'analisi di iali resti ha permesso considerazioni silla bioronologia della fauna e sullLa fauna i riferibile alVitkfanihiano superiore (Pleistocene Inferiore). Consideraiioni sul tipaotti fottili pr:ouengano da ui paleosuolo, mai segnalato in precedenzt.

3r6 P. ARGENTI, R, SARDELLA

stored at the St. Apollinare Museum (Tuccimei,I 896). Other isolat.d teeth, which Tuccimei generi-cally referred to Bos sp., Equrys sp.-, Sus sp. a1d Ceruussp., were collected and sto?ed af the scientific cabinetof the Archiepiscopal Seminary in Perugia (Tirccimei,

I 896).The fossil mammal remains of Villa Spinola need-

ed a systematic revision that could allow a correct andupdaied biochronological definition. Unfortunatelyrnort of the remains were lost so the Authors couldrevise only part of the fossil collection.

,^ ; F,IFIH:H) l)3! th,^The first occurrence of Pantltera gombaszoegensis

Kretzoi l-Panthera toscAna (Schaub)] has been con-sidered by Azzaroli (1977) as one of the main bio-events of the beginnitg of the Pleistocene (Late

Villafranchian, OTivola T.U.). At present, OlivolaF.U. is referred to the latest Pliocene (Gliozzi et Al.,1997). This large felid probably appeared in Asia dur-irg Pliocene and spread over Europe at lhq beginnilsof- the Pleistocen-e, until the end of the MiddlePleistocene; however, an African origin of this taxoncannot be ruled out (Hemmer, 1971; Hemmer et al.,2001). Panthera gombaszoegensis occurs in Italy _atOlivola (Magra Valley), in several localities of theUpper Vald"arno (Tusclny), in Val di Chiana(Tirscany) (Ficcarelli 6{ Torre, 1968), in the TiberRiver basin (Umbria) at Pantalla (Gentili et Al., 1997)and at Ellera di Corciano (Ambrosetti et dl., 1995), ittthe Thvernelle basin (Umbria) at Pietrafitta (Ge ndli etdl., 1997), and at Pirro Nord (Apulia) (Pantltera sp. inBeaumont, 1979 and unpublished data) in LateVillafranchian faunas. The Eurasian jaguar is still Pre-sent in Galerian faunas at Monte Peglia (Umbria)(Piperno et dl., 1984) _"ttd Slivia (Friuli VeneziaGiulia) (A-brosetti et al., 1979; Bon et al., 1992).The occurrence of Pantltera pardus in a LateVillafranchian faunal assemblag-e from Argentario(Tuscany) was pointed out by Baschieri 6{ Segre(1958). The fossil material of this localiry needs to berevised but it is possible that the fossil referred to theleopard could be related to Panthera gombaszoegensis.

This large felid is not very frequent in Italy; itschronoloei lal distr ibut ion

' incl ldes the

'Late

Villafr"tt.Tti"n and the Early Galerian (Gliozzi et al.,1997). In Europe this species was recorded in severallocalities with Villafranchian and Galerian faunas:Tegelen (Netherlands) (Koenigswald, 1960),Miedonia basin (Greece) (Koufos, 1992), \Testbury(Ehgland), Untermassfeld and Mosbach (G.1gany),Atapuerca (Spain) and L'Escale (France) (Tirryer,19i7 , with references), Koneprusy (Czech Republic)(unpublished data), Gombaszoeg and Vertdsszolos(Hungary) (Janossy, I 990).

Re-cent works focused the systematics of this largefel id (Hemmer et al . , 2001; O'RegxrI, 2003).

Text-fig. I - Location map of Villa Spinola.

Hemmer et al. (2001) refer the Late Villafranchianand Galerian fossil remains of this pantherine cat toEuropean chronosubspecies of the iaguar Pantheraonct' (Panthera onct toscAnA for the Late Villa-franchian specimens and Panthera onct gombaszoegen-sis for the Galerian ones).

In this paper the use of the taxonomical termPanthera gombaszoegensis is preferred because, even ifboth Euiasian and American jaguars show strongaffinities, the fragmentary nature and the scantinessof the remains cinnot exclude a specific distinctionberween the rwo groups.

The fossil remains from Villa Spinola belong toone specimen, represented by isolated canines, therwo hemimandibles, the second cervical vertebra,some thoracic vertebrae and fragments of the left andright femurs (Pl. 1, f igt . l -5).-

The canines are developed, conical, with the ryPi-cal longitudinal grooves, the jugal teeth show Pt-Poand PolM, indeies falling in the range of the lateVillafranchian Eurasian jiguar. The morpholory ofthe jugal teeth is similar

-to that of modern larg.e

felids, ih. lower carnassial shows a weak talonid, withthe protoconid and paraconid divided by a quite widefurrbw, larger than in other Pleistocene sPecies refer-able to the genus Panthera (Argant, I 99 I ).

The spe-imen from Villa Spinola shows a strong

Miralduolo

l t . ) |

V I I I Ot:i!!:

PoderePiana

f 'at t SPinola

Spinolo

Osperetton" ; , /

uS.Martirto i "

''---t€//iccr 3-

' tal

P ARGENTI, R. SARDELLA, EARLY PLEISTOCENE TARGE MAMMALS FROM VIL

1 a

2b

3b

Pl. I

1 b

EXPLANATION OF PLATE I

Figs. l-5 - Panth-era gombaszoegqnlis Kretzoi from Villa Spinola.I - left mandible: a) labial view, b) lingual view; 2 - right mandible: a) labial vie,labial vigw, b) lingual view; 4 - particu-lar view of the tiaces of the vegetable root,traces of the vegeiable roors on the left caput femuri.S c a l e b a r s = l c m .

upper r ight canine: a)particular view of the

3 1 8

Measures Villa Spinola

17,6rr,733,5

33,4

61,21 7 , 514,216,79,723,010,024,012,0

Thb. I - Measures (ntm) of the remains of Panthera gombaszoe-gensis from Villa Spinola. L - lenght;

'W' = width.

P, and is larger than the Late Villafranchian Pantf.ltftorn Olivolf ".d Upper Valdarno (Tab. 1),-but it is

smaller than the Galerian European ones. Accordi.g

to Hemmer et al. (2001), such characters have a

biochronological significance, thus the specime n

from Vil la "Spitto[i

can be placed in the late

Villafranchian group (referred by these Authors to P

oncA toscana).

THE SOUTHERN MAMMOTHMA]VT M UTH US M E RI D I O NAL I S

The southern mammoth, or a closely related form(Palombo, 1994; Ferretti, 1998), f irst occurred in

E.rrop. in Late Pliocene. It became widespl:"d

throughout the Northern Hemisph.ere (includi.g

Nottl i America) in the Early Pleistocene (Late

Vil lafranchian, Thsso F.U.) with the subspecies

Mammuthus meridionalis meridionalis (Azzaroli,

1977; Lister, 1996; Ferretti, 1998). European speci-

mens younger than those of the Thsso F.U. seem to

restii/ the occurrence of two distinct lineages 9.fsouthern mammoths, derived from M. meridionalis

merid.ionalis (Ferretti, 1999). The first lineage is reP-

resented in Italy by M. meridionalis uestinus Azzaroli,

1977 and related forms, as the one occurri.g at

Pietrafitta (lkome, 1995; Gentili et al., 1996), in a

mammal fauna referable to the Farneta F.U. These

specimens are large-sized with particularly.large tusks

r"d specialised sliull morplrolory; the molars do not

show any noteworthy differenle from those of M.

meridion-alis meridionalis. Fragmentary remains seem

to attest that this lineage survived in Italy until the

beeinnins of the Middle Galerian (Sl iv ia F.u.,

Mi'ddle Pieistocene; Ferretti, 1999).The second evolutionary lineage, which is widely

distributed, includes the material referred to M.

meridionalis ex 8r' tamanensis (= uoigtestd'ensis)

(Siniaya Balka, Russia, Dubrovo, I 964; and

Voigtsiedt, Germany, Dietrich, 1958) that existed in

the -

second part of Early Pleistocene (Foronova'

2001). This group has *oi. hypsodont molars, with

a higher tt,rf,'tb.i of plates; thise features probably

were an adaptation to a grass-r.ich diet. The forms

belongitg to this second gio.tp show metrical charac-

ters th"at lre berw een M. meridionalis meridionalis and

middle Pleistocene M. trogontherii (Pohlig, 1 885).

Their distribution in tVesiern Europe is from the

Colle Curti F.U. (Early Galerian) to the Slivia F.U.(Early Middle Galerian). In Italy jh. Mammuthus

,.-"i., from Imola (lmola Sands Formation; Slivia

F.U.; Ferretti 1999) may belong to this grouP.M. meridionalis meridionalii is well represented in

the Late Villafranchian (Olivola, Matassino, Thsso

local faunas) of Tuscany. The Late Villafranchianmammal assemblages suggest the occurrence of a rel-

atively cold climatlc phaiE, with the diffusion of for-

est/parkland biotopei (Gliozzi et al., 1997). In the

yo.ttg.t faunal assemblages referable to the Farnetap.u.

"M. meridionalis *tildionalis is replaced by M

meridionalis uestinus. This subspecies is considered a

hiehlv specialised feeder and its occurrence can be

li;kea with the diffusion of savannah parkland with

high trees, shrubs and grass. (Ferretti 6( Croitor,

ZdOl ). At Pietrafitta the abundance of this advanced

form of southern mammoth, together with the com-

position of the mammal assemblage, fit well with the

occurrence of a palustrine environment' surroundedby grassland ettii.h.d with rare trees and shrubs(Ambrosetti et al., 1992).

In Umbria this species was also found in a Middle

Pleistocene deposit at Promano (Cittl di Castello)(unpublished data).

The tusk of the elephant, found by Tuccimei in

1894-'95, at Colonnett" Properry, within the Villa

Spinola estate, was discovered berween the yellow

randr and the gravels, and today is._stored at the"soprintende n { Archeologica pgr. I'umbria" (D9

Aneelis, pers. comm.). The mandible, some ribs and

oth-'er fragments of the same specimen were found

near S. [Iartinello within the Villa Spinola estate(Perugia) (Calzoni, 1 956).

Tfi'e diameter of the tusk indicates that the speci-

men was an adult. The tusk is 3 m long, but now it is

divided into three Parts. The tusk is curved and rwist-

ed in the rypical shape of Mammuthus. The revision

of the tusk "llo*t to refer it to Mammutltus cf. M.

meridionalis.The hemimandibles were restored and their frac-

tures were filled at the time of the discov ery. Such

restoration does not allow the evaluation of the defor-

mation degree of the fossil remains (Text-fig. 2).The leli hemimandible (lnv. N". 291722) lacks

the symphysis behind the Posterior side of the alveo-

lus. ih.tooth is M2, the only one that was still work-

irg when the animal died. This tooth is rather worn-

P. ARGENTI, R. SARDELLA

L upper rieht CW ui,p.t tleht CHeighl of the mandibleat C-P/3 diastemaHeight of the mandiblebehind M/ IL left jugal teeth seriesL lower left C'W

lower left CL left P 13V left P/3L left Pl4W left P/4L left M/ I\7 left M/ I

out and some plates are lost. It has 9-10 plates, all inuse. The platei show a complete enamel loop, exceprthe last two; these are still divided into thiee ringseach. The occlusal surface is narrow. The talon lsdamaged.

The right hemimandible (lnv. N". 291721) lacksthe symphysis and the ascendine rAmus and shows the

rypical fe"iures of Mammuthus''meridionalis. The cor-pus is rather long with a lightly inflated labial side.The angle berween the ascendirg ramus and the cor-pus is quite wide.

VillaSpinola

right

VillaSpinola

left

GLFLGBFBPFQLLDLIGHKE

24522396861 l

)') )/- f- t./-

4,4430,43 ,9

228,'213?8985- 1 022,84,3230,23 ,5

Thb. 3 - Measures (mm) of the molars of Mammuthus meridio-nalis from Villa Spinola: GL = greatest mesio-distallength; FL = occlusal length; GB = greatest bucco-lin-g,r"J breadth; FB = occlusal breadrh; ?p = plate formu-Ia (including talons); QLL = ratio berween^GL and rhe

li t'ff ' d f i''53 kl ; J :,Tf; l" ko r nfi ,Tffi f ilr".1(GH/GB); E = medium thickness of the enamel of rheplate (from Ferretti, 1999; Maccagno, 1962).

The molar placed in the mandible is M2, it israther worn-out, with I I plates. These plates show acomplete enamel loop, except the last rwo ones,which are still divided into rings. The enamel is ratherthick. The molar is narro* "r? concave, its height isreduced.

The metrical and morphological features of thespecimen of Villa Spinola are within the variabiliryrange of M. meridionalis, with great affinities with thespecimens from the Early Pleistocene (lateVillafranchian Farneta F.U.) deposits of Pietrafittaand Scoppito (UAquila, Ab ruzzr) (Maccagno, 1962).They are bigger than the Middle Pleistocene speci-mens from Oriolo (Emilia Romagna) (Ferretti, 1999)and Promano (Umbria) (unpublished data). Theenamel frequency is quite high, similar to those of thePietrafitta, Promano and Oriolo specimens. For thesereasons, the specimen from Villa Spinola can bereferred rc M. meridionalis (advanced Form).

319

CONCLUSIONS

Accordi.g to the description given by Tirccimei(1896), the mammal fauna of Vil la Spinola comesfrom a fluvio-lacustrine deposit correlatable with theS. Maria di Ciciliano Formation (Basilici, 1995).From the south-western branch of the Tiber Riverbasin, Early Pleistocene vertebrate remains have beenrecorded in this formation, both from alluvialdeposits and paleosoils (Sardella et al., 1995; Gentiliet al., 1997; Petronio et al., 2000-2002) and theoccurrence of several late Villafranchian mammalassemblages and their biostratigraphical implicationshave been recently pointed out (Sardella et dl., inpress). Moreover, the well-known mammal assem-blage from Pietrafitta referred to the Farneta F.U.,

EARLY PLEISTOCENE LARGE MA]UIMALS FROM VILLA SPINOLA

Measures

Text-fig. 2 Mammuthus meridionalis Nesti (advanced form)from Vil la Spinola.A) left mandible; B) r ight mandible.Scale bar = l0 cm

VillaSpinola

left

Measures VillaSpinola

right

Length oral border oF the alveolusHeight of the mandible at midpointcheektooth row

266o f 180

255146

Tab. 2 - Measures (--) of the mandible of Mammuthus meri-dionalis from Villa Spinola.

320

comes from a localiry abo w 25 km from Perugia. Thetraces of chemical alterations on the skeletal remainswere probably due to the action of the soil and ofsmall

-plants. In particular, the right hemimandible

and both capiti of the femur belonging_to Pantheragombaszoegensis show rypical traces of roots (seeBehretrr*Eyer & Hill, 19^80). Such kind of evidencesuggests that the fossils came from a paleosoil, whichwas not recognised by Tirccimei or other laterresearchers.

The analysis of the fossil material coming fromVilla Spinola enables to refer it to Early Pleistocene.Unfortunately nothing can be said about the taxo-nomical position of the rhino, equids, suids, cervidsand bovids recorded in the deposit, but the occur-rence of a quite rich mammal assemblage can be tes-tified.

Thking into account the evolutionary degree ofthe southern mammoth and the Eurasi an jaguar themammal assemblage from Villa Spinola can bereferred to the late Villafranchian (probably FarnetaF.U.), confirming the occurrence in the S. Maria diCiciliano Formation of different mammal assem-blages.

ACKNO\TLEDGEMENTS

'We wish to thank Profl P. Ambrosetti (Universiry of Perugia)

and Dr. M. C. De Angelis ("soprintendenza Archeologica perI'Umbria') for allowing Ls to study the fossil material and for

-the

useful suggestions. W. also thank Prof. H. Hemmer and ananonymous referee for the critical revision of the manusctip!,Mrs. H. Terry and Dr. N. Minciotti flor improving the Englishtext. This work is supported by

- MURST grants

(Cofinanziamento I 999).

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