INTRODUCTION

The alluvial deposits of river Po and other Padana Plainrivers (Ticino, Adda, Lambro, ecc.), after major floodevent, yield fossil bones of large Quaternary mammals.Moose remains were recorded in 1822 by Breislak, althou-

gh without distinction between moose and giant deer,and then described in more detail in the famous mono-graph of Cornalia (1858-1871), completed with illustra-ted plates. Other works then followed: Zuffardi (1911)and Sacchi Vialli (1950) on the Pavese alluvions, Patrini

Mem. Sci. Geol. v. 54 pp. 51-63 11 figs 4 tabs Padova 2002– ISSN 0391-8602-

Pleistocene fossil Alcini (Cervidae, Mammalia) from Lombardy and Emilia Romagna (North Italy)

Marzia BREDA

Dipartimento di Geologia, Paleontologia e Geofisica dell’Università di PadovaVia Giotto1, I-35137 PADOVA, ItalyE-mail: marziabreda@hotmail.com

ABSTRACT - Fossil bones of large mammals are frequent in the alluvial deposits of the Po Plain and the moose were known since the beginning of the 19th Century. However a complete and up dated work on the fossil Alcini fromItaly was lacking. The numerous new remains shown in this work lead to better outline the distribution of moose inItaly, that shows a palaeobiogeographical meaning. In fact, the Alcini were limited to the Po Plain and never crossedthe Apennines to reach Tuscany, which, probably, represented a different bioprovince. The Italian moose remainsbelong, almost entirely, to the species Alces alces, now living in the northern European regions. Up to now, the ex-tinct genus Cervalces was signalized only twice in our peninsula: from Crostolo Creek near Reggio Emilia (Ambro-setti and Cremaschi, 1976) and from Fornaci di Ranica near Bergamo (Azzaroli, 1979). The specific identification ofthe Crostolo specimen has been here converted from C. gallicus to C. carnutorum, whether the Ranica remains areconfirmed as C. latifrons. Moreover, new finds of both the species are here described: one C. carnutorum from Leffe(Bergamo) and two C. latifrons respectively from San Cipriano Po (Pavia) and from an unknown locality of the Pa-vese alluvions. The finding of fossil moose alone, cannot lead to a precise palaeoenvironmental reconstruction, howe-ver their presence suggests a cold boreal environments with boggy areas dissected by water courses. In Italy, the fin-ding of fossil moose is an index of a cold episode.

K e y w o r d s : Alces alces, Cervalces latifrons, Cervalces carnutorum, Lombardia, Emilia Romagna, Midd-le Pleistocene-Holocene.

RIASSUNTO - In questo lavoro si analizzano i resti fossili di Alcini quaternari rinvenuti in Lombardia ed Emilia Ro-magna. Nei depositi alluvionali della Pianura Padana è frequente il rinvenimento di ossa fossili di grossi mammiferi,tra cui l’alce, segnalato già dall’inizio dell’Ottocento. I resti del grosso cervide sono stati analizzati in numerosi lavo-ri, per lo più datati, ma manca una trattazione aggiornata ed organica degli Alcini di queste regioni. Le numerose nuo-ve segnalazioni di questo lavoro consentono di delineare meglio la distribuzione paleogeografica dell’alce in Italia,che risulta limitata sostanzialmente alla Pianura Padana. Al di fuori di Lombardia ed Emilia Romagna, qui conside-rate, l’alce si trova, infatti, frequentemente nelle cavità carsiche di Veneto e Friuli Venezia Giulia (Breda, 2001a). Ladistribuzione padana dei reperti italiani würmiani di alce indica che il grosso cervide non è riuscito ad oltrepassarel’Appennino e a giungere in Toscana, sostenendo l’ipotesi della presenza di due diverse bioprovincie, una orientalepannonica ed una occidentale del Mediterraneo europeo. I resti italiani di alce appartengono quasi esclusivamentealla specie Alces alces, attualmente vivente nelle regioni settentrionali dell’Europa. Il genere estinto Cervalces è statofinora segnalato solamente due volte nella nostra penisola: sul torrente Crostolo (Reggio Emilia) da Ambrosetti e Cre-maschi (1976) e a Fornaci di Ranica (Bergamo) da Azzaroli (1979). In questo lavoro si assegna il palco del Crostolo,attribuito in precedenza a C. gallicus, a C. carnutorum, mentre si conferma la determinazione dei resti di Ranica a C.latifrons. Si segnalano inoltre altri resti di entrambe le specie: un frontale di C. carnutorum dal bacino di Leffe (Ber-gamo) e due palchi di caduta di C. latifrons rispettivamente da San Cipriano Po (Pavia) e da un’altra località impre-cisata delle alluvioni pavesi. Il ritrovamento dell’alce fossile non consente di ipotizzare un paleoambiente, in terminidi copertura arborea più o meno sviluppata, ma la sua presenza è indice di ambienti freddi di tipo boreale con corsid’acqua ed aree umide, quindi ritrovato fossile nella nostra penisola, testimonia una fase fredda.

P a r o l e c h i a v e : Alces alces, Cervalces latifrons, Cervalces carnutorum, Lombardia, Emilia Romagna,Pleistocene medio-Olocene.

(1926) on the remains from Arena Po and Agosti andBajetti (1966a; 1966b) on those from Acqualunga.

Since then, A. alces was recorded only in some gra-duation thesis (Dal Sasso, 1993; Magni, 1997; Smerini,2000), but a complete and up dated work, dealing withthe Alcini from Lombardy and Emilia Romagna, was stillmissing. The review of the moose remains from these re-gions, together with the work carried on by Breda (2001a)for North Eastern Italy, will complete the Italian picture.

Most of the Italian Alcini remains belong to the spe-cies A. alces, now living in the European northern re-gions. The genus Cervalces has been recorded in Italy onlytwice, respectively C. gallicus in the Crostolo Creek (Reg-gio Emilia) by Ambrosetti and Cremaschi (1976), and C.latifrons in the Fornaci di Ranica lacustrine basin (Berga-mo) by Azzaroli (1979). The identification by Azzaroli(1979), based on a male braincase and a fragmentary shedantler, is here confirmed. On the contrary, the Crostolomoose, consisting of another fragmentary shed antler, ishere attributed to the species C. carnutorum.

In this work, other two shed antlers of C. latifrons arerecorded: one from San Cipriano Po (Pavia), the otherfrom an unknown place in the Pavia province. Moreover,a right frontal bone of C. carnutorum is recorded from theLeffe basin (Bergamo). These remains, the first twolacking any stratigraphical indications, are however im-portant for the extreme rarity of this Middle Pleistocenespecies in Italy.

MATERIAL AND METHODS

A revision of the moose remains from Lombardy andEmilia Romagna regions, is here presented with descrip-tions, measurements, bibliographical references and cur-rent repositories. For the specimens already exhaustivelyanalysed recently, it will be referred to the respectiveworks for the description.

The moose were recognised among large artiodactyl re-mains, stored in numerous museums of the Po Plain.Although not all the specimens described in literaturewere found, as often their repositories indications weremissing, new remains were discovered from Petosino (So-risole, Bergamo), Leffe (Val Gandino, Bergamo), CavaMerlin (Trezzano sul Naviglio, Milan), San Cipriano Po-Portalbera (Pavia), Isola Serafini (Cremona), Rivalta diRodigo (Mantova) and other places. The moose is so re-corded from 21 localities in Lombardia and Emilia regions(Fig. 1) and in other not well identified places of the “Pa-vese” or “Po alluvions”.

The rich osteological collection of the Natural and Cul-tural Resources Department of Ferrara University wasused for comparison in order to recognize the bone frag-ments. The collection includes a complete skeleton of anadult male, lived in Norway and expressly prepared by theauthor. The remains from the different museums were de-scribed following the Barone (1980) nomenclature andcompared each other and with bones from present-dayanimals, to make considerations on sex, age and size of thespecimens.

The measures of antlers and jaws were gathered in ta-

bles at the end of the work. The measures, all in millime-tres, are those suggested by von den Driesch (1976), ad-ding the length (L) and breadth (B) of each tooth. Agraphical elaboration of metrical data was not possible,because measures were generally not comparable, due tothe bad state of preservation and fragmentariness of thematerial.

Unfortunately, most of the moose remains from Lom-bardy were found in alluvial deposits and lacks any stra-tigraphical information and, sometimes, the locality of di-scovery. Many remains were redeposited further down-stream, after fossilization took place, by periodical waterfloods. Downstream of the confluence between Po-Tici-no (Pavia province) and Po-Adda (Cremona province),for example, many mammal bones are found, probably asa consequence of the reduced velocity of the river current(Dal Sasso, 1993).

DESCRIPTION OF THE MATERIAL

CAVERNA GENEROSA

Caverna Generosa, at 1400 m above sea-level on Mon-te Generoso (S. Fedele d’Intelvi, Como) yielded severalfossil vertebrate remains, collected during a digging cam-paign in 1999 by some researchers of the University of Mi-lan, and studied in two graduation thesis (Smerini, 2000;Bona, 2001). A moose upper maxillary, now stored in theEarth Science Department of the University of Milan asMG 22/30-70, was recognized among the fossil bones. Itconsists only of the alveolar portion of the left uppermaxillary bone, which unfortunately is very fragmentary

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Fig. 1 - Map of the localities yielding fossil moose remains inLombardia and Emilia Romagna regions. (1: Caverna Genero-sa; 2: Trezzano sul Naviglio; 3: Vigevano; 4: Zibido; 5: Villan-terio; 6: Muzza; 7: Mezzana Corti; 8: San Cipriano Po; 9: ArenaPo; 10: Chignolo; 11: Petosino; 12: Ranica; 13: Leffe; 14: Ac-qualunga; 15: Pizzighettone; 16: Rivalta di Rodigo; 17: Isola Se-rafini; 18: Polesine Parmense; 19: Crostolo Creek; 20: Settepo-lesini di Bondeno; 21: Diamantina).

but includes P2-M1. The teeth show a crown, well swollenat its base, typical of Alcini; their advanced wear indica-tes the animal’s old age. The premolar tooth row is 65,5mm long. Measures of the individual teeth are the fol-lowing: LP2: 22,5; LP3: 22,6; LP4: 22,7; LM1: 26,5; BP2:21,8; BP3: 27,2; BP4: 29,8; BM1: 27,3.

TREZZANO SUL NAVIGLIO

A shed moose antler, which has been found in the Ca-va Merlin (Trezzano sul Naviglio, Milan), is now storedin the Civic Natural History Museum of Milan as V.1537.This antler, yellow in colour, shows well developed groo-ves of blood vessels. The palm is parted, by a deep inci-sion, into an anterior and a posterior portion, and breaksoff without letting hypothesise the number of tines.

VIGEVANO

Sacchi-Vialli (1950) describes a braincase with part ofthe antler beams, “proveniente da una cava di sabbia nel-le immediate vicinanze di Vigevano” (Pavia), stored in theCivic Museum of that village. The Museum being now clo-sed, it is not possible to analyse the braincase.

ZIBIDO

A fragmentary shed antler from Zibido (San Giacomo,south-west of Milan) is now exposed in the Civic NaturalHistory Museum of Milan. Most likely it consists of the ri-ght antler that bifurcates, by a large incision, into an an-terior and a posterior part. The anterior part preserves thebase of two tines and a third complete one, all of whichare directed forward. It is not possible to hypothesise thecomplete number of tines of the posterior part. The sur-faces are light in colour and preserve the vascular grooves.

TICINO ALLUVIONS

Cornalia (1858-71) describes a moose antler found inthe Ticino river. The finding, in a bad state of preservation,was given to the Museum of Milan by Count Carlo Her-mes Visconti. Cornalia describes and pictures it (pl. 15, fig.1) emphasizing the partition of palm into an anterior anda posterior portion. Unfortunately, this antler is not iden-tifiable in the Museum’s collections; it could have been de-stroyed in the fire during the Second World War.

VILLANTERIO

Zuffardi (1911) describes a left moose antler found“alla profondità di circa un metro, in mezzo alla sabbia ealle ghiaie del greto del Lambro...in località C. Mon-te...circa 7 km a nord di Villanterio” (Pavia). The authorcalls it Alces palmatus, synonimus of Cervus alces, in or-

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Fig. 3 - Fragmentary shed antler of A. alces from Zibido (Milan).Scale 15 cm.

Fig. 2 - Left upper maxilla of A. alces with P2-M1 from CavernaGenerosa (Como); A) occlusal view, B) lingual view, C) labialview. Scale 10 cm.

der to highlight “la presenza di una forte intaccatura,pressappoco in corrispondenza del tronco del corno, laquale viene a dividere tutta la palma in due parti: l’una an-teriore più piccola, l’altra posteriore molto grande”. Thespecimen is an almost complete shed antler, with wellpreserved surfaces and evident blood grooves, extendingfrom the beam to the entire palm. The anterior portionconsists of three long tines directed forward; the poste-rior portion is caudally elongated and is slightly concavedupward. The large posterior palmate portion has sevencomplete tines, except for the fifth that preserves only thebase. The specimen is now stored in the Geological Mu-seum of the University of Pavia as P-1. Its large size andelevated tine number let hypothesise that it belongs to astrong adult male.

From the same place, Sacchi-Vialli (1950) describesanother left antler fragment “costituito dalla pertica conrosetta in parte usurata, e dalla porzione prossimale del-la palma...il margine distale è completamente assente efratturato al di sotto di quella che doveva essere l’incisu-ra tra parte anteriore e parte posteriore della palma”. Thisspecimen is now not identifiable because Sacchi-Viallidoesn’t record its repositories.

MUZZA, LODI

Cornalia (1858-71) describes a shed antler from Muz-za (Lodi, Milan), already described by Meyer (1832) andthen kept in Frankfurt. The specimen was brought by M.Rüppel during his stay in Pavia in 1918-21 and was thendescribed by Breislak (1822), together with some giantdeer remains (the author didn’t distinguish between thetwo species). It is a right shed antler with a much conca-ved palm. The first five tines are complete with exceptionof the second one, consisting only of its base; on the otherhand, the posterior part of the palm breaks off without let-ting hypothesise the number of tines. Cornalia attributesthis antler to an old animal and pictures it in plate 15 (figs2 and 3).

MEZZANA CORTI

A braincase of an adult male, with part of the antlers,which is now stored in the private collection of Arch.Chiolini (Travaccò Siccomario, Pavia), comes from Mez-zana Corti (Pavia). The very well preserved skull has theoccipital, temporal, parietal and frontal bones, the lastones of which are interrupted on the orbit edge. The an-tlers, almost complete, are horizontally set on the skulland perpendicular to the sagittal plane. For a detailed de-scription, with pictures and measures, see Breda et al.(2002).

SAN CIPRIANO PO

An adult male braincase, now stored in the Archaeo-logical-Palaeontological Museum of Casteggio (Pavia),was found at San Cipriano Po (Pavia). The specimen isbadly preserved with much abraded surfaces. It consistsof the occipital, temporal, parietal and frontal bones, thelast ones broken off on the supraorbital foramens. These

antlers retain only the base and are also much abraded,with burrs already indistinguishable. The small size of thebraincase and the badly joined cranial sutures led to thehypothesis that it belonged to a not completely grown in-dividual. A careful description, with pictures and measu-res, is in Breda et al. (2002).

At San Cipriano Po was found a left shed antler too, nowstored in the Archaeological-Palaeontological Museum ofCasteggio (Pavia) as V.424. This antler is very incompleteand has badly preserved surfaces: the burr is quite com-pletely abraded; and, the beam breaks off straight after theopening of the palm. The short diameter of both, beam andburr, suggests that it belonged to a young male.

At last, in the Museum of Milan is stored, without ca-talogue number, an antler beam coming from San Ci-priano, too. The label identifies it as “Megaloceros eury-ceros italiae, dono Ferraresi”. It is a long shed antler beam,with dark smooth surfaces, without any tine. The burr hasan ellipsoidal outline and was less developed on the lowersurface. The beam is slightly curved and begins to flattendistally on a plane perpendicular to the major diameter ofthe burr. The beam breaks off, without opening in thepalm, at 373 mm from the shedding surface. Hypothesi-sing that the longer diameter of the shedding surface is theanterior-posterior one, and that the bending of the beamis directed forward, the antler would be the left one (seeBreda, 2001b). The lack of tines on the burr leads to ex-clude the attribution to a Megacerine; moreover, thelength of the beam is typical of the genus Cervalces. Thestrength of the beam suggests to identify it as C. latifronsfrom the Middle Pleistocene, and the big weight, due tothe heavy mineralization, is a confirmation of old age.

ARENA PO

Zuffardi (1911) describes an antler “tratto dal notogiacimento di Arena Po...riferibile ancora all’Alces pal-matus rappresentato da un semplice lembo di palma conquattro pugnali”. The author suggests that the specimen,now stored in the Geological Museum of the Universityof Pavia as P-17, is the anterior part of the antler of an oldindividual. The palmate portion, slightly concaved, is con-stituted by two almost parallel edges that continue in twotines, which are slightly outward curved. Between thesetwo tines, there are two shorter ones which are parallel tothe edges of the palm. This morphology reminds, rather,the extremity of the posterior palm, and is particularly si-

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Fig. 4 - Shed antler beam of C. latifrons from San Cipriano Po(Pavia). Scale 10 cm.

milar to the posterior portion of P-1, already described byZuffardi.

Another antler fragment from Arena Po (Pavia) is sto-red in the Geological Museum of the University of Paviaas A1I-1. It is a left shed antler, well mineralized and darkin colour, with smooth surfaces, that can be attributed toa relatively young animal, mainly because of the shortbeam diameter. The burr is much abraded and the beamopens gradually in the palm but breaks off before reachingthe base of the tines.

A left mandibular ramus with P4 has been found inArena too, and is now stored in the Natural Sciences Mu-seum of Voghera (Pavia), as 225. This specimen, lightbrown in colour, extends from the mandibular symphy-sis to the M1 alveolus, measuring 230 mm. It keeps onlyP4, which is very worn but complete.

Patrini (1926) describes even a “porzione basale di cor-no sinistro col rispettivo pugnale” and another “fram-mento di corno” from Arena Po, both attributed to themoose. These remains, then stored in the Museum of Mi-lan, were not identified in this research as they might ha-ve been destroyed during the Second World War bom-bing.

CHIGNOLO PO

From Chignolo Po (Pavia) comes a left mandibular ra-mus with P2-M3, now in the exhibitions of the Civic Na-tural History Museum of Milan, catalogued as V.1040.The jaw, brown in colour and well mineralized, lacks inthe ascending ramus and, partly, in the mandibularsymphysis. The maximum length of the piece is 282 mm,its maximum height (perpendicular to the length) is 70mm. All the check teeth are very well preserved and arelittle worn, which suggests an animal of young age (DalSasso, 1990). The transverse section of the mandibular ra-mus is well flattened, the swollen base of the teeth’scrowns and the hypoconide of P4 isolated in the labialwall, all confirm the attribution to the moose. The threemolars have a small ectostilyd on the labial wall betweenthe two lobes.

PAVESE ALLUVIONS

Numerous remains don’t possess the provenience lo-cality, but only a generic indication of where they werefound “Alluvioni Pavesi”:

- Sacchi-Vialli (1950) analyses a very well preservedbraincase of a moose, with part of the antlers (pl. 2, fig.1), now located in the collections of the classical lycée“Grattoni” in Voghera (Pavia). It preserves occipital, tem-poral, parietal and frontal bones with pedicles, broken offon the ethmoid meatus, just before the supraorbital fora-mens. The skull, recently reanalysed by Breda et al.(2002), belonged to a relatively young animal, because ofthe state of cranial sutures.

- Sacchi-Vialli (1950) describes also a right mandibu-lar ramus, extending from the mental foramen to the thirdmolar, with P4-M3. The author identifies it as A. alces dueto the flattened transverse section of the ramus and thetypical morphology of P4 with hypoconid isolated fromthe labial wall (pl. 1, fig. 5). The three molars are presentbut heavily damaged. Sacchi-Vialli states neither the lo-cation of collection, nor the place of repository.

- Shed antler stored in the Naturalistic Civic Museum“F. Lombardi” of Stradella (Pavia) under the cataloguenumber F.1179. The specimen, white in colour, has veryflaky and “decayed” surfaces with red clay sediment. Theburr is fairly well preserved and the beam has the bloodygrooves hardly visible. It should be a right antler, in factthere are two tines that seem to constitute the anteriorportion.

- Shed antler beam stored in the Naturalistic Civic Mu-seum “F. Lombardi” of Stradella (Pavia) under the cata-logue number F.1180. The sample is well fossilized butvery fragmentary, breaking off at the beginning of palm.The burr is completely abraded and the shedding surfa-ce is slightly oblique. The very small size testifies theyoung age of the animal.

- Right mandibular ramus stored in the NaturalisticCivic Museum “F. Lombardi” of Stradella (Pavia) underthe catalogue number F.1344. This specimen, well mine-ralized and dark brown in colour, extends from the cau-dal margin of the diastema to the third molar alveolus, fora full length of 245 mm. The following pieces are still pre-sent: parts of the P2 and P3 roots, a fragmentary P4 crown,a very small part of the M1 root and the whole M2 crown.The bony portion is broken between P3 and P4. The midd-le wear of M2 suggests an animal in adult age.

- Shed antler beam, very heavy, with a smooth and darksurface, now stored in the Natural Science Museum of Vo-ghera under the catalogue number 342. The beam is verylong and typical of the Alcini tribe because it lacks in ti-nes and signs of bifurcation. The shedding surface is qui-te circular and the beam becomes elliptical distally and itbreaks off before opening in to the palm at 300 mm fromthe shedding surface. The extreme length of this beam,

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Fig. 5 - Left jaw of A. alces with P2-M3 from Chignolo Po (Mi-lan); lingual view. Scale 10 cm.

Fig. 6 - Shed antler beam of C. latifrons from the Pavese allu-vions. Scale 10 cm.

excludes definitely its identification as A. alces. This an-tler is here attributed, because of morphology and di-mensions, to the species Cervalces latifrons of the Midd-le Pleistocene. The advanced mineralization of the speci-men suggests old age, in agreement with the systematicalconclusions.

PETOSINO

In Petosino (Sorisole), in the north-western outskirtsof Bergamo, is a gravely-sandy and clayey succession of la-custrine origin. These deposits have been exploited sin-ce the end of the 19th Century and the diggings broughtto light mammal fossil bones and human artefacts (Caffi,1938). Caffi (1930) signalises rhinoceros, bison and mam-moth remains. In the Civic Natural Science Museum “E.Caffi” of Bergamo are stored, under the catalogue num-ber 1422, some moose teeth, donated by eng. Sassi in1905. They consist of the upper teeth of an individual,free, with blackish crowns and reddish roots. There arethree fragments of a not identifiable molar (M1 or M2), allthe premolars and the two M3, all in a different state ofpreservation. The advanced wear testifies the old age ofthe animal. It was not possible to measure the premolartooth row due to incompleteness of the single teeth. Themeasures of the single teeth are the following: (right teeth)LP2: 24,1; LP3: 23,1; LP4: 22,9; BP2: 22,2; BM3: 31,1; (leftteeth) LP2: 24,6; LP3: 23,6; LP4: 22,9; LM3: 30,7; BP3:28,3; BP4: 29,1.

FORNACI DI RANICA

From the lacustrine basin of Ranica (Upper SerianaValley, Bergamo), two pieces of a large deer, which arenow stored in the Civic Natural Science Museum “E. Caf-fi” of Bergamo, were picked up from the clay, rich in fre-shwater shellfish and vegetal remains. They consist of abraincase and a shed antler beam which were identifiedand described by Azzaroli (1979) as the first, and up totoday the only, find of Cervalces latifrons in Italy.

The antler (catalogue number 1421) was found by Mr.Gavezzeni in 1929, but its exact location was not recor-ded (Azzaroli, 1979). The specimen shows the sheddingsurface, elliptical in section, the burr and the beam. Thelast one begins to flatten in a plane perpendicular to themajor diameter of the burr and breaks off at the begin-ning of the palm (the full length of the piece is 285 mm).This antler can easily be attributed to the Alcini tribe be-cause of the lack of tines on the beam. The beam’s lengthjustifies the identification as C. latifrons.

The braincase (catalogue number 2609) was found byMr. Milesi and Mr. Mora at the end of the 60’s in blue clayof freshwater origin, at about a depth of 10 meters. Theskull belonged to a strong adult male and consists only ofthe neurocranial portion with occipital, temporal, parie-tal and frontal bones, including pedicles. The facial por-tion, useful for the distinction between Alces and Cerval-ces genera, is lacking. But the occipital bone is massive andshows a broad and shallow outline, typical of Cervalces,in comparison to the narrow and deep occipital of Alces(Azzaroli, 1952; 1982). This broad occipital is necessary

in Cervalces to give insertion to the strong neck musclesthat need to support a heavier head because of the longerantlers in comparison to Alces. The large size of the spe-cimen together with the morphology of condyles and fo-ramen magnum (Breda, 2001c), support the specific iden-tification by Azzaroli (1979). For measures see Azzaroli(1979).

Both the above mentioned moose remains are lackingin a stratigraphical location. From a palynological analy-sis of two clay samples taken from the skull, and by com-paring these with the pollen diagram of the whole Rani-ca section, it was possible to infer the skull’s strati-graphical location. In fact, there is a strong similaritybetween the skull samples and the pollen zone RNC3,representing the basal marshy facies of the lacustrine se-quence of Ranica (Ravazzi et al., in press; Breda et al.,in progress). Infrared spectra were performed on the sa-me clay samples taken from the skull, supporting the cor-relation to the lower part of the Ranica section, becau-se of the high percentage of silicates and the scanty car-bonates (Pini et al., 2002). The deposits with normal po-larity magnetization are correlated to the Jaramillo sub-chron or to the Cobb cryptochron (Ravazzi et al., inpress).

LEFFE BASIN

The Leffe Basin (Bergamo) is located at 290-600 m al-titude, in the small Gandino Valley, tributary of the Se-rio Valley, and surrounded by high mountains. It consi-sts of a lacustrine sedimentary assemblage, 60m thick, al-ternated with three organic beds (Ravazzi and Mosca-riello, 1998), which yielded mammal remains since the se-cond-half of the 19th Century.

The Leffe mammals are assigned to two levels correla-ted respectively to the Farneta (upper level) and TassoFaunal Units (lower level) (Ravazzi, 1993). During a re-cent revision of this mammal fauna, now in progress bythe author, a moose right frontal bone was recognised inthe collection of the Civic Natural History Museum of Mi-lan; it still lacks a catalogue number. According to the oldlabel, the specimen was collected in the upper brown-coalbank, near the Re creek in the 50’s. It is presumed to be-long to an adult male. The specimen is well mineralized

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Fig. 7 - Braincase of C. latifrons from Ranica (Bergamo); occi-pital view. Scale 5 cm.

and shows the surface suture to the other frontal and tothe parietal bone; the endocranial cast of the brain is wellevident in the lower surface. The frontal is incomplete,lacking in the foremost portion (it breaks off in corre-spondence of the orbit) and in the pedicle extremity (theshedding surface or the burr). The pedicle is set horizon-tally and perpendicular to the sagittal body plane, typicalof the Alcini; it is elliptical in section (being compressedin a dorso-ventral plane), typical of the genus Cervalces incomparison to Alces (the diameters are 77 x 56 mm). Thesize of this frontal suggests the belonging to the speciesC. carnutorum; the presence, in the same brown-coalbank, of Archidiskodon meridionalis vestinus and Stepha-norhinus etruscus evolved form (Guerin, 1980), confirmsthat the bank is Late Villafranchian (Marchetti and Bre-da, in progress). Palynological analysis, of the sedimentpreserved in the bone, is in progress, in order to confirmthe stratigraphical position of the sample in the Leffe se-quence, by correlating it to the eleven vegetational and cli-matic cycles recognized in the complete pollen diagram(Pini and Ravazzi, 2002).

ACQUALUNGA

Agosti and Bajetti (1966a; 1966b) attribute two tibias,a radius, a pelvis and a fragmentary right antler to A. al-ces. The bones were deposited after a flood, together withsome elephant remains, on the sandy banks of the Oglioriver, at Acqualunga (Borgo San Giacomo, Brescia). As amatter of fact, only the antler identification is correct, asthe radius belongs to a megacerine, whereas the tibiasand the pelvis to a bovid. The antler, now stored in theCivic Natural History Museum of Brescia in the box 104,has the burr fully abraded and it is broken off a little af-ter the beginning of palm. The poor mineralization is re-sponsible for the flaky surfaces.

PO PLAIN ALLUVIONS

Numerous remains don’t possess the place of origin,but only the generic indication of where they were found“Alluvioni Padane”:

- Antler beam now stored in the Civic Natural HistoryMuseum of Milan under the catalogue number V.3482.The specimen is very badly preserved and light, with rou-gh surfaces without bloody grooves, because of the poormineralization. This antler beam, already described byMagni (1997), is too fragmentary to state if it is the rightor the left one.

- Braincase of an adult male, with both the antlers, nowstored in the Museum of Palazzo d’Arco, in Mantova.This skull, the first and one of the finest moose skullsfound in Italy, was described by Cornalia (1858-71), whopictured it in pls 16 and 17 (figs 1 and 2), and has recen-tly been reviewed by Breda et al. (2002). This specimenpreserves occipital, temporal, parietal and frontal boneswith almost complete antlers.

- Shed antler beam stored in the Civic Museum of Cre-ma and Cremasco (Cremona) under the catalogue num-ber st90968. The sample, with porous and brown surfa-ces, has a much abraded burr and breaks off at the be-

ginning of palm. It belonged to a young animal, giving thesmall diameters of burr and beam.

PIZZIGHETTONE

In 1962, at Cava nell’Adda, Pizzighettone (Cremona) wasfound a right shed antler, now in the exhibitions of theCivic Natural History Museum of Milan, under the cata-logue number V.352. The specimen, substantially com-plete, is light, with a porous grey and dark spotted surfa-ce. The burr tubercles are well developed, as well as thebloody grooves. The partition of the palm in both ante-rior and posterior portion is little evident with the forthtine in the middle of the incision. The posterior portionis backward lengthened. There are nine tines, all completeexcept for the sixth which keeps only the base. The largesize of the burr and the beam and the high number of ti-nes, justify the attribution to an old animal (Dal Sasso,1990).

RIVALTA DI RODIGO

Giacometti (1880) refers to a digging campaign carriedout at Rivalta di Rodigo (Mantova), at the end of the 19th

Century. Among the mammal bones, the author didn’t re-cognize a right A. alces jaw, now kept in the storehouseof the “Palazzo Ducale”, in Mantova. This jaw is broken

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Fig. 8 - Right shed antler of A. alces from Pizzighettone (Cre-mona). Scale 20 cm.

Fig. 9 - Right jaw of A. alces with P3-M3 from Rivalta di Rodigo(Mantova); lingual view. Scale 10 cm.

between M1 and M2 (the distal portion is catalogued as7051, the proximal one as 7054). The specimen lacks in:the distal part of the diastema starting from about themandibular symphisis, P2, and the upper part of the man-dibular angle. The mandibular condylus and the coro-noid process are present even if splintered at the edges.The well preserved teeth and the completely grown M3,both suggest an adult age.

ISOLA SERAFINI

A left shed antler, now in the Civic Natural HistoryMuseum of Milan under the catalogue number V.1532,comes from Isola Serafini (Cremona), at the Po-Addaconfluence. The antler surface is smooth and dark withsome bloody grooves on the beam. The burr is well pre-served, with developed tubercles on the superior edge.The specimen breaks off at the beginning of the palm.

POLESINE PARMENSE

A left shed antler from Polesine Parmense (Parma), isnow stored in the Civic Natural History Museum of Mi-lan under the catalogue number V.3481. This antler, de-scribed by Magni (1997), shows a dark, flaky surface, co-vering a lighter surface. The burr and the bloody groovesare quite well preserved and the palm is very concavedupward and parted in an anterior and a posterior portionby a deep incision. The anterior part bifurcates into twotines (only the bases are preserved); the posterior partopens in a wide palm that breaks off before the beginningof the tines.

CROSTOLO CREEK

On the top of the Calabrian levels with Arctica islan-dica, the erosion by Crostolo Creek exposed some conti-nental deposits that yielded many remains of Late Villa-franchian mammals. From these levels comes a fragmen-tary antler, identified as Cervalces gallicus by Ambrosettiand Cremaschi (1976), and now stored in the Civic Mu-seum of Reggio Emilia under the catalogue numberCRO3.

This specimen consists of the right shed antler beam,white and very flaky, and consolidated with paraloid. Thebeam is cracked along the bloody grooves, so that it con-tains sand and gravel from the finding place. The burr andthe proximal portion of the beam are elliptical in section(compressed in the dorso-ventral plane), then the beambecomes circular and begins to flatten distally in a plane

perpendicular to the major diameter of the burr, butbreak off before opening into the palm. The beam, about30 mm in length, is much shorter than that of the C. gal-licus holotype from Sénèze (about 60 mm), being of thesame diameter. For this reason, this specimen is here iden-tified as Cervalces carnutorum. The associated fauna, cor-related to the Farneta Faunal Unit because of the presenceof Mammuthus meridionalis (advanced form), Ursus etru-scus and Canis cf. C. etruscus, should better be compara-ble to the other European sites with C. carnutorum (Bre-da, 2001b; Marchetti and Breda, in progress).

SETTEPOLESINI DI BONDENO

At Settepolesini (Bondeno, Ferrara) the sand extrac-tion from a paleoriver bed formed a lake, about 35 hec-tares wide and 20 meters deep. Since 1997, the dredgerhas taken from the bottom of the lake, together with thesand, many bony fragments of large mammals (Galliniand Sala, 2000). Through sedimentological studies of thearea, and radiocarbon dating of some remains, the mam-mal fauna has been attributed to three different ages:Middle Würm, Late Glacial and Roman Time. The firstmammal association testifies, for the first time in Italy, thepresence of the typical “Mammoth-Steppe” (sensu Guth-rie, 1984; 1990); the second mammal association is againa Steppe association but less cold than the first one, withriparian forested areas, dominated by bisons. Numeroussub-fossilized remains of domesticated animals and manare recorded in the forested Holocene levels (Gallini andSala, 2001).

Among the material, now stored in the Palaeontologi-cal Museum of the University of Ferrara, the following A.alces remains are described below together with the cata-logue number:

- second anterior phalanx (47), complete and with wellpreserved surfaces. The measures are the following: Gl-pe: 62,9; Bd: 25,9; Bp: 29,5; SD: 20,5.

- shed antler beam (73), well mineralized and darkbrown in colour, with a much abraded burr. Because ofthe size of beam, the antler should belong to an adult ani-mal and it is thought to be the left one, considering thatthe anterior edge of the beam bends earlier to open in thepalm.

- fragmentary ascending ramus of the right jaw (168).The piece is brown, badly mineralized and it is composedby the articular condylus (27,5 mm wide) and part of thecoronoid process for a total length of 59 mm.

- shed antler beam (194). The burr is fully abraded andthe surfaces are poorly preserved and flaky. The sheddingsurface is oblique in relation to the beam’s axis, and smallin diameter; therefore, it should belong to a young animal.The antler breaks off at the beginning of the palm, but theanterior edge of the beam bends earlier, suggesting it isthe left antler.

- left shed antler (300). The strong beam testifies theold age of the animal. The burr and the surface are wellkept with evident bloody grooves. A good part of thepalm is preserved, with only one long tine, of which thetip is missing, laterally directed on about the same axis of

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Fig. 10 - Right shed antler beam of C. carnutorum from Crosto-lo Creek (Reggio Emilia). Scale 10 cm.

the beam. The posterior part of the palm should havebeen larger than the anterior one.

- right antler with part of the frontal bone (325). Thefrontal bone is upward concaved; it lacks in the anteriorportion and preserves the suture surface to the left fron-tal and to the right parietal bone. The endocranial cast ofthe brain is evident on the lower surface. The burr is welldeveloped, and a good part of the palm is preserved butall the tines are missing. Only the “saddle”, between an-terior and posterior part of the palm, is present.

- distal portion of right humerus (405). This piece cer-tainly belongs to a moose because of the trochlea morpho-logy (Pfeiffer, 1999): the proximo-ventral edge is hori-zontally set and the articular relief (to the groove in theproximal epiphysis of the radius) is well rounded. Typi-cal of moose, in comparison to the giant deer, is also themorphology of the lateral edge of the distal epiphysis: ina dorsal view, it shows a long “pinchement” and it opensgently in lateral direction, while it bends abruptly upwardin Megaloceros (Breda, 2001b; in progress). It was possi-ble to take only the following partial measures: Bd: 78; BT:64.

- atlas (414). The lips of the transverse processes areabraded, the caudal portion of the vertebral arch and thetip of the ventral tubercle are missing. The belonging toM. giganteus can be excluded as it shows a more cranio-caudally elongated vertebral body and more divergentand caudally elongated lips of the transverse processes(Breda, 2001b). GB: 149; GL: 108; BFcr: 97; BFcd: 92;GLF: 83.

The age of the bones from Settepolesini can be infer-red by their degree of mineralization, which is more pro-nounced in older levels. So, the moose bones 47, 73, 300and 325 should come from the horizons of the MiddleWürm, whilst the others could be related to the Late Gla-cial period.

DIAMANTINA

In 1933, during a drilling carried out by AGIP, at Dia-mantina, on the right bank of the Po river, at about 10 kmwest/north-western from Ferrara, was found a right A. al-

ces mandibular ramus. Maxia (1955) describes and pic-tures this jaw (pls 1-4), now in the Palaeontological Mu-seum of the University “La Sapienza”, in Rome. The bonyportion is poorly mineralized and yellow-brown in co-lour, and the teeth are blackish. The piece extends, for amaximum length of 252 mm, from the anterior edge of themental foramen to the second molar, lacking in all theascending ramus. There are all the check teeth except thethird one, and their low-medium grade of wear testifiesthey belonged to a young/adult animal (Maxia, 1955).

PALAEOGEOGRAPHICAL, CHRONOLOGICALAND PALAEOENVIRONMENTAL REMARKS

The numerous new identifications allow to better ou-tline the palaeogeographical distribution of the fossilmoose in Italy, which are limited to the Po Plain. Themoose were frequently found, besides in Lombardia andEmilia Romagna regions here considered, in the karst ca-vities of Veneto and Friuli Venezia Giulia regions (Bre-da, 2001a).

In Veneto, the first record of moose remains date be-ginning of last Century: Fabiani (1903; 1919), Dal Piaz(1922) and Battaglia (1924) described moose fragmentrespectively from Grotta di San Bernardino (Berici Hil-ls), the lacustrine clays of Borghel (Vittorio Veneto) andGrotta dell’Alce (Trieste karst). Since then, numerousother deposits yielded Alces remains, all analysed by Bre-da (2001a) and Breda and Gallini (2001).

There are no moose descriptions from Piemonte (Salaand Aimar, 1998) but it is not surprising, in fact, sedi-ments filling karst cavities in this region were scarcelystudied, and the flood plains are very small. However bi-son, mammoth and giant deer remains were found in themiddle-late Quaternary deposits of the Piemonte plain(Sala and Aimar, 1998). The lack of moose remains canbe link to their general rarity and to their living habits.Differently from other large ungulates, the moose is nota gregarious species, moving as single individuals. Mo-reover, the remains of animals lived in the Piemonte re-gion, could have been transported by river streams anddeposited in Lombardy, where the stream energy de-creased (Breda, 2001b).

The only exception to the Po Plain distribution of themoose in Italy, are some remains from Arene Candide(Cardini, 1980; Cassoli and Tagliacozzo, 1995) and BalziRossi caves (Boule, 1910), at the border between Liguriaand France. Likely, the moose arrived in Liguria, as in ca-ves of the southern French coast, from the Rhône valleyrather than from the Po Plain after having crossed theApennine (Breda, 2001b). Note however that the twoabove cited localities are prehistorical sites and remainscould have been subjected to mankind reworking andtransport.

A. alces reached the Italian territory during the LateGlacial, from the ex Yugoslavia, where it was widely wi-despread (Malez, 1986). The large deer crossed the Trie-ste karst plateau and the present northern Adriatic Sea,at that time above sea-level. From its first entrance in

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Fig. 11 - Fragmentary right antler with frontal bone of A. alcesfrom Settepolesini di Bondeno (Ferrara) 325. Scale 10 cm.

Italy, A. alces rapidly widespread throughout the Po Plain,where it settled uninterruptedly till its departure that oc-curred with the Holocene climatic warming. The Po Plain,closed to the north by the high Alps and to the south-we-st by the Apennines, prevented the latitudinal migrationsof some taxa, which, on the contrary, were possible in cen-tral Europe as a consequence of the continuous climaticchanges.

The distribution in the Padana Plain of the Italian“Würm” A. alces shows that the large cervid was not ableto overstep the Apennines to reach Tuscany, supportingthe idea of the existence of two separate paleobioprovin-ces: an Eastern or Pannonic province, that comprehendedalso the Po Plain, and a Western or European Mediter-ranean province. As for the moose, other cold species, ar-rived in Italy during the “Würm” Glacial along the sameeastern route, and then remained confined within the PoPlain (e. Ochotona pusilla and Sicista betulina). The pre-sence of a boundary between the two bioprovinces is te-stified, during the Late Pliocene, by the rich mammal as-semblage of Rivoli Veronese (Sala et al., 1994). During theEarly Pleistocene the two provinces were, as suggested bythe exceptionally high number of species of Monte la Me-sa deposits (Marchetti et al., 2000), slightly superposed atleast on the eastern Po Plain. Future moose discoveriesshould be more probable along the Adriatic coast, thanalong the Tyrrhenian coast.

While during the Late Pleistocene A. alces was wide-spread throughout the Po Plain, during the Holocene itwas probably confined only in the mountains, to takeshelter from climatic warming (Breda, 2001b). The morerecent specimen among the Italian A. alces is a telemeta-carpal bone, part of a burial of a Mesolithic grave at Mon-deval de Sora (Dolomites) at 2150 m above sea-level(Guerreschi, 1990), and dated, through radiocarbon,7140 y B.P. (Fontana F., pers. comm.).

Regarding Cervalces, there are two C. carnutorum re-mains, respectively from the Crostolo Creek and from theLeffe basin, both correlated to the Farneta Faunal Unit(sensu Gliozzi et al., 1997), and three C. latifrons. The C.latifrons skull from Ranica is correlated, through palaeo-magnetic and palynological data, to the subchrone Jara-millo or to the cryptochrone Cobb, corresponding to theColle Curti Faunal Unit (Ravazzi et al., in press). Proba-bly it represents the oldest finding of this species in We-stern Europe. The other two C. latifrons remains, from thePavese alluvions, cannot be dated, in any way, and theyare generically referred to the Middle Pleistocene.

The extreme rarity of the genus Cervalces in Italy is pro-bably linked to lack of pre-“Würm” sediments in the PoPlain, generally removed or covered by more recent de-posits. The San Cipriano Po area, where the two C. lati-frons shed antlers were here described, is relatively clo-sed to the piedmont area of the Apennines, and probablysheltered old alluvial deposits. The old age of the twoshed antlers from the Pavese alluvions, is supported bytheir high degree of mineralization, not comparable withthat present on other more recent Pleistocene remains.

Moose remains, although rare in comparison with tho-se of other Quaternary ungulates, have a great palaeo-biogeographic and palaeoenvironmental value.

Alcini are good indicators of marshy and riparian areas,although they can live in different vegetational zones. Forexample, A. alces was present at Settepolesini di Bonde-no (Ferrara) in the Middle Würm forest-steppe (Galliniand Sala, 2001) but lived even in the subsequent LateGlacial steppe, dominated by bison. Also Cervalces wasfound both in the forest faunal associations, e.g. Mauer(Soergel, 1914) and Bilshausen (Schmidt, 1934) and in thesteppe associations, e.g. Süβenborn (Kahlke, 1969). Infact, the moose, moving across vegetational “islands”,may penetrate in the tundra or steppe environments, bea-ring a wide range of temperature. Their distribution is notlimited to the taiga, and they can be found in the shrubforests along flood valleys. These valley biotopes survivedduring the abrupt Pleistocene climatic variations, whenthe forest degraded to a tundra-steppe association and vi-ce versa. This may explain why the Alcini morphology re-mained unvaried during the all Quaternary, besides thestrong climatic fluctuations (Sher, 1987).

This may explain why the moose is the only large mam-mal herbivorous that survived, at the end of the Pleisto-cene, to the arrival of the late glacial steppe that broughtto extinction mammoths and woolly rhinos. The small ri-parian forests, could not sustain large herds, but they we-re large enough, at the end of the Pleistocene, to assuremoose survival, why they lived as isolated individuals oras small temporary groups.

Presence of fossil moose indicates cold boreal envi-ronments with watercourses and boggy areas. The moo-se, today living in the northern European regions, foundas fossil remains in our peninsula, testify a sure cold pe-riod.

ACKNOWLEDGEMENTS

For giving me the opportunity to study the material I wish tothank: Dr. Marco Tonon (Civic Museum of Natural History ofBrescia), Dr. Mrs. Anna Paganoni (Civic Museum of Natural Hi-story “E. Caffi” of Bergamo), Dr. Miss Emanuela Zanalda (Uni-versity of Milan), Dr. Cristiano Dal Sasso (Civic Natural HistoryMuseum of Milan), Dr. Giuseppe Santi (University of Pavia), Dr.Carlo Cova (Civic Naturalistic Museum “F. Lombardi” of Stra-della, PV), Dr. Ferruccio Stella (Archaeological and Palaeonto-logical Museum of Casteggio, PV), Dr. Giuseppe Orlandi (Na-tural Science Museum of Voghera, PV), Dr. Mrs. Cinzia Galli(Natural History Museum of Cremona), Dr. Roberto Martinel-li (Civic Museum of Crema and Cremasco, CR), Dr. Mrs. SilviaChicchi (Civic Museum of Natural History of Reggio Emilia),Dr. Mrs. E. Starnini and Dr. Mrs. E.M. Menotti (Archaeologi-cal Superintendence of Lombardy - Mantova) and the Directorof the Department of Natural and Cultural Resources of Ferra-ra University.

Special thanks are due to Dr. Miss. Donatella Mellere (Uni-versity of Padova) and to Miss. Monica De Nardi (Montebellu-na) who helped me improving the English form. At last, I wishto thank Prof. Benedetto Sala (University of Ferrara) for the use-ful discussions about Alcinae and for having partly financedthis research (ex 40% funds Prof. Sala holder).

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61

Tab. 1 - Mandibular measures in millimetres: 3-15 after von den Driesch (1976); 16: breadth of the mandibular ramusrespectively to the height (15); HD: height of the diastema; BD: bredth of the diastema [Arena Po (Voghera Museum225); Chignolo Po (Milan Museum V.1040); Pavese Alluvions (Stradella Museum F.1334); Rivalta di Rodigo (PalazzoDucale, Mantova 7051 and 7054); Diamantina (after Maxia, 1955; University “La Sapienza”, Rome)]

Tab. 2 - Lower teeth measures in millimetres: tooth length (L) and tooth breadth (B) followed by the tooth name (cur-rent repositories as in Tab. 1)

Tab. 4 - Measures of the Cervalces antler beams in millimetres

Tab. 3 - Measures of Alces alces antlers in millimetres: 39-41 after von den Driesch (1976) with the respective antero-posterior (a-p) and dorso-ventral (d-v) diameters

APPENDIX

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Issued, November 2002