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ARTICLE IN PRESS
M.R. Palombo, M.P. Ferretti / Quaternary International 126–128 (2005) 107–136 119

primigenius of which it represents one of the earliest The Italian M. primigenius sample (Table 6)is
occurrences in Italy (see below). The remainder of the characterized by M3–m3 with a number of plates
material (Campo Verde and Capri) is represented by ranging from 20 to 24 (excluding talons), and thus does
molars whose characters fall in the overlap zone between not include specimens showing the extreme values (27–
M. trogontherii and M. primigenius and could then 28) recorded in central European and northern Asian
belong to either of these two species. populations. Mean values for enamel thickness of M3–
In conclusion, with the exception of the Tarquinia m3 is around 2 mm, while lamellar frequency is greater
skull, it is not possible, in our view, to provide a precise than 7 (modal values 8–9). A mandible from the Adda
systematic placement for these late Middle Pleistocene River near Cremona (Late Pleistocene; Caretto and
mammoths from the present material. Indeed, more Giacobini, 1983; MCSNM) shows a slender but
specimens are needed to solve this issue. relatively long symphyseal rostrum (Fig. 10), indicating
persistence of primitive morphologies among the Italian
2.1.5. M: primigenius (Blumenbach, 1799) woolly mammoth population.
Garutt et al. (1990) proposed as neotype of M.
primigenius a complete skeleton from the latest Pleisto- 2.1.6. Elephas ðPalaeoloxodonÞ antiquus Falconer and
cene of Siberia. Late woolly mammoth populations are Cautley, 1847
characterized by a smaller size with respect to older The type specimen is an incomplete mandible from
Mammuthus species and/or samples (Maschenko, 2002; England. The exact locality is unknown. However, from
Lister, 1996a; Kulczycki, 1955). M. primigenius is its preservation the mandible might have been retrived
characterized by its strongly curved tusks, M3–m3 with from terrace deposits of the Thames River, at Grays, a
(19) 20–27 plates (average 24), and an average lamellar locality correlated with the late Middle Pleistocene (Paul
frequency of 8. Enamel thickness in M3–m3 is usually Davies and Adrian Lister, personal communication,
less than 2.3–2.5 mm. 2003). Knowledge of the skeletal and dental variability
In Italy, abundant, though isolated remains (mostly of this species is however based on the more abundant
representing molars and post-cranials) of M. primigenius material from continental Europe, especially from
are known from Bucine (Arezzo; De Giuli, 1983; Mazza, Germany and Italy. The skull is characterized by a flat
1998; IGF, NHMB, MPF), Arezzo (various sites; dorsal profile, double-domed vertex, and specialized
Ferretti, 2000; IGF, MAA), and the Po Valley (various anatomy of the forehead, which is characterized by the
sites, especially between Pavia and Cremona; Airaghi, occurrence of a strong parieto-frontal ridge (but see
1917; Anelli, 1947; Sacchi Vialli and Pizzochero, 1958; below for intraspecific variability of this feature). The
Sala, 1986; Reggiani, 1995; MDSTP, MCSNM). Un- premaxillaries are fan shaped and extremely wide
fortunately, during the Second World War many distally. The jugal arch is relatively slender. The
specimens from the Po valley, housed at MCSNM, mandible is short with a weakly developed symphyseal
and from Arezzo, housed at MAA, were destroyed. beak. Tusks are large, and weakly curved. The molars
Among the latter material, also an incomplete skull went are hypsodont (HI 1.8–2.0 in M3), with a relatively
lost (Ricci, 1901a; Ferretti, 2000). Juvenile woolly narrow crown, characteristically wrinkled enamel with a
mammoth material was retrieved from a cave deposit mean thickness of 2.6–2.7 mm in M3–m3. M3 possess an
at Podere Cardamone, Lecce (Botti, 1890; Vaufrey, average of 16 plates with a maximum at 19–20. Incipient
1927; Rustioni et al., 2003; MPB; ITCL). Other wear figures on the occlusal surface are formed by a
occurrences of woolly mammoth in Italy are those from large oval central loop and two small lateral loops. At
Torrente Conca (Cattolica; Ferretti, 1997; MTR), Asolo an intermediate wear stage, complete enamel loops on
(Treviso; Reggiani and Sala, 1992; MCSNV), Riparo the occlusal surface are usually either of the convex–
Tagliente (Verona; Reggiani and Sala, 1992; MCSNV), convex or convex–concave type (according to the
Montecatini Terme (Lucca; Ricci, 1901b; IGF), Marche terminology of Kalb and Mebrate, 1993), even though
(various sites; Ferretti, in progress), Canale delle Acque variability is great. On both the anterior and posterior
Alte (Pianura Pontina, Latina; Palombo, 1994), Sette- enamel crest of each loop, one to two acute enamel folds
polesini di Bondeno (Ferrara; Gallini and Sala, 2001; are frequently present.
DSTF), Buca delle Iene (Alpi Apuane, Massa Carrara), The Italian material of E. (P.) antiquus currently
and Arene Candide (Savona; Cassoli and Tagliacozzo, available provides detailed knowledge of the entire skull
1994). Excluding Asolo, where a partial skeleton was and skeleton of this species, together with ontogenetic
found (Reggiani and Sala, 1992), all other localities details. In particular, comparisons of juvenile and adult
yielded only isolated dental and skeletal remains. The skulls clarify the development of the frontal ridge in
material listed above can be confidently referred to M. adults, a unique feature of this group. Complete E. (P.)
primigenius on the basis of the derived character of the antiquus skeletons were found in diatomaceous deposits
molars, namely elevated plate count, high lamellar cropping out at Riano (Maccagno, 1962b; MPR) and
frequency and thin enamel (Fig. 10). Grotte S. Stefano-Fonte Campanile (Viterbo; Trevisan,