The town of Montopoli Val d’Arno is located in Tuscany (central Italy), about 27 km east-southeast of Pisa and 40 km west-southwest of Florence. Most of the Montopoli collection comes from excavations made by Forsyth Major in 1880 and was found in gravelly sands alternating with marine littoral deposits. Continental mammal-bearing sands outcropped in at least two localities: “l’Uccellatoio” hill and “Poggio di Montevecchio”, both located a few kilometers southeast of the town of Montopoli (De Giuli and Heintz, 1974). The felid tooth described in the present work probably comes from the former locality. Fossil mammals from Montopoli have been analyzed by several authors since their discovery (Azzaroli, 1977, Benvenuti et al., 1995, De Giuli and Heintz, 1974, Ficcarelli, 1984, Forsyth Major, 1885 and Merla, 1949). According to published data the fauna is composed of the following taxa: A. pardinensis, Nyctereutes megamastoides, Pliocrocuta perrieri, Mammuthus gromovi, Stephanorhinus jeanvireti, Stephanorhinus etruscus, Equus livenzovensis, Axis lyra, Croizeratoceros ramosus, Eucladoceros falconeri, Procapreolus cusanus, Gazella borbonica, Leptobos stenometopon.

The tooth IGF 15358 from Montopoli (Fig. 2) is labeled as Acinonyx sp. in the current IGF catalog, although the identification “Viretailurus schaubi” was written with a pencil on the original paper catalog by an unknown past curator of the Museum. Viretailurus schaubi is an older synonym of Puma pardoides, according to Hemmer (2001). Tooth measurements were taken following Hemmer et al. (2004).

The left upper carnassial IGF 15358 is in good state of preservation, although the two anterior roots are broken (Fig. 2).

In occlusal view, the tooth shows a peculiar T morphology (Argant, 2004), caused lingually by the advanced position of the protocone, and labially by the bulging of the antero-labial base of the parastyle, that does not form a true ectoparastyle. This feature seems to be less evident in the extant P. concolor (Fig. 3I), which is however characterized by strong morphological variability in its dentition (Hemmer et al., 2004).

The protocone is a well-developed cusp, slightly oriented frontwards and noticeably low-placed toward the root, as one can note in anterior view. The triangular pit formed by the parastyle, paracone, and protocone is quite deep, and a clear ridge runs from the paracone tip to the protocone. The parastyle is globular and much worn on the labial side. The indentation between the para- and metacone is deep and open.

All these morphological characters have been described for P. pardoides from various Eurasian sites (Argant, 2004, Hemmer, 1965, Hemmer et al., 2004 and Viret, 1954).

It is evident that the tooth from Montopoli does not belong to a cheetah, Acinonyx sp., as indicated in the IGF catalog. In fact, the Villafranchian giant cheetah, A. pardinensis, is characterized by a larger and stronger carnassial, with a very reduced and posteriorly-placed protocone (Fig. 3H).

Morphometric values for IGF 15358 are very close to those of the holotype of “Panthera” schaubi from Saint-Vallier (Hemmer, 1965 and Viret, 1954), but smaller than those of the other specimens from the same locality (Argant, 2004) and of P. pardoides from Kvabebi (Hemmer et al., 2004). Also the Recent puma, P. concolor, shows higher mean values for the upper carnassial (Table 1).

More detailed comparisons are needed for Lynx issiodorensis. In fact, although this species is not reported at Montopoli, it was undoubtedly present in Europe at the beginning of the Middle Villafranchian (Werdelin, 1981). In addition, as Kurtén highlighted (1978), the overall morphology of the Issoire lynx was more “generalized” than extant lynxes, thus resembling a puma in some respects (larger head, longer neck, shorter limbs, and heavier body). Nevertheless, dental differences between L. issiodorensis and P. pardoides seem to be quite definite, as Viret (1954) stressed in describing the fossils from Saint-Vallier, where both species have been reported. Although the two felids are dimensionally similar, the upper carnassial of P. pardoides is on average slightly larger than that of L. issiodorensis (Table 1). The main morphological differences lie in the following characters (for a graphic comparison, see Fig. 3A–C for P. pardoides and Fig. 3D–F for L. issiodorensis): P. pardoides has a stouter upper carnassial on the whole, with a Panthera-like, massive, lingually-projected protocone. The parastyle is stout and globular as well. In addition, L. issiodorensis normally shows a well-developed ectoparastyle (more prominent than in the living lynx; Kurtén, 1978), which gives a sharp appearance to the antero-labial angle of the tooth (black arrows in Fig. 3D–F). On the contrary, the parastyle does not bulge labially (T-shaped occlusal outline of P. pardoides). The P4 protocone of L. issiodorensis is more slender than in P. pardoides; furthermore, its posterior margin strongly recedes in occlusal view (grey arrows if Fig. 3D–F), whereas in P. pardoides the protocone is more clearly isolated from the lingual wall of the tooth, coming off with a sharper angle.

The length and maximum breadth of the examined tooth are close to the smaller values for the extant leopard, P. pardus (Table 1), but the latter normally shows a stronger and stouter carnassial, with the absence of the typical T-shaped occlusal morphology (Fig. 3G), as in L. issiodorensis.

The Montopoli Faunal Unit is the earliest unit of the Middle Villafranchian (Rook and Martínez-Navarro, 2010). This large mammal assemblage provides a clear indication of the strong environmental change that occurred at the beginning of the Pleistocene, with the disappearance of some of the taxa with subtropical affinities that still characterize the previous Early Villafranchian Triversa FU, as well as the dispersal of herbivores such as a primitive species of the genus Mammuthus, the monodactyl horse (Equus cf. livenzovensis) (the so-called “elephant-Equus event” of Azzaroli, 1977), the large deer Eucladoceros tegulensis, and the bovid Gazella borbonica. The Montopoli FU – corresponding to the MN16b unit in the European MN-system of Mein (1975) – was originally included in the Early Villafranchian (Azzaroli, 1977 and Gliozzi et al., 1997), but the clear importance of the faunal turnover observable at the transition from the Triversa FU to the Montopoli FU suggested considering the latter as the basal FU of the Middle Villafranchian (Rook and Martínez-Navarro, 2010).

The Montopoli large mammal assemblage and the related Faunal Unit occur at the Gauss/Matuyama boundary (∼2.6 Ma) (Lindsay et al., 1980). Therefore the Early/Middle Villafranchian transition corresponds to the newly redefined Plio/Pleistocene boundary, lowered to the Global Stratotype Section and Point (GSSP) of the Gelasian Stage at Monte San Nicola, Sicily (Gibbard et al., 2010 and Rio et al., 1994).

Puma-like cats – which probably represent the most poorly known carnivores within the European Pleistocene faunas – spread into western Europe at the beginning of the Pleistocene and their diffusion can be considered one of the bioevents that characterized the Early/Middle Villafranchian faunal turnover (Hemmer et al., 2004). As suggested by several authors (Hemmer et al., 2004, Madurell-Malapeira et al., 2010a, Salles, 1992, Spassov, 2011 and Van Valkenburgh et al., 1990), the puma is closely related to the cheetah and the snow leopard (although recent genetic analyses (Johnson et al., 2006) seem to put the close affinities between the snow leopard and the Puma-Acinonyx clade up for discussion). At present, the incompleteness of the fossil record cannot provide clear information on the origin and systematics of these small-headed large cats, which are normally distinguished only on the basis of craniodental features.

The Early and Middle Villafranchian European mammal assemblages are characterized by the occurrence of different large felids such as two machairodontine cats (Homotherium and Megantereon), the giant cheetah A. pardinensis, and L. issiodorensis, besides P. pardoides. During the Late Villafranchian, the Eurasian jaguar Panthera gombaszoegnesis also spread into Europe, together with the giant hyaena Pachycrocuta brevirostris (“Pachycrocuta brevirostris event” of Martínez-Navarro, 2010) and the first wolf/jackal-sized dogs of the genus Canis (“wolf event” of Azzaroli, 1983) (Palombo et al., 2008). In the present paper, we confirm that P. pardoides was present in western Europe at least from the Plio/Pleistocene transition (while its first occurrence in eastern Europe – Kvabebi, Georgia – and Asia – Shamar, Mongolia – is probably slightly older; Hemmer et al., 2004). Therefore, keeping in mind that its youngest record is at Vallparadís, Spain, around the latest Early Pleistocene (Madurell-Malapeira et al., 2010b), we can conclude that this species was present in Europe for at least 2 Ma. Probably, the presumed high ecological adaptability of P. pardoides – inferred from the very different paleoecological features of the sites where this species has been found, as well as from the ecological data for the closely-related extant P. concolor – allowed this big cat to find space in the complex framework of the Early Pleistocene carnivore guild. It is not accidental that the disappearance of P. pardoides from the Eurasian fossil record at the end of the Early Pleistocene coincides with the arrival of an other ecologically flexible large cat, the leopard P. pardus (probably first recorded in the Vallonnet Cave at about 1.0 Ma, Moullé et al., 2006), which probably replaced the Eurasian puma occupying the same ecological niche. Unfortunately, even in this case the scantiness and ambiguity of the fossil record make the definition of this ecological “transition” unclear.

The occurrence of P. pardoides in the Montopoli assemblage is here documented thanks to the description of a left upper carnassial stored at the IGF. This fossil therefore can be considered as the earliest evidence of this species in Italy at the Plio/Pleistocene transition, when this felid spread into western Europe possibly from Asia. The Montopoli Faunal Unit, now referred to the Middle Villafranchian (Rook and Martínez-Navarro, 2010), can be related to an important faunal change driven by the climate change at the beginning of the Pleistocene, with the dispersal and diffusion of elephantids, gazelles, antelopes, and monodactyl horses. The puma-like cat P. pardoides was one of the top predators, together with the giant cheetah, machairodontine cats, and – later on – pantherine cats, that characterized the western Europe terrestrial ecosystems for hundreds of thousands of years. This felid became extinct at the end of the Early Pleistocene, when lions and leopards spread widely into Europe.