Sus Linnaeus, 1758

Guérin, Claude & Tsoukala, Evangelia, 2013, The Tapiridae, Rhinocerotidae and Suidae (Mammalia) of the Early Villafranchian site of Milia (Grevena, Macedonia, Greece), Geodiversitas 35 (2), pp. 447-489 : 470-472

Genus Sus Linnaeus, 1758

Sus arvernensis arvernensis

Croizet & Jobert, 1828

Four species of Suidae are known in the Pliocene and Early Pleistocene of Europe ( Guérin 1996; Arribas & Garrido 2008): Potamochoerus provincialis (Gervais, 1859) , exclusively of Ruscinian age, the Villafranchian Potamochoerus magnus Arribas & Garrido, 2008 , Sus arvernensis Croizet & Jobert, 1828 , of the Ruscinian (zones MNQ 14 and 15) and the Early Villafranchian (zone MNQ 16), and Sus strozzii Meneghini, 1858 , F. Major, 1881, of the Villafranchian (zone MNQ 16 to 19). e Milia Suidae belongs to S. arvernensis .

Sus arvernensis is a small species closer to the eastern boars, with a long rostrum of Sus verrucosus type, than to the recent European wild boar Sus scrofa Linnaeus, 1758 . Guérin et al. (1998) estimated its mass to be between 45 and 100 kg. He recognised two successive sub-species, S. arvernensis minor Depéret, 1890 of the Ruscinian and S. arvernensis arvernensis Croizet & Jobert, 1828 of the Early Villafranchian. e cheek teeth are still relatively simple and the last molars usually end in a single tubercle. Recall that the syntype of the species consists of a fragment of right maxilla bearing D2/, D3/, D4/ and M1/, and a mandible fragment bearing D/2, D/3, D/4 and M/1 on both sides, and the bud of the right M/2 ( Croizet & Jobert 1828: pl. 13, figs 3-5).

e Auvergne boar is defined in France in the Early Villafranchian of Perrier-les Étouaires, it is also found in Vialette and in Trévoux-Reyrieux in Ain ( Guérin et al. 1998); its primitive subspecies is known from the Ruscinian in the Perpignan region, in Cavaillé brickfield, in Serrat d’en- Vacquer (type-site of S. arvernensis minor ) and Mas Bruno near Saint-Estève, and Autrey near Gray in Haute-Saône ( Depéret 1890; Guérin et al. 1998). Guérin et al. (1998) pointed out that this species is also found in Great Britain (Red Crag of Suffolk), in Germany (Herbolzheim), in Hungary (Süttö), in Slovakia (Hajnacka and Ivanovca), in Moldova and probably in Georgia (Kvabébi), and in Turkey (Afyon-Dinar-Akçaköy and Çalta). ey also recall that the species is not rare in Italy (Tuscany: upper levels of Casino, Val di Pugna, Barga, Pieve Fosciana and Piedmont: Triversa, Villafranca d’Asti). Also in Italy, Campanino et al. (1994) described a third upper premolar of “ Sus minor ” discovered in Roatto near Villafranca d’Asti of the same age as the Dicerorhinus jeanvireti remains. Finally, Gliozzi et al. (1997) reported a Sus sp. of “small size”, in Collepardo in Anagni Basin (Lazio), which is probably S. arvernensis because of its size ( Guérin et al. 2004).

In Spain, Sus arvernensis is reported by Van der Made (1989 -90) in Gorafe IV (MNQ 14), by Montoya et al. (2006) in Alcoy-Mina (MNQ 14 or 13?) and by Mazo & Torres (1989 -90) in Piedrabuena (MNQ 16).

In Romania, Radulescu et al. (2003) and Radulescu (2005) indicate the presence of “ Sus minor ” in the Brasov Depression at Capena (MNQ 15b) and Vargha (MNQ 15b).

In Bulgaria, Spassov (2005) reported Sus arvernensis minor in Musselievo (Late MN 15).

Finally, with regards to Greece, Sus arvernensis may be present in Ptolemaida/ Servia ( Tobien 1981). In addition, two indeterminate suids are reported in the Ruscinian Damatria Formation of Rhodes Island, which could correspond to “ Sus ” minor according to Koufos (1986).

MATERIAL

Concerning the Auvergne boar, nine specimens have been recorded in the Milia faunal assemblage, six of which are measurable and listed as follows: – a right upper tooth row MIL 207 with P4/-M3 /; – a left mandible fragment MIL 696, with M/2 and the roots of other cheek teeth;

– a right mandible rostral fragment with part of the canine MIL 1283;

– a right upper canine MIL 511;

– a right lower canine MIL 1485;

– a left radius proximal fragment MIL 916.

DESCRIPTION

Upper tooth row and canine

It belongs to a young adult because P4/ and M2/ display no traces of wear, and M3/, although in place, is unworn ( Fig. 16 A-C). e principal cusp of P4/ is not quite central, it is slightly shifted buccally; M1/ and M2/ are typical of Suinae with their four principal tubercles; they have a peripheral cingulum which for the M2/ is slightly folded behind; the M3/ has in addition to the four principal tubercles, a large posterior axial tubercle. ere are many accessory cuspids on the anterior cingulum, on the posterior of the paracone and the metacone and around the metaconule; this tooth resembles the homologous tooth from Çalta ( Guérin et al. 1998: fig. 1E).

e dimensions of the cheek teeth are given in Table 13 (see Appendices). e dimensions of the upper cheek teeth fall within the range of variation of a sample from 5 to 28 specimens of Sus arvernensis , the lengths are close to the maximum values measured on our comparative sample, the widths are however somewhat lower than the observed average.

e canine MIL 511 is rather slender, its crown width is about 18 mm and the length about 15 mm.

Rostral fragment of mandible is fragment bears part of the canine, whose tip is broken ( Fig. 16E, F ). e lower canine has a triangular cross section, whose rounded hypotenuse covers approximately 18 mm and corresponds to the antero-lingual border. e postero-dorsal border without enamel is about 13 mm wide. e external border is about 17 mm. e inferior dihedral is acute. e cross-section and size of the canine are similar to those of the Çalta boar and as for the latter, the Milia boar approaches recent Asian Sus verrucosus ( Guérin et al. 1998) . e canine MIL 1485 is much stronger and its hypotenuse covers approximately 24 mm and corresponds to the antero-lingual border. e postero-dorsal bor- der without enamel is about 18 mm wide, and the external border is about 22 mm.

M/2

It is present in a hemi-mandible fragment, in which there are also roots of P/4 and M/1, and an anterior root of M/3 ( Fig. 16D ). It is barely worn and its morphology requires no special comment. e three dimensions of the M/2 are close to the average of nine to ten specimens of S. arvernensis (Appendices, Table 13).

Radius e proximal epiphysis of the Milia radius has DT = 27.5 mm for DAP = 19 mm. e same two dimensions are respectively 25 and 22 mm in Çalta specimens, and 26 and 19.5 mm for the two radii from Villafranca d’Asti.

CONCLUSION ON THE ANATOMICAL STUDY

OF SUIDAE

Because of its dimensions, in particular its upper cheek teeth, the Milia Suidae belongs to the nominate subspecies Sus arvernensis arvernensis , S. arvernensis minor being smaller.

BIOSTRATIGRAPHIC IMPLICATIONS

Sus arvernensis arvernensis is a marker of the Early Villafranchian (zone MNQ 16).

PALAEOECOLOGICAL IMPLICATIONS

Sus arvernensis was a species of open forest in relatively warm regions, well adapted to digging deep into the humid soils.

Figures

FIG. 16. — The Milia Sus arvernensis arvernensis Croizet & Jobert, 1828: A-C, right maxilla with P4/-M3/ MIL 207; A, occlusal view; B, lateral view;C, medial view; D, left mandible fragment with M/2 MIL 696, occlusal view; E, F, rostral fragment of mandible with part of the canine, MIL 1283 dex; E, lateral view; F, front view showing details of the canine section. Scale bars: 50 mm.