Tapirus Brisson, 1762

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 : 449-452

Genus Tapirus Brisson, 1762

Tapirus arvernensis arvernensis

Croizet & Jobert, 1828

Two species of tapirs are known in the Pliocene and Pleistocene of Europe ( Boeuf 1991; Guérin & Eisenmann 1994): Tapirus arvernensis Croizet & Jobert, 1828 and T. jeanpiveteaui Boeuf, 1991 . e latter is a small species that is currently known only from its type-locality at Barro, near Ruffec in Charente ( France); the assigned Pliocene age is not certain.

e Tapir of Milia is attributed to T. arvernensis .

Tapirus arvernensis is slightly larger and much more common than T. jeanpiveteaui . It is defined in Perrier-Les Étouaires (Puy-de-Dôme, France) and anatomically it is close to recent T. terrestri s (Linnaeus, 1758) from South America. e typeseries is composed of a left hemi-mandible with P/3-M/3, a juvenile hemi-mandible with D/1-D/4 and M/1, a right I2/, an atlas and an upper molar ( Croizet & Jobert 1828: pl. II, figs 1, 3, 5 and pl. XII, figs 4-6). Although smaller, T. arvernensis presents proportions similar to those of T. priscus Kaup, 1833 of the Late Miocene of Europe ( Guérin & Eisenmann 1994).

Its size corresponds to the recent T. pinchacus Fischer, 1829 of the Andes Mountains. Its total length reached 1.8 to 2 m, its shoulder height ranging from 75 to 80 cm and it weighed over 200 kg.

Tapirus arvernensis minor Michaux, Sigé & Sudre, 1976 , is a subspecies defined in the sands of Montpellier (Early Ruscinian, zone MNQ 14), which is smaller and more slender than the nominate subspecies. e authors revived a species name created in 1839 by M. de Serres, but later on abandoned it because it was a recent synonym of T. arvernensis . Tapirus a. arvernensis is known in the Late Ruscinian (MNQ 15) of Roussillon and the Villafranchian of the Massif Central ( Michaux et al. 1976).

Guérin & Eisenmann (1994) characterized T. arvernensis anatomically as follows: compared to T. terrestris , the mandible of T. arvernensis shows the same proportions as the Late Miocene European T. priscus , including a relative height of the corpus that differs from the extant species. e relative lengths of the upper premolars and molars differ from those of extant species: compared to T. terrestris the P1/ and P2/ are longer while the M1/, M2/ and M3/ are shorter. In T. arvernensis arvernensis the average lengths of the lower premolars are smaller than those of T. terrestris (they are the same as in T. pinchacus ), except for P/2 and P/4; P/2 and P/3 which are large; the posterior widths of P/3 and P/4 are roughly equivalent to those of T. priscus . e average length of M/1 is barely greater than that of the P/4. e relative width of P/3 and P/4 is the only characteristic that seems to distinguish European Miocene and Pliocene tapirs from extant tapirs (with the exception of T. pinchacus ). e long bones have proportions reminiscent of a small T. terrestri s with particularly robust diaphyses.

Tapirus arvernensis is known in the Ruscinian formations of Montpellier and Perpignan (MNQ14 and 15 zones) and is remarkably abundant in Vialette and common in Perrier-Les Étouaires, both sites of Early Villafranchian age (zone MNQ 16). It is at- tested by a tooth fragment in the Late Villafranchian site (zone MNQ 18) of Le Coupet in Haute-Loire ( Heintz et al. 1974), which is perhaps the youngest occurrence of European tapirs. However, if the remains of T. arvernensis of Tegelen ( Netherlands) are contemporary with Sus strozzii Forsyth Major, 1881 and Dicerorhinus mercki (Jäger, 1839) of the same deposits, this species may extend up to the very early Middle Pleistocene (the stratigraphic position of Tegelen large mammals is unclear, see this topic in Guérin 1980: 978, 979). Tapirus arvernensis is well represented in the Late Ruscinian and Early Villafranchian of Italy. us, it is found in the upper levels of the Casino, Meleto, Barga and Pieve Fosciana (Garfagnana), Sarzanello and Ponzano Magra in Val di Magra, in Gaville and Santa Barbara (both in the Upper Valdarno), and in Monticchiello in Tuscany. It is also known in Monte Biancano and Vignola in Emilia Romagna, in Villafranca d’Asti and Triversa in Piedmont, in Livergnana and Sasso di Glosina near Bologna, in Spoleto in Umbria, in Castel San Pietro (Nera Montoro) in Latio ( Kotsakis 1986; Rustioni 1992). According to Rook & Rustioni (1991), it is surely present in the V3 level of Baccinello in Tuscany, probably of Latest Turolian age (MN 13). Tapirus arvernensis is also known from the Early Villafranchian of Hajnacka in Slovakia ( Fejfar 1964; Janstova 2004). In Romania, Radulescu et al. (2003), reported T. arvernensis in the Dacian Basin in Malusteni (MNQ 15a), and in the Brasov Depression in Capena (MNQ 15b), Vargha (MNQ 15b), Iaras 1 (MNQ 16), Araci- Fântâna Fagului (MNQ 16a) and in the Ilieni Basin (MNQ 16).

e species had not hitherto been studied in Greece. ere is only a short reference by Paraskevaidis (1977) of three upper cheek teeth of Tapirus sp. from Servia (W. Macedonia), without description and measurements, but the present location of the specimen is unknown.

MATERIAL

– A fragment of right hemi-mandible of a juvenile MIL 649, with the alveoli of D/2 and D/3, with D/4 and M/1; M/2 is unerupted, thus it is not measurable.

– Calcaneum of an adult MIL 1282 dex.

DESCRIPTION e mandible fragment and teeth

On the mandible fragment ( Fig. 2 A-C), the height of the corpus between D/4-M/1 reaches 46.5 mm and its transverse diameter at the same level is 27 mm. For nine specimens of T. arvernensis the height between P/4 and M/1 averages 46.1 mm (range: 42 to 49.8 mm). e lower cheek teeth have the characteristic appearance of the family with their two transversal ridges and anterior and posterior cingula, particularly well distinguished on the M/1 and M/2 ( Fig. 2 A-C).

e dimensions of the teeth are given in Table 1 (see Appendices), the specimen belongs to juvenile, the D/4 is slightly worn, and the M/1 is not worn at all; the M/2 is not measurable. e Milia M/1 slightly exceeds the average length of 13 specimens of Tapirus arvernensis arvernensis , and the two widths are equal to the maximum values measured on a sample of 11 specimens of the same subspecies; thus, in Milia we are dealing with the nominate Villafranchian subspecies, larger than the Ruscinian one.

Janstova (2004) gives some dimensions of undifferentiated M/1 and M/2 which were collected during the new excavations in Hajnacka: length 22 to 23 mm for five specimens, anterior width 14 to 17 mm for 12 specimens.

Calcaneum is bone looks very rhinocerotoid ( Fig. 2D, E ) with elevated tuberosity, the beak (= foremost part of the bone) extending at least as far as the tuberosity, and the relatively small sustentaculum tali perpendicular to the vertical axis of the bone.

e dimensions of the calcaneum are in Table 1 (see Appendices).

Compared to a sample of nine to ten specimens of T. arvernensis from the Ruscinian of Montpellier (one specimen) and from the Early Villafranchian of Vialette (all others), the Milia calcaneum is particularly large: the observed maximum values of its dimensions exceed all the others

Owing to the largest size of the nominate subspecies this tends to confirm its subspecific identification, but the insufficient size of our comparison sample do not allows us to know precisely the extreme values of the calcaneum dimensions and then prevents us to be certain of it.

Conclusion of the anatomical study of the tapir Because of their size, the remains of the Milia tapir, which are larger than those of Tapirus arvernensis minor , can therefore be attributed to T. arvernensis arvernensis .

BIOSTRATIGRAPHIC IMPLICATIONS

While Tapirus arvernensis minor is characteristic of the Early Ruscinian (zone MNQ 14), T. arvernensis arvernensis is a subspecies typical of the Late Ruscinian (zone MNQ 15) and the Early Villafranchian (MNQ 16), its presence in more recent levels seems exceptional.

PALAEOECOLOGICAL IMPLICATIONS

A consensus has been reached that the fossil tapirs with anatomical characteristics similar to those of extant species, which is the case of T. arvernensis , indicate similar ecological requirements: warm climate, dense forest, moist environment and proximity to sufficient amount of water ( Eisenmann & Guérin 1994; Guérin & Eisenmann 1994).

Sub-order CERATOMORPHA Wood, 1937

Figures

FIG. 2. — The Milia Tapirus arvernensis arvernensis Croizet & Jobert, 1828: A-C, fragment of right juvenile hemi-mandible MIL 649; A, occlusal view; B, buccal view; C, lingual view; D, E, right calcaneum MIL 1282 of adult; D, medial view, E, posterior view. Scale bar: 50 mm.