Order Carnivora Bowdich, 1821

Family Mustelidae Fischer de Waldheim, 1817

Genus Eomellivora Zdansky, 1924

Eomellivora wimani Zdansky, 1924

Locality. Ravin de la Pluie (RPl), Axios Valley, Macedonia, Greece (Fig. 1).

Age. Late Vallesian, MN 10; GPTS = ∼9.3 Ma

Material. Left upper first molar, LGPUT-RPl-252.

Measurements. M1: 9.1 × 19.7, L_lingual = 12.2

Description: The cusps of the tooth are worn and low, indicating a relatively old individual (Fig. 2). The tooth has a strong constriction in the middle of its width. The paracone is strong, but the metacone is weak and quite worn (barely distinguishable). The paracone is surrounded by a strong mesio-labial cingulum. The protocone is large, situated in the mesial part of the lingual lobe of the tooth. It is connected through the pre-protocrista (nomenclature according to Bryant et al., 1993) with the paracone. It is surrounded mesially, lingually and distally by a strong cingulum; although it shows advanced wear, it is clear that the cingulum consists of small cusps. The mesial extension of the lingual cingulum is smaller than the distal one. The posterocrista is slightly distinguished in its origin from the paracone but cannot be followed because the enamel is slightly broken.

Discussion: An extensive revision of the genus Eomellivora, as well as its synonymy and systematics, are given by Wolsan and Semenov (1996), whose conclusions are accepted in this article. Eomellivora, compared to the modern Mellivora capensis, is clearly larger (Fig. 3). Apart from its smaller size, the M1 of M. capensis has a longer lingual than buccal mesiodistal diameter, a relatively larger paracone, and no metacone. The protocone is surrounded by a stronger and more prominent cingulum than those of LGPUT-RPl-252 and Eomellivora wimani.

The comparison of LGPUT-RPl-252 with the M1 of E. wimani from China (Zdansky, 1924: p. 61; taf. XII, Fig. 1) indicates that they are morphologically and metrically (Fig. 3) similar. However, the metacone is better developed in the Chinese sample but this is possibly due to the more advanced attrition of the studied specimen. The metacone of Eomellivora is vestigial (Wolsan and Semenov, 1996) and attrition decreases it rapidly. A maxillary fragment with P4-M1 and an isolated M1 (MNHN-TRQ-1005, 1006) from the Turkish locality of Middle Sinap are referred to E. piveteaui (Ozansoy, 1965: p. 23: pl. I, Figs. 1–5) and recently to the subspecies E. w. piveteaui (Wolsan and Semenov, 1996). A direct comparison of LGPUT-RPl-252 with the Sinap M1 indicates that they have similar morphology and size (Fig. 3). Because the Sinap teeth are less worn, their metacone and posterocrista seem to be stronger and more expressed. A skull and an incomplete mandible of a large mustelid from Grebeniki, Moldavia were described as Perunium ursogulo (Orlov, 1948) and later synonymized with E. wimani (Ozansoy, 1965 and Wolsan and Semenov, 1996). LGPUT-RPl-252 is similar to the Grebeniki M1 (Orlov, 1948: Figs. 2 and 5) in occlusal morphology and size (Fig. 3). One M1 from the Early Vallesian Spanish locality of Los Valles de Fuentidueña was described as a new species under the name E. liguritor and later synonymized with E. wimani (Crusafont Pairo and Ginsburg, 1973 and Wolsan and Semenov, 1996). The morphological characters of the Spanish Eomellivora are similar to those of E. wimani. The differences noted for the latter, such as the more developed lingual cingulum and the slightly less developed protocone (Crusafont Pairo and Ginsburg, 1973) are not as indicative. The M1 size of E. liguritor is quite similar to that of E. wimani from China, Grebeniki and LGPUT-RPl 252 (Fig. 3), confirming the synonymy of E. liguritor with E. wimani (Wolsan and Semenov, 1996). A new species of Eomellivora, under the name Eomellivora tugenensis is referred from Ngorora, Kenya (Morales and Pickford, 2005). Its M1 resembles that of E. wimani but it is much smaller (Fig. 3). Eomellivora is rare in eastern Mediterranean, and known only from Middle Sinap (Turkey) until now. Its presence at RPl marks its first appearance in Greece and the Balkan Peninsula.

Family Hyaenidae Gray, 1821

Genus Plioviverrops Kretzoi, 1938

Plioviverrops orbignyi (Gaudry and Lartet, 1856)

Locality. Ravin des Zouaves-5, RZO, Macedonia, Greece (Fig. 1).

Age. Early Turolian, MN 11; GPTS = ∼8.2 Ma.

Material. Right mandibular fragment with m1–m2, LGPUT-RZO-166.

Measurements. m1 = 9.0 × 4.5; m2 = (5.2) × –.

Height of the mandibular corpus below m1 (lingual) = 11.7

Height of the mandibular corpus below p4 (lingual) = 11.0

Thickness of the mandibular corpus below m1 = 6.1

Description: The studied specimen preserves the mandibular corpus from the p4 to the mandibular angle and the two molars; the m2 is broken disto-buccally (Fig. 4A). The mandibular corpus is shallow and quite thick. The masseteric fossa is deep and its anterior border ends below the distal border of the m2. The m1 is short and wide with very high cuspids; the trigonid cuspids (proto-, para- and meta-conid) are of almost similar height. The talonid is well distinguished from the trigonid by a deep valley and bears a high entoconid and a weak hypoconid (Fig. 4A₃). The labial cingulum is well developed. The m2 preserves only the paraconid, which is high.

Discussion: Extensive comparisons of P. orbignyi with the other species of the genus and with those of Protictitherium, as well as its stratigraphic distribution, have been given recently (Koufos, 2006b and Koufos, 2009). The high trigonid cuspids, the bicuspid and well separated talonid, as well as the deep valleys of the carnassial in the studied specimen characterize P. orbignyi and allow its attribution to this species. The dimensions of the studied carnassials are also in the range of variation of P. orbignyi, confirming its attribution.

Genus Adcrocuta Kretzoi, 1938

Species Adcrocuta eximia (Roth and Wagner, 1854)

Adcrocuta eximia leptoryncha Bonis and Koufos, 1981

Locality. Ravin de la Pluie (RPl), Axios Valley, Macedonia, Greece (Fig. 1).

Age. Late Vallesian, MN 10; GPTS = ∼9.3 Ma.

Material. Right upper canine, LGPUT-RPl-269; left mandibular fragment with i2, c-m1, LGPUT-RPl-272.

Measurements. C:16.2 × (11)

Mandible: Length anterior c-distal side of condyle = 194.5; Height ventral border of angular process-dorsal side of condyle = 43.4; Symphysis length = 60.0; Height in front of p2 (lingual) = 43.0; Idem in the middle of p3 (lingual) = 40.8; Idem in p4 (lingual) = 43.0; Idem in m1 (lingual) = 46.2; Diastema c-p2 = 10.0; Alveolar length c-m1 = 106.2; Alveolar length p2-m1 = 81.2; Alveolar length p2-p4 = 55.1.

i2: 7.0 × 5.5; c = 15.0 × 13.3; p2 = 13.7 × 9.0; p3 = 18.3 × 11.5; p4 = 20.5 × 11.7, m1 = 27.6 × 11.7, L _{m1 tal.} = 5.0 (the talonid length is measured from the distal wall of the protoconid to the distal end of the talonid).

Description: The hemimandible LGPUT-RPl-272 lacks the ascending ramus and the teeth i1 and i3 (Fig. 4B). The mandibular corpus is elongated and shallow for its size, with two mental foramina that are connected and situated below the p2. The symphysis is strong and inclines abruptly backwards. The masseteric fossa is large and oval, and its anterior border is well behind the m1. The dentition is well preserved and the p1 is absent. The premolars are narrow and their main cuspid inclines backwards. The p2 is small without an a.a.c. and with a small p.a.c (Fig. 4B_1,3). The p3 bears a small a.a.c. and p.a.c., as well as a strong distal cinglum. The p4 has a strong a.a.c. and a smaller p.a.c., situated on a large distal projection of the distal cingulum. The carnassial is narrow, without metaconid, and it has a small and bicuspid talonid. The isolated upper canine has the typical morphology and size of A. eximia (Fig. 4E).

Discussion: The general morphology of the studied mandible and teeth fits that of A. eximia; however, the RPl A. eximia differs from the Pikermi one, representing the new subspecies A. e. leptoryncha; it is known by a skull and a badly preserved mandibular fragment (Bonis and Koufos, 1981). The absence of the p1, the more slender teeth, the smaller premolars, and the narrower m1 of the studied mandible distinguish it from the typical A. e. eximia of Axios Valley, Pikermi and Samos. On the other hand, its dental dimensions are similar to those of A. e. leptoryncha (Fig. 5) and allow its attribution to this subspecies.

Adcrocuta eximia (Roth and Wagner, 1854)

Locality. Ravin de la Pluie (RPl), Axios Valley, Macedonia, Greece (Fig. 1).

Age. Late Vallesian, MN 10; GPTS = ∼9.3 Ma.

Material. Left maxillary fragment with DP2-DP4, LGPUT-RPl-244; fragment of a left DP3, LGPUT-RPl-271.

Measurements. The measurements are given in Table 1.

Description: The DP2 is small and relatively short and wide. In its mesiolingual corner there is a small projection of the cingulum, like a vestigial a.a.c., whereas the p.a.c. is distinct and low (Fig. 4C₂). The DP3 is elongated and narrow, with a very low protocone extended lingually from the middle of the tooth; the protocone is worn and its height does not exceed the root of the tooth (Fig. 4C₂). The parastyle is moved mesially and it is distinguished like a cusp; in its mesiolingual corner there is a small cuspule (Fig. 4C₂). The metastyle is blade-like, elongated and slightly curved buccally. The DP4 is triangular and low-crowned. Both paracone and metacone are very low and they are connected by a thin crest, running from the mesial border of the paracone to its apex and through the metacone's apex to its distal border (Fig. 4C₂). The protocone is more developed than the other cusps but still low, and it is situated in the lingualmost angle of the tooth. The paracone is projected buccally, and between it and the metacone there is a deep buccal sinus. The DP3 fragment lacks its mesial part with the parastyle, and it is similar to the one described but more worn (Fig. 4D).

Discussion: The species A. eximia is well known and widespread in the Late Miocene, covering the whole of Eurasia from Spain to China. Although it is a common taxon, the remains of its deciduous dentition are few. The upper deciduous dentition is known by two fragments from Pikermi (Gaudry, 1862–67:pl. 12, Figs. 4 and 5) and another two from Perivolaki (Koufos, 2006b: pl. 4, Fig. 6). Another maxillary fragment is known from the Turkish locality of Akkaşdaği (de Bonis, 2005). Some undescribed material of A. eximia with deciduous teeth from Samos is included in Brown's collection at AMNH (Kostopoulos, pers. comm.). The Pikermi specimen MNHN-PIK-3025 preserves the DP3-DP4 and the just erupted M1. The M1 resembles that of A. eximia and its size falls within the range of variation for this species (Fig. 6), certifying its assignment to A. eximia. The milk teeth studied are similar in morphology and size to those from Pikermi and Perivolaki (Table 1) and can be attributed with certainty to A. eximia. A maxillary fragment of Hyaenictis graeca with the DP3-DP4 and P4-M1 is known from Pikermi (Gaudry, 1862–67). A direct comparison of the studied material with a cast of H. graeca suggests that the general morphology of its deciduous dentition is similar to LGPUT-RPl-244; however, the DP3 protocone is higher and more projected lingually, the metacone of the DP4 is higher and the teeth are slenderer than in A. eximia (Table 1). As mentioned above, the RPl Adcrocuta belongs to the subspecies A. e. leptoryncha. The deciduous teeth studied could belong to this subspecies but it is impossible to confirm this, becasue they have no differences from the Pikermi ones; hence they are referred to as A. eximia subsp. indet.

Genus Hyaenictis Gaudry, 1861

?Hyaenictis sp.

Locality. Ravin de la Pluie (RPl), Axios Valley, Macedonia, Greece (Fig. 1).

Age. Late Vallesian, MN 10; GPTS = ∼9.3 Ma.

Material. Right P2, LGPUT-RPl-270.

Measurements. P2: 20.3 × (10.8) mm.

Description: The studied tooth is badly preserved, lacking parts of its mesial and lingual base (Fig. 4F_2,3). The distolingual part is also broken postmortem and lacks the cingulum. The mesial cingulum seems to project mesiolingually but the presence of the a.a.c. cannot be checked. The main cusp is relatively low and without inclination. The p.a.c. is large and relatively high, although it is worn. The distal cingulum is not projected but it forms a slight edging in the distolingual part of the tooth. The buccal cingulum is weak.

Discussion: The studied P2 has hyaenid features and must belong to a species of this family. Excluding the smaller ictitheres, several large hyaenids are known from the Late Miocene of Greece. The most common and well-known is A. eximia. The P2 of A. eximia differs from the one studied in its more robust features, the almost straight mesial and distal border, the higher main cusp, the presence of a large a.a.c., the larger and more mesio-distally compressed p.a.c., and its larger size (Fig. 7).

Chasmaporthetes bonisi is a large hyaenid known from Ravin des Zouaves-5, (RZO) and Dytiko-1 (DTK) of the Axios Valley (Koufos, 2000 and refs. cited). The P2 studied differs from that of C. bonisi, in having more slender features, a lower main cusp, a more angular distal border, absence of the distal projection of the distal cingulum, and smaller size (Fig. 7). A large hyaenid named Lycyaena chaeretis is known from Pikermi and Samos (Forsyth Major, 1894 and Gaudry, 1862). The P2 studied is significantly larger than that of L. chaeretis (Fig. 7) and differs from it in having a lower main cusp, more slender features, and a less mesio-distally compressed p.a.c. The P3 of L. chaeretis is similar in size to LGPUT-RPl-270, but it differs from it in having a more elongated-elliptical occlusal aspect, a slenderer feature, and a stronger p.a.c.

Another large hyaenid known from Pikermi is H. graeca (Gaudry, 1862–67). A direct comparison of the studied tooth with a cast of H. graeca from the specimen NHMW-PIK-A.4715 indicates some similarities, such as the general aspect of the P2, the strong lingual cingular projection, the large and narrow p.a.c. and the similar size (Fig. 7). The above-mentioned comparisons suggest that LGPUT-RPl-270 is morphologically and metrically closer to Hyaenictis. However, the sole known and badly preserved specimen cannot allow certain determination; hence it is referred to ?Hyaenictis sp.

Family Percrocutidae Werdelin and Solounias, 1991

Genus Dinocrocuta Schmidt-Kittler, 1976

Dinocrocuta sp.

Locality. Xirochori-1 (XIR), Axios Valley, Macedonia, Greece (Fig. 1).

Age. Late Vallesian, MN 10; GPTS = ∼9.6 Ma.

Material. Right maxillary fragment with DP2-DP4, LGPUT-XIR-5.

Measurements. See Table 1.

Description: The specimen preserves well the DP2-DP4; the DP4 lacks its lingual half, but its alveolus is distinct, allowing estimation of its breadth (Fig. 4G). The DP2 is relatively short and robust without an a.a.c. but with a strong mesial cingular projection, a large p.a.c. and a strong lingual cingulum in the distal half of the tooth (Fig. 4G₂). The DP3 is elongated and relatively narrow with a weak protocone restricted to a basal lingual projection (Fig. 4G₂). The parastyle is moved mesially, forming a large cusp, well separated from the paracone by a deep valley. In the mesio-buccal base of the parastyle there is a strong cingulum. The paracone is high and narrow with a well developed cuspule situated in its mesio-lingual border; the cuspule is connected with the paracone's apex by a slight crest. The metacone is straight, short and low. The preserved part of the DP4 includes its buccal half, which bears a low paracone that is strongly projected bucco-distally, and a very small metacone, situated distally and separated from the paracone's projection by a deep sinus (Fig. 4G₂).

Discussion: The deciduous dentition studied differs remarkably from that of A. eximia. In addition to its larger size (Table 1), there are some morphological differences. The DP2 of LGPUT-XIR-5 has a clear and large mesial cingular projection, which is absent in A. eximia (Fig. 4C and G) and it is more robust than the DP2 of A. eximia; its robusticity index (R.I. = B × 100/Length is 54 versus 47–49 in A. eximia (Table 1). The protocone of the DP3 in LGPUT-XIR-5 is more developed and the metacone relatively shorter than that of A. eximia; the index L_metacone × 100/LDP3 is 32 in the studied DP3 versus 36 in A. eximia. The DP3 of LGPUT-XIR-5 is more robust than that of A. eximia; its R.I. is 43 versus 50–56 in A. eximia (Table 1). The studied DP4 is remarkably different than that of A. eximia. It is shorter and less triangular than in A. eximia; the paracone's projection is stronger and the metacone is smaller than in A. eximia. A maxillary fragment with DP2-DP3 (MNHN-TRQ-1053) of a large hyaenid from Middle Sinap (Turkey) was described as Hyaena senyüreki and later transferred to Percrocuta (Howell and Petter, 1985, Ozansoy, 1965 and Schmidt-Kittler, 1976). It differs from the specimen studied in having more slender teeth (Table 1), a stronger protocone in the DP3, and a slightly smaller DP2. A fragment of the DP3 and a DP4 of a large hyaenid from Çandir, Turkey are described as Percrocuta aff. tungurensis (Schmidt-Kittler, 1976: p. 55; Figs. 49 and 50). The stronger protocone of the DP3, the stronger metacone of the DP4 and its smaller size (Table 1) distinguish the Çandir teeth from the ones studied.

The studied deciduous dentition is clearly larger than that of A. eximia and different from D. senyüreki. Two hyaenas larger than A. eximia are known from the Vallesian of the Axios Valley: D. salonicae, discovered in an unknown locality near the village of Diavata and D. gigantea, found at Pentalophos-1 PNT (Fig. 1), (Koufos, 1995). A direct comparison with these two species is not possible, but the large size of LGPUT-XIR-5 is an indication that it possibly belongs to one of these species. Based on these observations the studied specimen must belong to Dinocrocuta, but its specific determination is pending; hence it is referred to Dinocrocuta sp.

Family Felidae Fischer de Waldheim, 1817

Genus Metailurus Zdansky, 1924

Metailurus parvulus (Hensel, 1862)

Locality. Ravin de la Pluie (RPl), Axios Valley, Macedonia, Greece (Fig. 1).

Age. Late Vallesian, MN 10; GPTS = ∼9.3 Ma.

Material. Right maxillary fragment with C-M1, LGPUT-RPl-268; isolated right P4, LGPUT-RPl-267

Measurements. The measurements are given in Table 2.

Description: The dentition of LGPUT-RPl-268 is badly preserved, but the isolated carnassial LGPUT-RPl-267 is better preserved (Fig. 8A and B). The canine is broken in the crown, has lost the enamel layer, and cannot provide accurate measurements. Its root is elongated, flattened bucco-lingually, and relatively strong. Based on its size, the canine is quite strong for the size of the animal. The P3 is broken, lacks its lingual and mesial part, and bears a small and well distinguished p.a.c. The carnassial is elongated and relatively narrow with a strong parastyle in front of the paracone. Along its mesial border there is a small cingular projection, like a vestigial ectoparastyle (Fig. 8A₂). The protocone is broken in LGPUT-RPl-268 but it is well preserved in LGPUT-RPl-267 (Fig. 8A₂, B₃). It is a small cusp at the base of the paracone and far behind the mesial border of the parastyle. The metacone is blade-like, relatively low, and curved buccally. It is distinguished from the paracone by a deep buccal valley (Fig. 8A₃). The M1 is reduced, situated vertically to the P4; it has an elliptical occlusal shape and bears a small cusp, situated buccally (Fig. 8A₂).

Discussion: Metailurus parvulus is known from several Eurasian localities (NOW, 2011); it was originally described from Pikermi as Machairodus parvulus by a mandibular fragment with p3–p4 (Hensel, 1862: p. 568, Fig. 6). Later, it was recognized in Halmyropotamos, Chomateres and Samos (Forsyth Major, 1894, Koufos, 2009, Melentis, 1967 and Symeonidis, 1978). In Europe, it is known from the Late Vallesian locality of Montredon and in China from the Early Vallesian locality of Lantian-12 (NOW, 2011).

The two specimens from RPl have a dental morphology similar to that of M. parvulus from the various Greek localities. The elongated and narrow P3 with a clear p.a.c., the elongated upper carnassial with a small and low protocone situated well behind the mesial border of the parastyle, the large parastyle with a small mesial cingular projection like ectoparastyle and the very small M1 of the studied material are characteristic of M. parvulus. Although the canine of LGPUT-RPl-268 is badly preserved, its measured dimensions are closer to those of M. parvulus; the size of the other teeth from RPl is also similar to this species (Table 2). The comparison of the carnassial size indicates that M. parvulus is clearly distinguished from M. major and both studied specimens are situated in the ranges of variation of the previous species (Fig. 9). Hence, the studied specimens can be attributed with certainty to Metailurus parvulus. The studied material is the first evidence for its presence in the Vallesian of Greece and the wider eastern Mediterranean region.

Genus Machairodus Kaup, 1833

Machairodus giganteus (Wagner, 1848)

Locality. Ravin des Zouaves-5 (RZO), Axios Valley, Macedonia, Greece (Fig. 1).

Age. Early Turolian, MN 11; GPTS = ∼8.2 Ma.

Material. Right upper canine, LGPUT-RZO-205; left p4, LGPUT-RZO-240; left humerus, LGPUT-RZO-238; left calcaneum, LGPUT-RZO-239.

Measurements. Cs: 37.5 × 14.0, H_t = 178.7, H_m = 81.5, H_d = 99.0

p4: 30.2 × 12.7

The measurements of the postcranials are given in Table 3.

Description: The upper canine LGPUT-RZO-205 is almost complete, lacking only the end of its apex (Fig. 10A). It belongs to an adult individual, because its mesial border is quite worn. The canine is strongly curved backwards; its mesial curvature is larger than the distal one (Fig. 10B₅). The canine is also slightly curved buccally. Its lingual surface is flattened and the buccal one is convex. The distal border of the crown is serrated (Fig. 10A₃); the serrations of the mesial border have disappeared because of advanced wear. The root is very strong, wide and thick, corresponding almost to half of the total height. The studied p4 is badly preserved because it is crushed in the middle; its posterior root is also broken lingually (Fig. 10B). The tooth is elongated and narrow with three cuspids. The a.a.c. is strong and it is situated slightly lingually; it is surrounded by a strong buccal and mesial cingulum. The lingual part of the tooth is damaged and the cingulum is broken. The p.a.c. is weaker than the a.a.c. and it is situated on a strong distal cingular projection. The distal cingulum is worn but it seems to be strong and elevated. The roots are very strong and deep (Fig. 10B_1,2).

The humerus LGPUT-RZO-238 is almost complete, lacking the greater and lesser tubercles in its proximal part and the anterior part of the distal articular surface (Fig. 10C). The head of the humerus is well preserved (Fig. 10C_1,2), slightly curved and quite large; its mesio-distal diameter is 63.3 mm and the buccolingual 58.3 mm. The deltoid tuberosity is distinct and ends ∼160 mm from the head of the humerus. The diaphysis is strongly curved distally; in lateral view, the diaphysis starts to curve at about the end of the deltoid tuberosity (Fig. 10C_1,2). The transverse diameter of the diaphysis is tear-shaped. The coronoid fossa is wide, shallow and not connected to the olecranon fossa; there is no supratrochlear foramen. The olecranon fossa is deep and oval (Fig. 10C₃); its maximum dimensions are 43.4 × 38.8 mm. The medial epicondyle is strong and strongly projected laterally; above it there is a large supracondylar foramen (Fig. 10C₃). The lateral epicondyle is relatively weak and quite smaller than the medial one.

The calcaneum LGPUT-RZO-239 is well preserved and almost complete; the calcaneal tuberosity is slightly damaged and some parts are broken (Fig. 10D). The articular facet for the cuboid is rounded and its dimensions are 27.9 × 24.5 mm. The anterior talar articular surface is elongated and narrow with a sinus along its anterior border; its dimensions are 31.8 × 14.2 mm. The posterior talar articular surface is quite large and elliptical. The calcaneal sulcus is deep and wide, distinguishing well the anterior and posterior talar articular surfaces. The talar shelf is pointed and less projected. The calcaneal tuberosity, although broken, seems to be large with a deep sulcus.

Discussion: Two main large machairodonts are known from the Late Miocene of Europe: Machairodus aphanistus from the Vallesian and M. giganteus from the Turolian, which are well separated by several characters (de Beaumont, 1975). A third smaller taxon, named M. alberdiae, is only reported from Spain (Ginsburg et al., 1981). The upper canine, although it has similar morphology in the two main taxa, is more flattened in M. giganteus (Anton et al., 2004). The index L (= bucco-lingual diameter) × 100/B (= mesio-distal diameter) expresses the flattening of the canine (Table 4). Apart from the Charmoile specimen, which has a high index, the rest of the material of M. aphanistus group has similar index values to M. giganteus. There is great variation in the size of the upper canine in the two species, due to sexual dimorphism (Anton et al., 2004 and refs. cited). The mean size of the canine of the M. giganteus group is larger than the canine of M. aphanistus group (Fig. 11). The overlap between the two species is great and indicates the above-mentioned sexual dimorphism. The two available upper canines of M. alberdiae are close to the minimum values of M. aphanistus group and could represent another smaller taxon. The canine of M. giganteus group has a stronger curvature than that of the M. aphanistus group; both curvature angles of the studied canine are closer to those of the previous group (Fig. 10E). The dimensions of LGPUT-RZO-205 are closer to those of the M. giganteus group. They exceed the mean dimensions of M. aphanistus group, being near its highest values, and they are closer to the mean values of the M. giganteus group (Fig. 11).

The available p4 is directly compared with a cast of a mandibular fragment from the Axios Valley (MNHN-SLQ-936) that was described as M. aphanistus and later transferred to M. giganteus (Arambourg and Piveteau, 1929 and Koufos, 2000). The studied p4 is morphologically similar to that of MNHN-SLQ-936 with three cuspids and well developed distal cingulum; its size is also very close to that of MNHN-SLQ-936 (28.9 × 12.5 mm), confirming their similarity. The studied p4 is compared to a cast of M. aphanistus (NHML-EPP-M.49667a) from the type locality of Eppelsheim, (de Beaumont, 1975 and Pilgrim, 1931). The main cuspid of the p4 in NHML-EPP-M.49667a is broken but it seems to be high and strong. The a.a.c. is situated slightly lingually, as in the studied one but weaker. Likely the distal morphology of the tooth resembles that of LGPUT-RZO-240, but the p.a.c. is also weaker. The cingulum is strongly developed in both specimens. Pilgrim (1931) noted that the p4 has two p.a.c. He possibly considered the strong elevation of the distal cingulum as the second cuspid.

The known postcranial remains of Machairodus are rare and the comparisons limited. In the description of the Pikermi material (Gaudry, 1862–67) referred four humerus remains. He noted that they are large and the transverse diameter of the proximal articular facet is 87 mm, whereas the distal one is 90 mm. I found in the MNHN a complete humerus and a distal part of one from Pikermi. Their size and general shape are similar to those of LGPUT-RZO-238 (Table 3). Some remains of humeri from Pikermi are also described as belonging to M. giganteus (Roussiakis, 2002); the available dimensions of the distal part of humerus seem to be slightly smaller (Table 3). Some hind bones from Pikermi, described as “Epimachairodus”?taracliensis, possibly belong to M. giganteus (Pilgrim, 1931 and Roussiakis, 2002). Among them there is a calcaneum, whose dimensions are similar to those of LGPUT-RZO-239 (Table 3). Based on these similarities the studied postcranials from RZO could belong to M. giganteus.

M. giganteus is rare in the Axios Valley localities. It is recorded by a fragmentary skull and a mandibular fragment from Ravin-X (R-X), (Arambourg and Piveteau, 1929 and Koufos, 2000). A single metapodial in the MNHN collection is similar in size to this species; it is without locality indication but the type of fossilization resembles that of the Vathylakkos localities. The specimen from RZO is the only one found during our excavations in the Axios Valley since 1973. The species is also known from Samos (Forsyth Major, 1894 and Koufos, 2009). Personal observations in the old Samos collections, housed in NHML, NHMW and BSPM showed the presence of the species in the NHMW collection by a maxillary fragment and two upper canines. All are without locality indications and thus impossible to correlate them with the local stratigraphy. The species is also mentioned from the localities of Halmyropotamos and Kerassia, Evia Island (Melentis, 1967 and Roussiakis et al., 2006). All these Greek localities are dated from early to middle Turolian, MN 11-MN 12, (Koufos, 2006a). The genus Machairodus has a wide distribution, covering the whole of Eurasia from Spain to China (NOW, 2011) and thus it is a useful biostratigraphic marker.

As mentioned above, several articles have been published on the Axios Valley carnivores, describing the material found in the area since 1972. Recently a revision of the old and new material of the Axios Valley carnivores was given by Koufos (2000). In spite of forty years of field work and the great quantity of the collected fossils, the carnivore remains are few compared to other mammal groups. In this respect the Late Miocene fauna of the Axios Valley differs from those of Pikermi and Samos, which are comparatively rich in carnivore taxa (Koufos, 2006a). Although relatively poor, the carnivore fauna of the Axios Valley contains several taxa unknown from Pikermi and Samos, enlarging our knowledge of the Late Miocene carnivores of Greece. The studied carnivores, as noted in the introduction, originate from three different localities of the Axios Valley.

The Late Vallesian (MN 10) locality of XIR includes a poor mammal fauna. Two carnivore taxa, A. eximia and Protictitherium crassum, have been described earlier (Koufos, 2000). The new material indicates the additional presence of a giant hyaena belonging to Dinocrocuta after those of Pentalophos 1 and Diavata (Koufos, 1995 and Koufos, 2000). Although the age of XIR is clear (Late Vallesian, MN 10; ∼9.6 Ma), the age of the other two localities is questionable. The PNT fauna is quite peculiar, including some taxa different from those of XIR and RPl and indicating a possible older age than them; for this reason an Early Vallesian (MN 9) age is quite possible for PNT (Koufos, 2006a and Koufos, in press). The exact site of Diavata is unknown; except D. salonicae, the Diavata fauna includes some hipparion remains belonging to a form similar to the Vallesian/Early Turolian ones of the Axios Valley (Koufos, 1985). All these indicate that the dinocrocutas are certainly present in the Vallesian of Greece. However, they are absent from the Turolian faunas of the Axios Valley, as well as from the other Turolian faunas of Greece.

The available carnivore faunas from the three referred localities are quite poor (Fig. 13). Despite this, I have tried to make their guild structure diagrams and check whether there are indications for the palaeoenvironment similar to those provided by previously published results. The XIR carnivore fauna is poor, but its guild structure diagram (Fig. 12A) provides some information on the palaeoenvironment. Dinocrocuta is included in this diagram as a genus; its body mass and diet are considered similar to those of D. salonicae and D. gigantea, the two possible species to which the XIR specimen may belong. The XIR hyaenas belong to the large ones (30 ≥ 100 kg) and they are bone/meat eaters with a terrestrial way of life. Such forms characterize open modern faunas, like Serengeti (Morlo et al., 2010), as well as open fossil ones, like Samos, Pikermi (Nagel and Koufos, 2009). On the other hand the presence of Protictitherium crassum, a scansorial, insectivorous and small (3–10 kg) carnivore, is an indication for the presence of some higher vegetation (bushes, shrubs, small trees).

The earlier known carnivores from RPl were few, including A. e. leptoryncha, Protictitherium aff. gaillardi and Protictitherium aff. intermedium (Koufos, 2000). The study of the new material added three new taxa: E. wimani, ?Hyaenictis sp. and Metailurus parvulus. The felid M. parvulus is well documented in RPl and it is new for the Axios Valley fauna. Although it is well known in the Turolian faunas of Greece (Pikermi, Samos, Kerassia, Halmyropotamos), its presence in RPl is the first one in the Vallesian of Greece and southeastern Europe. The large mustelid E. wimani is rare in Eurasia and the single tooth found in RPl is the first evidence of its presence in the Balkan Peninsula. The third taxon found in RPl is ?Hyaenictis, represented by an isolated tooth very similar to H. graeca from Pikermi. This is the first indication of the possible presence of the genus in the Axios Valley and in the Vallesian of Greece and Eastern Mediterranean. The RPl carnivore fauna is richer than XIR one and its guild structure diagram is more representative (Fig. 12B). The RPl ?Hyaenictis is given in the diagram, using the characters of H. graeca from Pikermi (Fig. 13). The presence of large, terrestrial-cursorial, meat/bone-hypercarnivorous elements and the abundance of hyaenids are indications of an open environment. On the other hand, the two scansorial, insectivorous and small ictitheres suggest the presence of small trees, shrubs and bushes (Morlo et al., 2010 and Nagel and Koufos, 2009). A recent analysis of the Vallesian carnivores in the eastern Mediterranean suggests an open environment, similar to a modern open savannah with small trees and bushes, confirming the indications from the XIR and RPl guild structure diagrams. These indications also fit well with previous results for the Vallesian environment of the eastern Mediterranean and more precisely for the RPl suggested environment (Koufos and Konidaris, 2011 and refs. cited).

The new collection from RZO does not include any new carnivore taxon, but it offers several new pieces of evidence for the presence of Machairodus. The RZO carnivore fauna includes six species (Fig. 13) and its guild structure diagram is quite representative (Fig. 12C). The dominance of the hyaenids is clear in the diagram; five of the six taxa are hyaenids, and there is also a large-sized felid (Fig. 12C). The locomotor character of the carnivores, mainly terrestrial and cursorial, indicates an open environment, such as the modern Serengeti ones and the fossil ones of Pikermi and Samos. The carnivorous-meat/bone-hypercarnivorous diet is another indication of an open environment, whereas the small, insectivorous and scansorial Plioviverrops orbignyi is an indication of the presence of some small trees and bushes (Morlo et al., 2010). The RZO carnivore guild structure diagram is closer to that of Samos and Pikermi (Nagel and Koufos, 2009), as well as to the Turolian one of the Greco-Iranian Province (Koufos and Konidaris, 2011). All indications provided by the RZO carnivore guild structure diagram fit well with the previous results for the RZO and the Turolian eastern Mediterranean palaeoenvironment (Koufos and Konidaris, 2011 and ref. cited).