taxonID	type	description	language	source
03F887FBFF99FFF9FF3FF9A3FD9831E4.taxon	description	Family Cercopithecidae GRAY, 1821 Subfamily Colobinae BLYTH, 1863	en	Tsoukala, Evangelia, Nagel, Doris, Youlatos, Dionisios, Crégut-Bonnoure, Evelyne, Vlachos, Evangelos, Spassov, Nikolai (2024): Primates And Carnivores From Late Miocene And Earliest Late Pliocene Sites Of Macedonia, Northern Greece. Fossil Imprint 80 (2): 362-389, DOI: 10.37520/fi.2024.027
03F887FBFF99FFFBFF4DF961FB9E3227.taxon	description	Text-fig. 3 L o c a l i t y. Thermopigi. M a t e r i a l. Humerus left SIT 986, tibia right SIT 980, calcaneus left SIT 1100. D e s c r i p t i o n. The humerus retains a curved diaphysis, with proximal and distal metaphyses (Text-fig. 3 a, Tab. 1). It shows a high degree of modification or deformation due to neotectonic events, which is quite common – it has been observed in several bones of large mammal remains from Thermopigi. Epiphyses are missing, likely due to the young juvenile (neonate?) age of the individual. The diaphysis shows a curvature in the middle and torsion of a relatively large cercopithecid. The maximum height of the chord from the axis of the bone is 15.22 mm. The distal portion is slender, whereas the proximal portion, particularly the collum area, is robust. The tibia retains the distal end, with a small part of diaphysis (Text-fig. 3 b, Tab. 1). The epiphysis was not fused, and a suture is visible around the entire perimeter of the distal end. The part of the shaft appears relatively robust, is posteriorly rounded and anteriorly flat. The anterior surface is partly eroded. The malleolus tibiae is well developed with a sub-triangular facies articularis malleoli. The facies articularis distalis tibiae is sub-square but narrow, whereas the facies articularis malleoli fibulae is semicircular and wide. The cross-section of diaphysis at 40 mm from distal end is relatively rounded. The calcaneus is complete, and well preserved (Text-fig. 3 c, Tab. 1). It is robust, with a slightly concave squarishshaped tuber below it, and laterally, a well-marked fossa. The lateral side of the calcaneal body is eroded, and the posteromedial and disto-medial edges of the sustentaculum tali are broken. From the lateral and posterior view, the sustentaculum tali curves anteriorly and medially respectively in relation to the manubrium. The articular surface for the cuboideum is broad and hollow. Detailed descriptions of the calcaneus can be found in Youlatos et al. (in press). D i s c u s s i o n. The morphology of the humeral shaft implies a rather pronounced deltopectoral ridge, although it is difficult to judge, as it is rather incomplete. The shaft appears relatively robust and seems to be mediolaterally compressed. This morphology is reminiscent of more terrestrial Macaca LACÉPÈDE, 1799 species. All these characters suggest quadrupedal adaptations related to terrestrial activities (Harrison 1989). The tibial morphology is similar to most terrestrial cercopithecids (e. g., Papio DESMAREST, 1820), but different from the more medially expanded shafts of more arboreal colobines (DeSilva et al. 2010). The medial malleolus is moderately developed, triangular, and standing at an almost vertical angle to the trochlear facet. The overall morphology is similar to that of most terrestrial cercopithecids like Papio and differs substantially from the medio-laterally robust and protruding malleolus of hominoids (DeSilva et al. 2010). The malleolus bears a slightly concave articular surface, ending distally at a pronounced ball-shaped area, a feature typical to cercopithecids, but absent in hominoids (Harrison 1989). Although the antero-distal part of this end is broken, the bulbous malleolar end is reminiscent of that of more terrestrial cercopithecines (e. g., Papio, Theropithecus I. GEOFFROY, 1843). This condition is different from that observed in the more arboreal colobines and from that in Paradolichopithecus NECRASOV et al., 1961 from Vatera (Sondaar et al. 2006). In distal view, the antero-posterior axis of the malleolus is set parasagitally, a condition observed in more semi-terrestrial and terrestrial cercopithecines (e. g., Macaca, Theropithecus), and is different from the medial tilt of most arboreal colobines (e. g., Colobus ILLIGER, 1811) (Youlatos 1994). The postero-distal part of the epiphysis and the malleolus are slightly damaged, and the groove that accommodates the tendons of the posterior tibialis and the long digital flexor is not visible (Ford 1986). In this character, SIT- 980 is different from Mesopithecus pentelicus, where the groove is deep (Ingicco 2008), but is similar to most terrestrial cercopithecids. On the distal surface, the trochlear facet has a trapezoid shape, is smoothly concave, and mediolaterally narrow. The latter morphology is typical of cercopithecids (Harrison 1989), and more particularly that of more terrestrial forms (Laird et al. 2018). The facet bears an underdeveloped, relatively concave median antero-posterior ridge. The ridge is slightly perpendicular to the medio-lateral plane of the bone, is bordered by shallow medial and lateral depressions for articulation with the trochlear surface of the talus, and ends anteriorly to a V-shaped tubercle. The protuberance of the tubercle is reminiscent of that in Macaca and Theropithecus, and differs from that of more arboreal colobines (Youlatos 1994, Laird et al. 2018). The posterior border of the trochlear facet is smooth and relatively low. The anterior border appears sharper and relatively higher, and bears the anterior tubercle. On the lateral side of the bone and towards the anterior part of the trochlear facet lies the relatively shallow crescent of the fibular facet. This morphology is similar to that of cercopithecids. Overall, these characters of the distal tibia suggest terrestrial quadrupedal activities. The calcaneal features indicate a typical cercopithecid calcaneus, morphologically different from extant and fossil hominoids (Rose 1986, Strasser 1988). The proximal calcaneo-astragalar facet is relatively short, squared off, tightly curved, and tilts obliquely and faces more dorsally, whereas fossil and extant hominoids possess longer, gently curved, and acutely tilted facets (Langdon 1986, Rose 1986, Strasser 1988, Gebo 1989). Additionally, there is a distinct pressure facet for the fibula-calcaneal ligament on the lateral side of the proximal calcaneo-astragalar facet, a feature absent in hominoids (Strasser 1988). More distally, the middle and distal calcaneo-astragalar facets on the sustentaculum tali are discontinuous and acutely oriented relative to each other. This condition is different from the confluent facets of fossil and extant hominoids (Rose 1986, Strasser 1988). The middle facet is almost flat, with a disto-medial orientation, whereas the distal one is also flat and adjacent to the calcaneocuboid facet (Ford 1988). At the distal end, the calcaneocuboid joint is dorso-plantarly low and relatively flat, with a very shallow medial pit, but not rounded and deeply concave, as in most hominoids (Rose 1986). These features support the attribution of SIT- 1100 to cercopithecids. The overall size of the calcaneus appears larger than that of M. pentelicus from Pikermi, and is more like that of large colobines, such as Semnopithecus DESMAREST, 1822 and Pygathrix LINNAEUS, 1771. However, as calcaneal features that distinguish between the two subfamilies demonstrate considerable variation, related to both phylogenetic and ecological constraints, it is difficult to classify this calcaneus at a subfamilial level. The calcaneal body is relatively robust and high, and is similar to that of extant colobines, and to a lesser extent to Mesopithecus. It is generally straight, a condition more reminiscent of terrestrial cercopithecines (Pina et al. 2011). On the dorsal surface of the calcaneal body lies the proximal calcaneo-astragalar facet. In SIT- 1100, the facet is quite low, relatively narrow (like in Erythrocebus SCHREBER, 1775 and Semnopithecus), comparatively short, and tilts disto-medially, similar to that of Papio and Erythrocebus. It is generally reminiscent of the morphologyofsemi-terrestrialandterrestrialcercopithecines. Finally, on the dorso-lateral side of the facet, the pressure facet is smooth and short, but relatively wide. This condition is more similar to that of more terrestrial cercopithecids. The proximal region of the calcaneus is relatively high and curves medially, and its relative size seems intermediate between Papio, Erythrocebus, and Semnopithecus and the more arboreal colobines (e. g., Colobus) and cercopithecines (e. g., Cercopithecus LINNAEUS, 1758). On the lateral side near the proximal end of the calcaneus, SIT- 1100 possesses a marked relatively deep fossa that accommodates the calcaneofibular ligament. This condition is similar to that of Mesopithecus and the terrestrial cercopithecines, and probably indicates a strong ligament that counteracts lateral tensions and transversely stabilizes the foot. At the proximal end of the calcaneus, the tuber calcanei, where mm. triceps surae insert, is very wide and relatively low. It ends dorsally on a rounded surface and proximally forms an extended, lightly concave surface. This morphology is similar to that of Mesopithecus from Pikermi and that of more terrestrial (e. g., Erythrocebus) and semi-terrestrial (e. g., Chlorocecus J. E. GRAY, 1870, Cercocebus É. GEOFFROY, 1812) cercopithecines. Compared to Mesopithecus pentelicus from Kryopigi, the SIT- 1100 calcaneus has a more squarish tuber than that of KRY, which is distinctly smaller in size and has an oval-shaped tuber, in proximal view. In SIT- 1100, the distal region of the calcaneus appears relatively long and morphologically reminiscent of the calcanei of semi-terrestrial Macaca. On the distal and medial part of the calcaneus, the sustentaculum tali, which accommodates the middle and distal astragalo-calcaneal facets, is relatively short and narrow. The middle facet faces medio-plantarly and is relatively flat and relatively steeply inclined. The distal facet is also steeply set and is relatively flat. The overall morphology is similar to that of most terrestrial and semi-terrestrial cercopithecines (e. g., Macaca). The surface between the facets lacks the prominent fossa observed in some calcanei of Mesopithecus, indicating a weak astragalar neck ligament (Lewis 1989, Youlatos 1990). On the plantar side of the sustentaculum, the wide and very shallow groove that runs on the latero-plantar side of the calcaneal body suggests a less developed hallucal flexor. On the distal end of the calcaneus, the calcaneocuboid facet of SIT- 1100 is quite flat and relatively rounded, with a very shallow medial pit. This morphology is similar to that of Mesopithecus and terrestrial cercopithecines (e. g., Papio, Theropithecus I). The facet appears slightly symmetrical, similar to Papio and Semnopithecus, when compared to the highly asymmetrical facets of more arboreal colobines (e. g., Pygathrix). Moreover, on the plantar side of the calcaneocuboid facet, there is a quite well-developed plantar tubercle, similar to that observed in more terrestrial cercopithecines, such as Papio and Theropithecus. A well-developed tubercle indicates a strong calcaneocuboid ligament that stabilizes the joint (Langdon 1986). Overall, these characters most likely suggest quadrupedal adaptations on more stable, terrestrial substrates. In Greece, Mesopithecus delsoni was established for the first time in Ravin des Zouaves 5, Axios Valley, Macedonia (type locality), of Late Miocene age and biozone MN 11 (Bonis et al. 1990). In addition, probable occurrences M. cf. delsoni are reported at Perivolaki (MN 12; Koufos 2006 b) and Ravin-X in Axios Valley (MN 11; Koufos 2022 d). In the revision, Koufos (2022 d) referred to Mesopithecus sp. at Nikiti 2 in Chalkidiki (MN 11), and in Perivolaki and Vathylakkos (MN 12). On the other hand, Mesopithecus pentelicus is established at Pikermi in biozone MN 12, as well as at Chomateres (Attica), Dytiko 1, 3 (Axios Valley) and Kryopigi (Chalkidiki). Therefore, based on the presence of this early form of Mesopithecus and the rest of the associated fauna, the stratigraphic age of the Thermopigi locality can be estimated between late MN 11 and pre-Pikermian MN 12 biozones.	en	Tsoukala, Evangelia, Nagel, Doris, Youlatos, Dionisios, Crégut-Bonnoure, Evelyne, Vlachos, Evangelos, Spassov, Nikolai (2024): Primates And Carnivores From Late Miocene And Earliest Late Pliocene Sites Of Macedonia, Northern Greece. Fossil Imprint 80 (2): 362-389, DOI: 10.37520/fi.2024.027
03F887FBFF97FFF0FF2FFF3AFEDD3331.taxon	description	Text-fig. 7 L o c a l i t y. Promachonas. M a t e r i a l. i 2 right PRC 104, p 4 left PRC 103, p 4 frag. right PRC 106, m 1 ant. frag. left PRC 102, m 2 left PRC 101, m 2 right PRC 100, m 3 frag. left PRC 105. The tooth material is probably from one mandible. D e s c r i p t i o n. The lower second incisor is well-preserved (Text-fig. 7 d, Tab. 3), slightly worn, with about half its single root length. The bicuspid fourth lower premolar PRC 103 (Text-fig. 7 c) is complete with two roots: the mesial one is approximately 9 mm long, while the distal one is broken. The PRC 106 fragment (Text-fig. 7 g) retains the anterior half of the tooth with the talonid missing. The metaconid and the protolophid are well developed in both teeth (Tab. 3). The lingual cuspid is larger than the labial. The trigonid basin is well distinct and relatively deep, whereas the talonid basin is broad. The first lower molar fragment PRC 102 (Text-fig. 7 e) is slightly worn and preserves only the anterior lobe with well-developed metaconid (Tab. 3). The second lower molars PRC 101 (Text-fig. 7 a) and PRC 100 (Text-fig. 7 b) are bilophodont and slightly worn. The metaconid is distinctively higher than the hypoconid (Tab. 3). The hypo-, para- and metacristae are intense. There is an anterior cingulum. The former also preserves both roots, whereas the latter is about half length of the two roots. The transverse crests are well developed. The third lower molar fragment PRC 105 preserves only the talonid (Bm 3 tld = 4.83 mm) with the well-developed hypoconulid (H = 4.5 mm) (Text-fig. 7 f), half hypoconid and the well-developed entoconid (H = 5.9 mm). According to the description above, the teeth could belong to the same individual, therefore the MNI seems to be 1. D i s c u s s i o n. The primate material from Promachonas (PRC) consists of isolated, complete, or partial teeth. In the broader area of Serres, in Maramena of biozone MN 13 / 14, south of PRC, the cercopithecid Mesopithecus pentelicus was initially identified through teeth (Küllmer and Doukas 1995), which was later attributed to Mesopithecus sp. (Koufos 2009 b). There is a marked difference in the heights of the labial and lingual cuspids of the lower molars between Mesopithecus (KRY) and Dolichopithecus (Megalo Emvolo MEV and PRC), with the lingual being more pointed and higher than the buccal in the latter specimens. The most significant remains of the PRC monkey are the two second lower molars. Their proportions are typical of lower molars. They are slightly larger than those of Mesopithecus delsoni from Hadjidimovo (late MN 11), Bulgaria (Bogdanova et al. 2023); even one of the three male individuals from Hadjidimovo sample practically reaches their size (see Koufos et al. 2003). At the same time, the cuspids of the PRC specimen seem to be more similar to those of Dolichopithecus: more pointed (higher and mesio-distally narrower) than those of Mesopithecus, and their notches are deeper. In terms of size, they differ slightly from the m 2 of the type specimen of Dolichopithecus balcanicus from Tenevo. The cuspids of the m 2 from PRC do not appear to be as elongated as in the latter specimen, but this is possibly due to some extent to the greater individual age of the individual. The PRC teeth are slightly smaller and shorter than those of MEV, whose cuspids are also more pointed. Compared to the tooth dimensions of Dolichopithecus from the Pliocene of Pridnestrovie (Mashchenko and Marareskul 2011), Megalo Emvolo (Koufos et al. 1991) and Tenevo, as well as to the morphology of the Tenevo Dolichopithecus (Spassov and Geraads 2007), the PRC is tentatively attributed to D. balcanicus from the Latest Pliocene of Bulgaria, which is distinguished from the common European species, D. ruscinensis by its smaller size, much shallower mandibular corpus and marked congestion of the premolars (Spassov and Geraads 2007). In Greece, Pliocene cercopithecids are mainly reported from cranial material of the middle Villafranchian, biozone MN 17, from Vatera, Lesvos Island [Paradolichopithecus arvernensis (DEPÉRET, 1929)] (De Vos et al. 2002, van der Geer and Sondaar 2002, Lyras and van der Geer 2007), Karnezeika, Peloponnese (cf. Paradolichopithecus sp.) (Sianis et al. 2022), and Dafnero, Kozani (P. aff. arvernensis) (Kostopoulos et al. 2018). Earlier cercopithecids are reported from the Ruscinian biozone MN 14 – 15 in the Ptolemais basin (Dolichopithecus ruscinensis) (Doukas and De Bruijn 2002) and Megalo Emvolo (Koufos et al. 1991), the latter tentatively included in D. balcanicus (Spassov and Geraads 2007). The find from Promachonas represents possibly the same geologically younger Balkan Dolichopithecus species, which is distinguished from the common European species, D. ruscinensis by its smaller size, much shallower mandibular corpus and congestion of the premolars (Spassov and Geraads 2007).	en	Tsoukala, Evangelia, Nagel, Doris, Youlatos, Dionisios, Crégut-Bonnoure, Evelyne, Vlachos, Evangelos, Spassov, Nikolai (2024): Primates And Carnivores From Late Miocene And Earliest Late Pliocene Sites Of Macedonia, Northern Greece. Fossil Imprint 80 (2): 362-389, DOI: 10.37520/fi.2024.027
03F887FBFF90FFF2FF16FA97FECF3F56.taxon	description	Text-figs 8, 9 M a t e r i a l. Thermopigi: Skull with P 2 – M 1 left SIT 1421; 2 P 2 right SIT 531 and left 940, P 3 left SIT 1057, mandible with p 2 – m 1 fr. left SIT 713 + 714; m 1 right SIT 529, anterior pad, d 4 fr right SIT 1280, p 3 right SIT 530; coprolite SIT 1359. Platania: A maxilla with the right and left P 2 – P 4, PLD 464. D e s c r i p t i o n. Thermopigi: The corroded skull SIT 1421 bears the very poorly preserved left tooth row P 2 – P 4 and M 1, nasals, nasal cavity, infra-orbital foramina, frontal with the orbital cavities and ecto-orbitals, post-orbital constriction to the parietals, most of the left zygomatic arch, and a palate showing a deep depression below the metastyle of the left P 4, in front of the molar, followed by the choanae (Text-fig. 8 a 1 – 3). The toothrow is damaged (Text-fig. 8 a 4); only traces remain that allow us to make a few measurements (Tab. 4). There are traces of the mesial accessory cusp of the P 2 and P 3, and the metastyle of the upper carnassial. The worn and slender M 1 rotates 90 degrees inwards; paracone, metacone and protocone are barely distinguishable. There is palatal asymmetry, on the right side of the maxilla, where paleopathological evidence indicates a type of exostosis bearing the roots of P 2 and P 3. Of the two isolated P 2 s, SIT 940 retains only the upper part of the unworn crown, the pointed tip and the two well-defined crests (Text-fig. 8 e). The preserved dimensions are: length 12 mm, breadth 8 mm. The P 2 SIT 531 is almost complete and only a little worn (Text-fig. 8 d). Two sharp anterior and posterior crests extend from the tip of the tooth, the latter meeting the well-distinct posterior accessory cusp (Tab. 4). The P 3 SIT 1057 is almost complete, missing only the mesial part with the accessory cusp (Text-fig. 8 c). The main cusp is unworn and strong with two sharp crests extending from the tip anterior and posterior of the tooth. The latter terminates in the well-developed posterior accessory cusp; dimensions in Tab. 4. The left mandible SIT 713 + 714 preserves the corpus with p 2 - m 1 (Text-fig. 8 b 1 – 3). The slightly worn tooth row consists of the complete p 2, p 3, the talonid of the damaged p 4, and the mesial half of the lower carnassial. The alveolus of the single root p 1 is also preserved (Depth = 4.45 mm), as well as parts of the symphysis. The corpus is relatively high and straight at its preserved inferior border. There are two mental foramina below p 2: the larger below the middle and the smaller below the posterior border of the tooth. The p 2 is short but robust, with a small anterior and a larger posterior accessory cuspid. There is a lingual cingulum. The larger p 3 bears a strong anterior and a posterior accessory cuspids on the mesio- and distolingual sides of the tooth. There is a faint buccal cingulum. The p 4 is damaged, and only the well-developed distal accessory cuspid is preserved on a large, raised base of the distal cingulum around the distal area of the tooth. The mesial part of the lower carnassial is preserved with a mesial cingulum at the crown base, whereas only the root of the distal part of the tooth remains. Dimensions of the mandible and teeth are in Table 4. From the lower deciduous carnassial d 4 SIT 1280, only part of the blade is preserved (Text-fig. 8 h). It is slightly worn. The paraconid and protoconid form an angle of 115 ° at 13 mm between their peaks. The preserved dimensions are a length of 15.73 mm and breadth of 6.0 mm. The nearcomplete p 3 SIT 530 is well-preserved and unworn. The protoconid is relatively high and the distal crest is sharp. The mesiolingual accessory cuspid is weak, while the distolingual one is larger (Text-fig. 8 f). Dimensions are in Tab. 4. From the carnassial SIT 529, a half-fragment bears the unworn paraconid of a juvenile. The mesiolingual crest is strong. The preserved dimensions are a length of 13.10 mm and breadth of 10.05 mm (Text-fig. 8 g). The well-preserved coprolite SIT 1359 shows a typical constriction at one third of its total length and a shallow depression at one of its ends (Text-fig. 8 i). Dimensions are length 47.31 mm, breadth 37.08 mm and height 24.49 mm. Platania: Maxilla PLD 464 preserves both right and left tooth rows with P 2 – P 4, nasal cavity and the left infraorbital foramen (ca. 6.5 mm) above and medial to the P 3 roots (Text-fig. 9 a – c). Both tooth rows are well-preserved; only the protocone of the right carnassial is missing. The highcrowned tooth rows are slightly worn and with lingual cingula. P 2 is oval shaped with a palatal cingulum, with sharp anterior and posterior crests, a vestigial anterior and a small posterior accessory cusp. P 3 is large, with sharp mesiopalatal and distal crests, a small anterior and a larger posterior accessory cusp. The anterior part of the tooth is broader than the posterior. The upper carnassial is large, long, and robust. The parastyle is well-developed, the paracone high and the metastyle forms a long blade. The protocone is relatively weak, confined to the base of the tooth, posteriorly to the anterior border of the parastyle, typical for Adcrocuta (dimensions in Tab. 4). D i s c u s s i o n. The PLD maxilla is described and discussed here to compare the two richest Turolian mammal sites, located close to each other in eastern Macedonia: Thermopigi and Platania. Thermopigi Adcrocuta (SIT) appears less robust than Platania (PLD) hyena. They generally fall within the variation between Axios Vallesian and Axios Turolian (Text-figs 10, 11). When comparing SIT and PLD maxillae, it should be noted that (a) longer tooth row (even damaged) P 2 – P 4 and carnassials in SIT than in PLD, (b) index B / LX 100 for P 2: SIT = 56.16, PLD = 71.10, (c) P 3 higher but slenderer in SIT than in PLD and the B / HX 100 index: SIT = 71.35, PLD = 84.53. The morphology of the Platania P 2 – 3 (relatively narrow, with a strong mesial additional cusp and a strong lingual cingulum) indicates an earlier geological age of the Adcrocuta from Platania, and places it apparently in the pre-middle Turolian stage of premolar evolution of this hyena, close to some other Balkan samples from the Vallesian / Turolian boundary (Spassov et al. 2018, 2019). This morphology seems to support the above-assumed biochronological position of the locality. Also, it is noteworthy to compare these two faunal associations of Thermopigi and Platania with Adcrocuta, as already discussed in earlier publications on rhinoceroses (Tsoukala 2018), proboscideans (Konidaris and Tsoukala 2020), bovids (Vasileiadis et al. 2019), turtles (Vlachos and Tsoukala 2014) and the paleoenvironment. The bite marks of Adcrocuta appear comparatively more often in the bone-remains of Platania than in those of Thermopigi. Especially in hipparions, young rhinos and bovids, there is a common presence of bitten and broken bones, presumably making it a feeding site for this top carnivorous scavenger. Adcrocuta eximia upper teeth compared Adcrocuta eximia lower teeth compared In Greece, Adcrocuta eximia is widespread and known from at least 20 sites dating from the late Vallesian to the late Turolian (Koufos 2022 b). The dental dimensions are close to those of Adcrocuta eximia from sites of Northern Greece, such as Axios Valley (late Vallesian, MN 10), Nikiti (early Turolian, MN 11) (Koufos 2000, 2012, 2016) and Kryopigi Kassandra (MN 12 / MN 13) (Lazaridis 2015). It must be noted that the upper carnassials are more robust in the SIT and PLD specimens than those of the other sites (Text-figs 10, 11).	en	Tsoukala, Evangelia, Nagel, Doris, Youlatos, Dionisios, Crégut-Bonnoure, Evelyne, Vlachos, Evangelos, Spassov, Nikolai (2024): Primates And Carnivores From Late Miocene And Earliest Late Pliocene Sites Of Macedonia, Northern Greece. Fossil Imprint 80 (2): 362-389, DOI: 10.37520/fi.2024.027
03F887FBFF8EFFEFFF0DFAE5FD6D33A8.taxon	description	Text-fig. 14 L o c a l i t y. Milia. M a t e r i a l. Tibia distal left MIL 974. D i s c u s s i o n. This distal tibia represents a felid. This specimen is smaller than that of the scimitar-toothed cat Homotherium FABRINI, 1890, which is one of the three carnivores from the MN 16 a mammalian assemblage of Milia (H. crenatidens (WEITHOFER, 1889), Ursus etruscus CUVIER, 1823, and Agriotherium sp.; Tsoukala et al. 2014). In addition, its intermalleolar notch is deeper than that of Homotherium (Ballesio 1963). The other scimitar-toothed cats, present during the Late Pliocene, are Dinofelis ZDANSKY, 1924 and Megantereon. In Dinofelis, the medial side is more concave than in the Milia tibia, because the medial malleolus is medially more prominent, and the line passing through the base of the medial malleolus and the lowest point of the central malleolus is less oblique (Werdelin and Lewis 2001: fig. 21). Concerning Megantereon cultridens (CUVIER, 1824), the distal articulation shows some similarities, but the distal width is greater than that in the Milia specimen (Christiansen and Adolfssen 2007). The Villafranchian Acinonyx pardinensis (CROIZET et JOBERT, 1828) has a shorter central malleolus and an intermalleolar notch less deep with, in cranial view, the inflection point of the notch’s curve in a more medial position (Hemmer 2001: pl. 137). The Milia tibia could belong to a felid with a size close to that of Puma pardoides (OWEN, 1846). This species could be a good candidate. The dimensions are of the same order as the individuals from La Puebla del Valverde (Madurell-Malapeira et al. 2010) and Untermassfeld (Hemmer et al. 2004), two localities that are more recent than Milia (Tab. 6). However, their morphology is not exactly the same: cranially, the inflection point of the intermalleolar notch’s curve has a rather more medial position than in Milia, and it is similar to the current P. concolor (LINNAEUS, 1771). The cochlear grooves of the Milia tibia are more asymmetric. Consequently, the attribution of the Milia tibia is under question. We are sure that it cannot belong to a Homotherium, Dinofelis, or a cheetah. This single and incomplete specimen does not provide enough material to conclude with certainty that it belongs to a Megantereon female (the Senèze skeleton analysed by Christiansen and Adolfssen (2007) is undeniably a male) or a new species of Puma. In any case, regardless of the genus, this would most likely represent the oldest record in Eurasia: previously, the first appearance of Megantereon dated from the early Villafranchian (middle Pliocene; Berta and Galiano 1983) (first known in Europe probably in Les Étouaires; see Hemmer and Kahlke 2022), and that of Puma is at ca. 3.1 Ma in Asia (Mongolia) and around 2.8 Ma in Europe, i. e., MN 16 b (also the French locality of Les Étouaires; Hemmer et al. 2004, Nomade et al. 2014). For the time being, it is more reasonable to leave the nomenclature open.	en	Tsoukala, Evangelia, Nagel, Doris, Youlatos, Dionisios, Crégut-Bonnoure, Evelyne, Vlachos, Evangelos, Spassov, Nikolai (2024): Primates And Carnivores From Late Miocene And Earliest Late Pliocene Sites Of Macedonia, Northern Greece. Fossil Imprint 80 (2): 362-389, DOI: 10.37520/fi.2024.027
03F887FBFF8FFFE9FF35FA1DFDDE3188.taxon	description	Text-figs 15, 16 L o c a l i t y. Thermopigi. M a t e r i a l. Cranium with right I 1 – I 3, C and left, P 1, P 2, P 3, P 4 SIT 1494. D e s c r i p t i o n. The skull is deformed and compressed due to neotectonic activity in the area, but bears most of the teeth in good condition. It shows a short rostrum, and preserves the nasal and frontal bones, the anterior part of the zygomatic arch, the right orbit, and the small infraorbital foramen located above the anterior root of P 4 and the protocone (Text-fig. 15 a, d, e). There is a small diastema (ca. 3 mm) between I 3 and C. The posterior part of the skull is missing, as well as the sagittal crest. The almost unworn dentition and the half-raised right P 3 suggest a young individual. All incisors are well-preserved and show V-shaped palatal cingula. In addition, the much larger I 3 s exhibit slight torsion. The robust canines may indicate a male individual. They show wrinkled enamel at their base, and two well-defined anterior and posterior longitudinal crests. The P 1 is a small, single-rooted tooth with the main cusp, and a distinct basal cingulum well-developed. The P 2 is doublerooted, has a single main cusp and a crest extending along its midline, weaker anteriorly and more distinct posteriorly, terminating on a small tubercle on the posterior margin of the low, relatively distinct cingulum (Text-fig. 15 b). The P 3 is half-raised, with a strong main cusp and a basal cingulum. The upper carnassial, P 4, is typically robust and three- a: palatal view, b: details of teeth – occlusal view, c: details of P 4, M 1, d: cranial view, e: right lateral view. Scale bars 20 mm. rooted. The parastyle is weak but distinct, and includes a small, conical parastylar cusp. The paracone is broad and bears a pair of low crests along the anterior surface; the medial crest terminates just to the parastylar cusp, posterior to the parastyle. There is a relatively deep inflection of the anterior portion of the P 4, located between the parastyle and protocone. The protocone is low and extends anteromedially from the base of the paracone, projecting nearly as far forward as the parastyle. The metastyle is mesially broad, but tapers distally. As regards the cingulum, the condition of the specimen does not allow any description. The paracone on the M 1 is externally rounded, large (5.8 mm × 6.4 mm) and dominates over the metacone (3.8 mm × 4.3 mm). Both paracone and metacone are relatively short. The inner lobe is broad, expanding posterolingually to form a broad talon, with a slight constriction. Protocone and paraconule are joined by a pronounced crest. A cingulum surrounds the entire lingual part of the molar, and buccally there is a platform, more developed in the paracone area. It is further bounded by a well-developed cingulum, which in its union with the paracone forms a rudimentary parastyle (Text-fig. 15 c). Dimensions of skull are in Table 8. D i s c u s s i o n. Many studies have described the expansion, evolution and extinction of Plesiogulo. The giant wolverine was the largest terrestrial mustelid, originating in Asia and migrating to North America between 7.0 and 6.5 Ma ago (Kurtén 1970, Hendey 1978, Harrison 1981, Alcalá et al. 1994, Haile-Selassie et al. 2004, Samuels et al. 2018 and references therein). It is known from several Middle to Late Miocene and Pliocene sites across Eurasia, North America, and Africa, and has long been discussed as a possible relative of Gulo. The earliest occurrence in Europe is from the Middle Miocene (MN 6, ca. 15.2 – 12.5 Ma) (Samuels et al. 2018). There is a question if Plesiogulo crassa was ecologically replaced in the Pliocene by the leopard-sized wolverine Plesiogulo monspessulanus VIRET, 1939 (Montpellier, MN 13), as some of its characters are intermediate between those of the Late Miocene form and * Koufos 2006 c, ** Kurtén 1970, Alcalá et al. 1994	en	Tsoukala, Evangelia, Nagel, Doris, Youlatos, Dionisios, Crégut-Bonnoure, Evelyne, Vlachos, Evangelos, Spassov, Nikolai (2024): Primates And Carnivores From Late Miocene And Earliest Late Pliocene Sites Of Macedonia, Northern Greece. Fossil Imprint 80 (2): 362-389, DOI: 10.37520/fi.2024.027
