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Palaeontology of the upper Miocene vertebrate localities of Nikiti (Chalkidiki Peninsula, Macedonia, Greece)

Artiodactyla§ Dimitris S. Kostopoulos Laboratory of Geology and Palaeontology, Department of Geology, Aristotle University of Thessaloniki, 54124 Thessaloniki, Greece

A R T I C L E I N F O

A B S T R A C T

Article history: Received 5 November 2014 Accepted 19 January 2016 Available online xxx

The artiodactyl assemblage of the upper Miocene fossil locality Nikiti 2, Northern Greece, is rediscussed in the light of new data provided by a second, five-year long round of fieldwork. The significantly enriched artiodactyl material allows revising and updating previous systematic interpretations. Along with the previously recorded Helladotherium duvernoyi, Gazella cf. capricornis, and Nisidorcas planicornis, two more giraffids (Palaeotagus rouenii, and Palaeotraginae indet.) and two additional bovid taxa (Gazella pilgrimi, and Palaeoryx cf. pallasi) are recognized. Bovid material previously referred to Tragoportax aff. rugosifrons is reinterpreted here as partly belonging to T. amalthea, and partly to a second boselaphine taxon close to Miotragocerus. Furthermore, the medium-sized spiral horned antelope originally referred to as cf. Ouzocerus is now attributed to a primitive morphotype of Palaeoreas lindermayeri. As a whole, the updated artiodactyl assemblage from Nikiti 2 includes three giraffid and seven bovid taxa. According to local biochronological evidence and the primitiveness of both Palaeoreas and Nisidorcas, an early Turolian age is suggested for the Nikiti 2 fauna. ß 2016 Elsevier Masson SAS. All rights reserved.

Keywords: Giraffidae Bovidae Mammalia Taxonomy Late Miocene Greece

1. Introduction The fossiliferous site of Nikiti (Koufos et al., 1991) is located at the northernmost corner of the Sithonian branch of the Chalkidiki peninsula, N. Greece. The site is placed into the upper part of the homonymous lithostratigraphic formation of sands, sandstones and red-beds outcropping ENE of the Nikiti village (Koufos, 2016). The site includes four fossil localities with Nikiti 1 (NKT) and Nikiti 2 (NIK) being the most productive in terms of density and completeness of fossil mammal remains. Stratigraphically, NIK stands ca. 20 m above NKT and within the same alternations of reddish silty sands with gravel intercalations (Kostopoulos and Koufos, 1999: fig. 1; Koufos, 2016). An age between 9.3 and 8.7 Ma has been proposed for NKT (Koufos, 2006), whereas Kostopoulos (2009a) correlated NIK to the lower part of the early Turolian, with an estimated age between 8.7 and 8.0 Ma. Previous systematic works on the mammal remains from NKT and NIK focused on the sharp taxonomic differences between these two faunal assemblages that tightly frame the Vallesian/Turolian boundary in N. Greece (e.g., Koufos, 2006) and the hominoid/cercopithecine turnover (Andrews et al., 1996). Two rounds of systematic excavations took place in Nikiti. The first one lasted six years starting with the discovery of the NKT locality in 1990 (Koufos et al., 1991). Giraffid and bovid material discovered at that time from NIK were described by Kostopoulos §

Corresponding editor: George D. Koufos. E-mail address: [email protected]

et al. (1996) and Kostopoulos and Koufos (1999), respectively. According to these authors, the artiodactyl assemblage of NIK includes: Helladotherium duvernoyi, Tragoportax aff. rugosifrons, Gazella aff. capricornis, Nisidorcas planicornis, and cf. Ouzocerus sp. A second round of five years of excavations (2005–2009) focused on the NIK locality and greatly improved its fossil record. Numerous new specimens of giraffids and especially bovids, including fairly complete skulls and postcranials in natural articulation were unearthed at this time. These new findings are discussed here; they allow an overall reappraisal of the artiodactyls of the NIK locality. 2. Material and methods The fossil artiodactyl material from NIK is stored in the Laboratory of Geology and Paleontology of the Aristotle University of Thessaloniki (LGPUT). The present study focuses on dental and postcranial material of giraffids and cranial and dental material of bovids from NIK; bovid postcranials, though very abundant, are not included here and will be part of a separate forthcoming study. The basic morphology of most identified species has been already given in previous works (Kostopoulos et al., 1996; Kostopoulos and Koufos, 1999). Additional descriptions are included wherever necessary. The main goal of this paper is to update and revise previous determinations of the artiodactyl assemblage of the NIK locality, according to the additional evidences provided by the second fieldwork season.

http://dx.doi.org/10.1016/j.geobios.2016.01.011 0016-6995/ß 2016 Elsevier Masson SAS. All rights reserved.

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Lower teeth are in lower case, and upper teeth in upper case letters; the premolar to molar row length ratio is indicated as P/M for the upper and p/m for the lower tooth row. Measurements are given in millimeters; all row data are given as Supplementary Tables S1–S17. Dental terminology generally follows Heintz (1970) but lower premolars of giraffids are described according to Geraads et al. (2013). Statistical analyses were achieved using the PAST software (Hammer et al., 2001). Institutional abbreviations: AMNH, American Museum of Natural History, NY; AMPG, Athens Museum of Geology and Paleontology, National and Kapodistrian University of Athens; LGPUT, Laboratory of Geology and Paleontology, Artistotle University of Thessaloniki; MNHN.F, Museum National d’Histoire Naturelle, Paris; MMTT, Maragheh collection in the Laboratory of Evolutionary Biology, Howard University, Washington; MTA, Natural History Museum, General Directorate of Mineral Research and Exploration, Ankara; NHMA, Aegean Museum of National History, Samos Island, Greece; NHML, Natural History Museum, London; PIM, Palaeontological Institute Mu¨nster. Locality abbreviations: NKT, Nikiti 1, Greece; NIK, Nikiti 2, Greece; RZO, Ravin des Zouaves 5, Greece; PXM, Prochoma, Greece; VTK, VLO and VAT, Vathylakkos 1, 2 and 3, respectively, Greece; PER, Perivolaki, Greece; PIK, Pikermi, Greece; HD, Hadjidimovo, Bulgaria; MAR, Maragheh, Iran; AK, Akkas¸dag˘i, Turkey. Nomenclatural and measurement abbreviations: L: length; W: width, H: height; LPM/Lpm: length of the upper/lower premolar-molar series; MNI: minimum number of individuals; NISP: number of identified specimens; n: number of specimens. Additional abbreviations are given in Supplementary material. 3. Systematic paleontology Class Mammalia Linnaeus, 1758 Order Artiodactyla Owen, 1848 Suborder Ruminantia Scopoli, 1777 Family Giraffidae Gray, 1821 Genus Helladotherium Gaudry, 1860 Type-species: Helladotherium duvernoyi (Gaudry and Lartet, 1856); Pikermi, Greece; late Miocene. Helladotherium duvernoyi (Gaudry and Lartet, 1856) Figs. 1, 2 Studied material: Part of left mandibular ramus with p2-m3, NIK-1; part of right mandibular ramus with p2-m3, NIK-1804; left mandibular ramus, NIK-1057; scapula, NIK-766; radio-ulna, NIK1151; tibia, NIK-1805; talus, NIK-70, NIK-1017; posterior first phalanx, NIK-1096. Measurements: See Tables S1–S3. Description: LGPUT NIK-1804 seems to belong to the same individual as LGPUT NIK-1, although discovered ten years later. All studied tooth rows represent senile to very old individuals. The foramen mental of the mandible LGPUT NIK-1057 is large, ovalshaped and opens 100 mm in front of the p2 (Fig. 1). The caudal edge of the symphysis is placed at the level of the mental foramen. The ventral profile of the horizontal ramus between the mental foramen and the p2 is well concave (Fig. 1). The molar part of the horizontal ramus deepens significantly from p2 to m3. The ascending ramus of the mandible is wide ventrally and tapers gently to the top. Behind the m3, the anterior margin of the ascending ramus forms a 70o angle with the alveolar level, whereas the caudal margin is weakly concave in the upper part. The angle of the mandible is wide, and projects posteroventrally; it is marked by a thick, crest-like lateral lip. The vascular notch is weakly marked ventrally. The coronoid process is wide and slightly curves backwards with its tip remaining in front of the posterior level of

Fig. 1. Helladotherium duvernoyi from Nikiti 2, Greece. Mandibular ramus NIK-1057 in labial view and system of measurements (Table S1). Measures 9 and 10 represent ventral width of the mandibular corpus in front of p2 and behind m3, respectively; measure 15 corresponds to the transverse (mediolateral) diameter of the mandibular condyle. Scale bar: 10 cm.

the mandibular condyle. The condyle is subhorizontal and strongly projects lingually. The pterygoid fossa is wide and shallow, whereas the masseteric fossa is not marked. Dental features of NIK-1804 (Fig. 2(A)) are identical to those of LGPUT NIK-1 (Fig. 2(B)), which basic tooth morphology has been already given by Kostopoulos et al. (1996). The p2 is primitively simple and long (representing more than 80% of the p3 length), with a barely traced anterolingual stylid, which is, however, much more developed and distinct in LGPUT NIK-1057 (where the length of the p2 reaches 90% of the p3 length). The parasinusid of p3 closes early in wear, so the anteriorly directed paraconid fuses with the parastylid. The anterior valley (mesosinusid) of p3 should be originally open, but closes 10–13 mm above the base of the crown, not because of the junction of metaconid with the paraconid but due to thin cingular wings. At the same tooth, the postmetacristid is long in LGPUT NIK-1 and NIK-1804, reaching the lingual point of the hypoconulid. The entoconid of LGPUT NIK-1 and NIK-1804 is short, so the metasinusid and telosinusid communicate until an advanced stage of wear. In LGPUT NIK-1057 the metaconid seems

Fig. 2. Helladotherium duvernoyi from Nikiti 2, Greece. A. Right toothrow p2-m3, NIK-1804 in occlusal (A1) and labial (A2) views. B. Left toothrow p2-m3, NIK-1 in lingual view. Scale bar: 5 cm.

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to be more transversally placed and well distinguished from the entoconid that runs parallel to it. The p4 is fully molarized in all specimens but the contact between the postmetacristid and entoconid is still visible in advanced wear stage. The molars are wide, have a rather strong parastylid and their anterior and posterior lobes are tightly settled, separated by a deep narrow labial notch. The p4 and m3 of LGPUT NIK-1057 lack the large labial accessory pillars seen in NIK-1 and NIK-1804. The p/m ratio ranges from 61.7 to 67.2%. Remarks: In contrast to the NKT locality, giraffids are rather scarce in the NIK sample, both in number of specimens and species. Kostopoulos et al. (1996) already ascribed a worn p2-m3 series (LGPUT NIK-1) and an astragalus (LGPUT NIK-70) to H. duvernoyi, a species that is further documented in the new material by two additional tooth rows and several postcranials. Morphologically (Fig. 1), the mandible LGPUT NIK-1057 fits perfectly with the single known and sufficiently preserved mandibular ramus ascribed to Helladotherium from Maragheh (de Mecquenem, 1925: fig. 8), as well as with the complete sivatheriine left mandibular ramus AMNH 19684 from Dhok Pathan, Siwaliks, attributed to Bramatherium or Hydaspitherium (considered as synonyms by Geraads and Gu¨lec¸, 1999a). The mandible of the similarly large-sized Samotherium from China and the Aegean region (Table S1) differs from those specimens in the smaller mental foramen, the proportionally longer anterior (i.e., pre-p2) portion, the longer symphysis with its caudal edge well-behind the mental foramen, the much less concave or even flat ventral profile of the horizontal ramus in front of the p2, the caudally less deepened molar portion of the horizontal ramus, the less steeply raised (forming a 50o angle with the alveolar level) anterior margin of the proportionally wider ascending ramus, the almost straight caudal margin of the ascending ramus running subparallel to the anterior one, the more localized angle of the mandible, and the much more curved caudally coronoid process that reaches or marginally exceeds the caudal level of the condyle. Additionally the imprint of the masseteric muscle is clear on the left mandibular ramus NHMA MTLA311 of Samotherium major from Samos (Kostopoulos, 2009b), the pterygoid fossa is placed more ventrally, and the condyle is rather symmetrically placed when compared to the coronoid process in dorsal view. By its absolute length, long premolar row compared to the molars (Table S2), less advanced p3 morphology, wide p4 and molars, extremely large hypoconulid on m3, and strong lingual relief of the molars (Fig. 2), the dentition of the NIK species is easily distinguished from that of the similar-sized Samotherium ForsythMajor, 1888 from Samos (Kostopoulos, 2009b) and Vathylakkos, Axios Valley (Geraads, 1974, 1978), and resembles in all dental respects H. duvernoyi from Pikermi, Greece (Wagner, 1860: fig. 23), Maragheh, Iran (de Mecquenem, 1925: pl. II, fig. 1), and Hadjidimovo, Bulgaria (Geraads et al., 2005). Unfortunately, the NKT Helladotherium is known only by a few postcranials and thus there is no basis for a comparison with the NIK dentitions. The available radio-ulna (LGPUT NIK-1151), tibia (LGPUT NIK1805), tali (LGPUT NIK-70, NIK-1017) and proximal phalanx (LGPUT NIK-1096) (Table S3) are similar to those of H. duvernoyi from NKT, Perivolaki, Pikermi, Samos and Kerassia (Kostopoulos et al., 1996; Iliopoulos, 2003; Kostopoulos and Koufos, 2006; Kostopoulos, 2009b and pers. data), indicating a complete morphological and metric concurrence (Fig. 3). The scapula LGPUT NIK-766 belongs to an adult individual; hence, a full comparison with the rather young and partly destroyed scapula LGPUT PER295 of H. duvernoyi from Perivolaki (Kostopoulos and Koufos, 2006) is limited. Nevertheless, both specimens show a pentagonalshaped glenoid cavity, a straight spine, moderately projecting supraglenoid tuber with rounded lip, and subparallel lateral

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borders of the neck in lateral view. In contrast, Samotherium (Bohlin, 1926: pl. VIII, fig. 7) shows a sigmoidally developed spine, the tuber is more pronounced cranially, the acromion is more proximally placed compared to the glenoid cavity, and the lips of the neck are strongly concave. Several authors have made convincing important arguments for the possible synonymy of Helladotherium with the Asian late Miocene sivatheriines ascribed to the genera Bramatherium Falconer, 1845, Hydaspitherium Lydekker, 1876, and Vishnutherium Lydekker, 1876, which in turn may represent a single genus (e.g., Pilgrim, 1911; Matthew, 1929; Hamilton, 1978; Geraads and Gu¨lec¸, 1999a). Dental and mandibular features are also in favor of such a hypothesis, as evidenced by the remarkable similarity between LGPUT NIK-1057 and AMNH 19684 and by the small variations in dental features within the Helladotherium sample; hence, the p3 of LGPUT NIK-1, NIK-1804 and HD-5157 from Hadjidimovo (Geraads et al., 2005) are of ‘‘Vishnutherium-type’’ (Matthew, 1929: fig. 48), whereas the p3 of NIK-1057, MNHN.F MAR-882 from Maragheh (de Mecquenem, 1925) and the Pikermi p3 illustrated by Wagner (1860) are of ‘‘Bramatherium/Hydaspitherium-type’’(Colbert, 1935: fig. 181). However, pending a full taxonomic revision of the Siwaliks late Miocene sivatheriines, I shall continue to use Helladotherium as a valid designation for the European taxon. Genus Palaeotragus Gaudry, 1861 Type-species: Palaeotragus rouenii Gaudry, 1861; Pikermi, Greece; late Miocene. Palaeotragus rouenii Gaudry, 1861 Studied material: Left P2-(part)M1, NIK-1823; distal part of humerus, NIK-1810; radius of young individual, NIK-1809; metacarpal III + IV, NIK-1807; calcaneum, NIK-1912; part of cuboscaphoid, NIK-1914; proximal metatarsal III + IV, NIK-1808; first phalanx, NIK-1913; first, second and third phalanges, NIK1915. Measurements: See Table S4. Remarks: The presence in the NIK locality of a second, small giraffid with elongated and slender limb bones is documented only by a recently discovered P2-M1 tooth row and a few postcranials. P2–P4 length is about 50 mm. Both P2 and P3 have subcircular occlusal outlines and strong parastyle and paracone rib. P4 is more square-shaped in occlusal view, with somewhat flattened lingual wall, strong parastyle and metastyle, and slightly convex paracone rib. By their small size and morphology, the NIK upper premolars match closely those of Palaeotragus rouenii from Pikermi and Dytiko (Geraads, 1978; pers. data). The available postcranial elements also resemble those of Palaeotagus rouenii from SE Europe (e.g., Godina, 1979; Iliopoulos, 2003; Kostopoulos and Sarac¸, 2005; Kostopoulos and Koufos, 2006; Kostopoulos, 2009b). Kostopoulos et al. (1996) also refer to Palaeotragus cf. rouenii several postcranials from the NKT locality, but comparison between the two Nikiti samples indicates fundamental morphometric differences. As a whole, the NKT Palaeotragus has a long but stouter radius similar to P. rouenii, shorter but equally wide metacarpals, significantly shorter and more robust metatarsals, shorter and more robust tibia, and longer and wider tali. These differences clearly exceed those of intraspecific variability and refute the original species assignment for the NKT Palaeotragus – an issue that will be presented in of a forthcoming revision. Palaeotraginae indet. Studied material: Part of first phalanx lacking proximal epiphysis, NIK-211; proximal epiphysis of first phalanx, NIK-264 (young); second phalanges of young individuals, NIK-1916 (posterior?), NIK-1917 (anterior?). Measurements: See Table S4.

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Fig. 3. Metrical comparison of Helladotherium duvernoyi postcranials from Nikiti 2, Greece. Data from Kostopoulos et al. (1996), Iliopoulos (2003), Kostopoulos and Koufos (2006), and Kostopoulos (2009a).

Remarks: A few giraffid phalanges from the NIK locality are dimensionally in between those of Palaeotragus rouenii and Helladotherium duvernoyi, indicating the presence of a third giraffid taxon in the locality. These second phalanges are proportionally similar to those of Bohlinia attica from Dytiko (Axios Valley, Greece; Geraads, 1974, 1979) but differ in the absolutely and relatively deeper diaphysis, the more squareshaped proximal outline, and the lesser development of the lateral distal fossa. As these specimens are morphologically closer to phalanges of Palaeotragus and Samotherium they are referred to as Palaeotraginae indet., awaiting more material for a better identification. Family Bovidae Gray, 1821 Genus Tragoportax Pilgrim, 1937 Type-species: Tragoportax salmontanus Pilgrim, 1937; Dhok Pathan, Siwaliks, Pakistan; late Miocene. Tragoportax amalthea (Roth and Wagner, 1854) Figs. 4, 5(A)

1999. Tragoportax aff. rugosifrons (partim) - Kostopoulos and Koufos, p. 195. Studied material: Skull, NIK-1688; partial skull, NIK-1687; frontlets of young individuals, NIK-1135, 1689; partly preserved hornless skull, NIK-1684; palate, NIK-423, 1685; right P2-M3, NIK1141; mandible, NIK-1181; left part of mandible, NIK-1675; lower tooth rows, NIK-425, 433, 1676, 1691. Measurements: See Tables S5, S6. Description: The skulls LGPUT NIK-1688 (Fig. 4) and NIK-1687 show all main diagnostic features of Tragoportax Pilgrim, 1937: wide skull at the frontals and occipital region, rather low braincase, moderately bent down face compared to the basicranial axis, deep lachrymal fossae, grooved basioccipital, strongly developed and well-depressed dorsal rugose area marked by thick temporal ridges, raised frontals between the horn-cores forming a wide intercornual plateau, anticlockwise twisted horn-cores inserted moderately apart on the frontals and having a strong anterior keel, weak posterolateral keel and triangular basal cross-section. Both frontlets of young individuals (LGPUT NIK-1135 and NIK1689) are as wide at the orbits as LGPUT NIK-1688 and already

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Fig. 4. Tragoportax amalthea from Nikiti 2, Greece. Skull NIK-1688 in lateral (A), dorsal (B), and ventral (C) views. Scale bar: 5 cm.

show strong temporal ridges, but the rugose area and the intercornual plateau are not yet fully developed and the horncores are much more slender than those of adults and lack torsion. Nevertheless, the horn-cores of LGPUT NIK-1135 show an incipient anterior keel, and a flattening of the posterior surface, characters which confer a roughly symmetrically triangular cross-section to the horn-core base. A hornless skull (LGPUT NIK-1684) is provisionally ascribed to the same species because of its shallowness, the rather posterior position of the orbit compared to the toothrow, the development of the lachrymal fossa, and the ridged ventral border of the orbit, features that recall LGPUT NIK-1688.

Remarks: Kostopoulos and Koufos (1999) ascribed all boselaphine-like material discovered during the first fieldwork season in NIK to Tragoportax aff. rugosifrons, but Spassov and Geraads (2004) mentioned that some of the described postcranials should, in fact, belong to Miotragocerus Stromer, 1928. The new material allows confirming the presence of two boselaphine taxa in this locality. The skulls LGPUT NIK-1688 and NIK-1687 show all main diagnostic features of Tragoportax Pilgrim, 1937 (Pilgrim, 1937; Spassov and Geraads, 2004; Fig. 4; Table S5). Compared to T. rugosifrons (Schlosser, 1904) from Samos, the NIK species differs in the relatively longer braincase compared to the face, the weakly converging caudally (instead of being parallel) lateral braincase

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Fig. 5. Comparison of the mandibles of Tragoportax amalthea (NIK-1675; A) and ?Miotragocerus sp. (NIK-764; B) from Nikiti 2, Greece. Scale bar: 5 cm.

sides, the weaker bent basicranial axis with respect to the palate (30o instead of 45o; both specimens not deformed), the lower face with narrower lachrymal fossa and regularly sloping down nasals, the caudally prominent nuchal crest, the roughly semicircular (instead of trapezoidal) occipital outline with well-convex dorsal edge, and the more backward-inclined and twisted horn-cores, with sharper anterior keel. This set of morphological features closely approaches the NIK sample to T. amalthea crania from Pikermi and Halmyropotamos (NHML M10836, M11420; MNHN.F PIK2287; AMPG PA2084/91, AMPG 1967/38; Melentis, 1968, and pers. obs.) as well as from Maragheh, Iran (Kostopoulos and Bernor, 2011), where I prefer now to include it. On the other hand, supposedly differentiating features between T. amalthea and T. rugosifrons horn-cores such as the stronger keel demarcations and exostoses (rugosities) seen in several specimens of the former species may depend on the much better sample reflecting individual and ontogenetic variability. Upper and lower dentitions of the two NIK boselaphine-like species are extremely variable and greatly overlapping in both morphology and size (Table S6). Therefore affiliation of individual specimens is rather provisional, based on the dental features of the preserved crania and partly on other characters such as the position of the infraorbital foramen compared to the alveolar level, the deepness of the mandibular corpus, the height of the ascending ramus, etc. As a whole, the present taxon is characterized by a relatively small P2 with weakly developed anterior complex, a long p2 when compared to the p4, and a most frequently open anterior valley on p4. The P/M and p/m ratio are, on average, 77.2% (n = 8, standard deviation = 0.94;) and 72.8% (n = 6, standard deviation = 2.12;), respectively (Table S6); both values fall within the ranges of T. amalthea from Pikermi and Maragheh and T. rugosifrons from Samos (Roussiakis, 1996; Kostopoulos, 2006, 2009c; Kostopoulos and Bernor, 2011). Genus Miotragocerus Stromer, 1928 Type-species: Miotragocerus monacensis Stromer, 1928; Oberfo¨hring, Austria; late Miocene. ?Miotragocerus sp. Figs. 5(B), 6 1999. Tragoportax aff. rugosifrons (partim), Kostopoulos and Koufos, p. 195 Studied material: Partial skull with associate mandible, NIK764; anterior part of skull with palate, NIK-422, 1690; lower dentitions, NIK-1103, 1140, 1680, 1681. Measurements: See Tables S5, S6.

Remarks: The second boselaphine species from NIK is documented by a fairly complete skull (LGPUT NIK-764; Fig. 6) and several upper and lower tooth rows unearthed during the second round of field work, as well as the palate LGPUT NIK-422 previously ascribed to Tragoportax aff. rugosifrons (Kostopoulos and Koufos, 1999: 195). The long, high, narrow and posteriorly curved down braincase, the subtriangular occipital face, the large round orbits placed rather rostrally (anterior margin of orbits above M2/ M3 level), the high and weakly sloping down face, the narrow intercornual plateau, the moderately developed dorsal rugose area, the long basioccipital marked rostrally by a longitudinal crest, the large upper premolars, and the tear-shaped cross-section of the horn-cores (Fig. 6) exclude this taxon from Tragoportax and bring it closer to the Turolian boselaphine species currently grouped under Miotragocerus Stromer, 1928 (sensu Spassov and Geraads, 2004). Nevertheless, the saber-like shape of the horn-cores and the rather advanced dentition with small p2 and frequently molarized p4 are important differences from known species of the latter genus, making inconclusive a definite generic affiliation. In most of its features the studied NIK sample resembles T. curvicornis Andre´e, 1928 from Samos (Kostopoulos, 2009c) and some of the Axios Valley skulls referred to as T. rugosifrons (Bouvrain, 1994), especially LGPUT PXM 91 and Vathylakkos Ravin C-1a. The mandible LGPUT NIK-764 (Fig. 5(B)) differs from that of the previously described T. amalthae (LGPUT NIK-1675; Fig. 5(A)) in the higher ascending ramus, the wider and longer coronoid process, the narrower mandibular incisure, the longer and deeper pterygoid fossae, the more convex ventral edge of the horizontal ramus between m1 and m3, and the smaller mandibular foramen. As mentioned above, dental morphometric features of this taxon greatly overlap with those of T. amalthea from the same site but the P2 is proportionally larger, with a better developed anterior complex and an usually marked parastyle; the p2 is smaller compared to the p4, and the p4 frequently exhibits a continuous lingual wall. The upper premolar row represents 75.6% of the molars (n = 2) and the lower premolar row 69.7% of the molars (n = 5; standard deviation = 4.8). Genus Gazella Blainville, 1816 Type-species: Capra dorcas Linnaeus, 1758; lower Egypt; extant. Gazella pilgrimi Bohlin, 1935 Fig. 7(B) Studied material: Partially preserved skull, NIK-1811; frontlets, NIK-813, 1813, 1814, 1817; left horn-core, NIK-700, 1815; right horn-core, NIK-1816, 1818; palate, NIK-1899; right P2-M3, NIK-607; part of mandible with i1-m3 sin. and i1, p2-m3 dex., NIK778; right mandibular ramii, NIK-589, 1888, 1890, 1898, 1902; left mandibular ramii, NIK-13’, 590, 1891, 1893-1895, 1900, 1901. Measurements: See Tables S7, S9, S10. Description: The horn-cores are moderately long (L: 140– 150 mm) with flattened lateral surface, oval cross-section throughout most of their length except in the apical part, and distinctly grooved surface, especially medially and anterolaterally (Fig. 7(B)). The horn-cores are slightly curved and moderately inclined backwards (about 50o in lateral view), and they either diverge each other moderately from the base (258) or run subparallel at the base and more strongly diverge in the distal 2/3. The pedicle is three times longer anteriorly than posteriorly. The supraorbital foramina are small, placed in narrow, subtriangular to elliptical pits, settled widely apart on the frontals. The postcornual fossa is small and deep. The interfrontal suture is open and pinched along the entire frontal length. The skull is wide at the orbits and has a globular braincase slightly inclined when compared to the face (Fig. 7(B)). The endoparietal is narrow, trapezoidal shaped, and represents 1/4 of the parietal dorsal length. The face is short with

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Fig. 6. ?Miotragocerus sp. from Nikiti 2, Greece. Skull NIK-764 in lateral (A), dorsal (B), ventral (C), and caudal (D) views. Scale bar: 5 cm.

the anterior margin of the orbit placed above the middle of M2. The ventral border of the orbit is marked by a sharp jugal crest that prolongs on the face until above the front of the M2. The infraorbital foramen is large and opens above the front of P3. The lachrymal depressions are partly deformed but they seem to be small and moderately deep. The occipital is rather shallow and wide, subtriangularly outlined, with a prominent external occipital crest that makes occiput facing bilaterally. The foramen magnum and the occipital condyles are relatively small; the bicondylar width represents 60% of the bimastoid width. The mastoid is relatively restricted on the occipital plane and faces posterolaterally. The basioccipital slightly narrows to the front, and bears crest

like posterior tuberosities weakly extended laterally, and small, closely settled, and ventrally upraised anterior tuberosities separated by a median groove; the lateral borders of the basioccipital between the anterior and posterior tuberosities are slightly convex. The U-shaped choane and the pterygoid fossae open at the level of the posterior lobe of M3. The premolars are relatively short compared to the molars (P/M ratio: 66–69%, n = 4; p/m ratio: 53–64%, n = 6); the P2 is wide (WP2/LP2 > 80%); the P2 and P3 have a trapezoidal occlusal outline and a thin parastyle that marginally reach the labial wall; the p3 and p4 usually have a pinched hypoconid, a weak paraconid, an open anterior valley and posteriorly directed metaconid that closes the posterior valley; the

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Fig. 7. A. Skull of Gazella cf. capricornis, NIK-1812 (Nikiti 2, Greece), in lateral (A1) and dorsal (A2) views. B. Skull of Gazella pilgrimi, NIK-1811 (Nikiti 2, Greece), in lateral (B1) and dorsal (B2) views. Scale bar: 5 cm.

m1 and less often the m2 bear a basal pillar. A couple of specimens (LGPUT NIK-1817, 700) show much shorter (L: 95–100 mm), almost straight and less mediolaterally compressed horn-cores that, however, have similar posterior inclination, comparably long anteriorly pedicles, and equally deep longitudinal grooving as the rest of sample; they may represent young individuals of the same species. Remarks: The first fieldwork session allowed the identification of a single gazelle species at NIK (Kostopoulos and Koufos, 1999). The presence of a second, actually more abundant gazelle was revealed during the second round of field work by a skull, several frontlets and mandibles, altogether representing a minimum of eight individuals (MNI based on either horn-core or mandibular specimens; Tables S7, S9, S10). Horn-core size and morphology (Figs. 7(B), 8; Table S9) match perfectly with G. pilgrimi from the Axios Valley and Samos (Bouvrain, 1996; Kostopoulos, 2009c). Bouvrain (1996: 116) suggested that the horn-cores of G. pilgrimi may exhibit a shortening and an increase of posterior curvature from early to middle Turolian, but the NIK data fail to confirm this. In all studied samples, the anterior length of the horn-core falls usually between 130 and 160 mm (n = 18) with a couple of exceptions from the Ravin de Zouaves 5 locality (Axios Valley, Greece), where it may reach 200 mm. At the same time, backward curvature seems to be variable in each local sample but with a curvature index (Kostopoulos, 2005: 750) always less than 120 (i.e., 103–115 in NIK (n = 7); 103–109 in RZO (n = 4), 106–118 in VAT (n = 4), and 105–112 in PER (n = 5)). However, the compression index, both at the base of the horn-cores and at seven centimeters above it, does show a weak reduction from NIK to Perivolaki and between the

two samples of Samos (Fig. 8); it may indicate a general trend through time. The skull morphology, absolute dimensions and proportions of the NIK gazelle are fully compatible with the few available and partially preserved skulls of this species from the Axios Valley (namely LGPUT VTK-17, 114; LGPUT PXM-49 and MNHN.F Slq809) discussed by Arambourg and Piveteau (1929), and Bouvrain (1996, 2001), especially regarding the occipital and basioccipital features, as well as the position and shortening of the face compared to the braincase (Table S7). The AMNH 20580 skull from the Q5 site of Samos, tentatively ascribed to the same species (Kostopoulos, 2009c: 355), differs from the continental Greek sample by its stronger temporal lines, wider choanae that open slightly in front of the pterygoid fossae, more retired occipital condyles, less narrowing basisphenoid and longer tooth row. In all available NIK and Vathylakkos specimens, as well as in most Prochoma ones, the postcornual fossae are rather narrow, small and moderately deep, whereas in one specimen from Prochoma (LGPUT PXM-62), the two frontlets from Perivolaki and in most Samos specimens the postcornual fossae are significantly wider and usually shallower. Tooth row morphology and size, and premolar shortening compared to the molars of the NIK sample closely match those from the Axios Valley (Bouvrain, 1996, 2001) in every respect (Table S10). Gazella cf. capricornis (Wagner, 1848) Fig. 7(A) Studied material: Partially preserved skull, NIK-1812; partially preserved skull with mandible, NIK-1142; frontlet, NIK-699; right horn-core, NIK-1819; basal part of left horn-core, NIK-1820; right

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Fig. 8. Boxplots of horn-core mediolateral compression at the base (left) and at 7 cm above the base (right) in Gazella pilgrimi samples from different Greek sites. NIK: Nikiti 2; RZO: Ravin des Zouaves-5; VAT: Vathylakkos-3; PXM: Prochoma; PER: Perivolaki; PIMSamos: Samos-PIM collection; MTLAB: Samos-MTLA/B sites. Data from Kostopoulos (2006, 2009b) and pers. obs.

P2-M3, NIK-607, 1845. Possibly ascribed: right mandibular ramus with p4-m3, NIK-1892; left mandibular ramus with p3-m3, NIK1889, 1896; right mandibular ramus with p4-m3, NIK-1897. Measurements: See Tables S8, S9, S11. Remarks: The presence in the NIK locality of a gazelle similar to the typical Pikermi taxon, Gazella capricornis, was already known (Kostopoulos and Koufos, 1999). The new material from NIK barely enriched the sample of this species, from which the horn-core specimen LGPUT NIK-700 should be excluded, as it may belong to a young individual of G. pilgrimi. Of similar size and unfortunately known mostly by their horn-cores and dentitions, G. capricornis and G. pilgrimi stand as the two dominant gazelles of the earlymiddle Turolian of Southern Balkans and Turkey. More complete skull remains of these two common species are scarce, limiting diagnostic features (e.g., Bouvrain, 1996; Kostopoulos, 2009c; Kostopoulos and Bernor, 2011). This fact, along with the apparently great dental and horn variability, leaves room for reasonable doubts and taxonomic confusion (e.g., Solounias, 1981; Bibi and Gu¨lec¸, 2008). The fairly complete, though moderately deformed skulls LGPUT NIK-1142 and NIK-1812 (Fig. 7(A); Tables S8, S9) match quite well the general morphological and metric features of G. capricornis skull as summarized by Kostopoulos (2009c: 351) and Kostopoulos and Bernor (2011: 655); they allow extracting some important differential characters compared to the G. pilgrimi skull from the same site (LGPUT NIK-1811; Fig. 7(B)) as well as those from the Axios Valley (Arambourg and Piveteau, 1929; Bouvrain, 1996, 2001). Hence, LGPUT NIK-1142, 1812 and G. capricornis differ from G. pilgrimi in (Fig. 7):  the slightly larger absolute size;  the shorter (L < 120 mm), more uprightly inserted (> 608), more posteriorly curved (curvature index usually > 120), and thinner grooved horn-cores that may run parallel to each other or uniformly converge from the base;  the anteriorly much shorter pedicles (anterior length less than half of the posterior one;  the larger, triangular to tear-shaped supraorbital pits;  the larger postcornual fossae;  the smaller infraorbital foramen;  the thinner ventral border of the orbit with only anterior and posterior remains of a jugal crest;

 the longitudinally domed nasals;  the larger endoparietal representing about half of the parietal dorsal length;  the slightly longer and less globular braincase with relatively larger occipital condyles (bicondylar width against width at the anterior tuberosities of the basioccipital < 2 in G. pilgrimi, and > 2 in G. capricornis);  the hexagonal outlined occipital plane with occiput facing mostly posteriorly;  the longer exposed mastoid on the occipital plane, facing mainly caudally;  the stronger and much more laterally expanded posterior tuberosities and the larger and bulbous anterior tuberosities of the basioccipital, which lateral borders are variably concave;  the slightly deeper mandible with more vertical ascending ramus;  the longer P2 (L/W for P2 < 80 (n = 8) of G. capricornis from Pikermi, Axios Valley and NIK, vs. > 80 (n = 8) for G. pilgrimi from NIK and the Axios Valley) with a stronger parastyle;  the longer upper premolars compared to the molars (P/M > 70 (n = 10) for G. capricornis from Pikermi, Axios Valley and NIK, vs. < 72 (n = 8) for G. pilgrimi from NIK and the Axios Valley);  the well-developed paraconid on p3 and p4 (always weak in G. pilgrimi).

Distinction of the lower dentition of these two species is less obvious than previously thought (e.g., Kostopoulos, 2009c: 351– 352; Tables S10, S11): the lower premolar to molar ratio shows a greater overlap than for the upper dentition (p/m > 60 (n = 8) for G. capricornis from Pikermi vs. < 64 (n = 8) for G. pilgrimi from NIK and the Axios Valley), and the posterior valley of the p4 appears closed in all four specimens of G. capricornis from NIK and in 30% of those of Pikermi (n = 20; Roussiakis, 1996; pers. obs.), whereas it is always closed in G. pilgrimi. Genus Nisidorcas Bouvrain, 1979 Type-species: Nisidorcas planicornis (Pilgrim, 1939); Perim Island, India; late Miocene. Nisidorcas planicornis (Pilgrim, 1939) Fig. 9

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Fig. 9. Nisidorcas planicornis from Nikiti 2, Greece. A. Skull NIK1848 in lateral (A1), ventral (A2), dorsal (A3), and caudal (A4) views. B. Skull NIK-1847 in lateral (B1) and dorsal (B2) views. Scale bar: 5 cm.

1999. cf. Ouzocerus sp. (partim) - Kostopoulos and Koufos, p. 208. Studied material: Skull, NIK-11, 101,577, 1846, 1847, 1848, 1859, 1860, 1862, 1866; frontlet, NIK68, 69, 571, 572, 574, 576, 657, 782, NIK-1849–1858, 1861, 1863–1865; left horn-core, NIK575’, 1867–1878; right horn-core, NIK-12, 257, 1879–1887; upper tooth rows, NIK-597, 598, 600–603, 605, 606, 608, 683–685, 777, 821, 1078, 1136, 1903, 1904; lower tooth rows, NIK6, 583, 585– 588, 593, 595–597, 599, 600, 682, 1019, 1101, 1859, 1904–1911. Measurements: See Tables S12–S14. Description: Craniodental and horn-core morphological features agree with those reported by Bouvrain (1979; 1992) and Kostopoulos (2006) with the following additions: the craniofacial angle (measured along frontals) ranges from 1308 to 1558; the basicranial angle (basioccipital-palate) is about 1508; the lachrymal fossa is rather small, not well defined and shallow; the jugolachrymal suture passes from the middle level of the anterior orbital rim; the infraorbital foramina are variable in size, and pocketing; the nasals are 15–25% longer than the frontals (along midsagittal plane); the ethmoidal fissure is closed; the orbital borders are quite prominent (width of the skull at the orbits appears 40% larger than at the braincase); the foramen oval is moderate to large, facing laterally; the choanae open at the same level with the pterygoid fossae, at the back end of M3; the upper premolar row represents 62–77% of the upper molar row length and the lower premolars represent 51–61% of the molars; the horncore posterior inclination and divergence angles are equal (458). Remarks: N. planicornis is by far the most abundant bovid in the NIK assemblage, known by dozens of cranial and postcranial specimens. Based on the material unearthed from the first fieldwork period (Kostopoulos and Koufos, 1999), Kostopoulos (2000) estimated the minimum number of the individuals (MNI) of this taxon at 17. The new material includes, however, 7 additional

fairly complete skulls (Fig. 9), 15 frontlets and 26 isolated horncores (13 of them from the left side), suggesting a much higher number of individuals (i.e., MNI = 45 based only on skull and horncore specimens). The frontlet LGPUT NIK-68, previously referred to as cf. Ouzoceros sp. (Kostopoulos and Koufos, 1999: 208) is now believed to be a large Nisidorcas individual. The morphological and metric features, as well as the population and time variability of the species have been thoroughly discussed by Bouvrain (1979, 2001), Ko¨hler (1987), Kostopoulos and Koufos (1999), and Kostopoulos (2000, 2006). The NIK material of N. planicornis permits, however, a better definition of some diagnostic features mentioned by Bouvrain (1992: 55–56) and to complete missing ones (Tables S12–S14). Bouvrain (2001) referred to as ?Nisidorcas sp. two horned cranial specimens from Vathylakkos (LGPUT VTK-12 and VTK115). Morphological comparison of these specimens with Nisidorcas crania from NIK fully confirms their attribution to N. planicornis. The skull of N. planicornis from Vathylakkos (as extracted by combination of LGPUT VTK-66, 85, 95, 115) is slightly longer (5% in most linear length measurements) and significantly wider (> 10% in most linear width measurements) than that of NIK, especially in the orbital region (Table S12). Additionally, the infraorbital foramina of the Vathylakkos sample are larger, the auditory bullae are more elongated, the foramina oval are larger and well in front of the anterior tuberosities of the basioccipital, and the posterior tuberosities are much more developed in ventral sense. The horn-cores from NIK differ from those of all other Greek samples (RZO, PXM, VTK, PER; Table 1) in the tighter twisting (> 1), the absence of spiraling, the weaker compression especially distally, and the thick-swollen posterior keel in its basal part. As, however, the basic skull, horn-core and dental morphology remains remarkably stable through time, these differences do not seem sufficient for a distinction at the

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Table 1 Comparison among Nisidorcas planicornis populations from several sites.

Skull width at the horn-cores (lateral) W supraorbitals Supraorbitals Postcornual groove Horn-core length Horn-core torsion Horn-core spiralling APD base trending Posterior keel (basal) Anterior Keel Horn-core divergence Horn-core MLC at base Horn-core MLC at 7 cm above base (mean) LPM Lpm

NIK

RZO

PXM

VTK

PER

Kayadibi

Maragheh

Type (BMNH M37264)

67.0–79.0 Mean 73 27–36 Mean 32.5 Small Deep 130–190 Mean 160 Tight, > 1 No a/p Thick-swollen No

77–80 Mean 79 ?

[73.5]

84.0

[60–65]

84

?

[30]

?

77–86 Mean 80 27

30–34

38.2

?

? Deep 160

Small Shallow 160

Small Mod. deep  180

Medium (n = 1) Deep 150–170

?medium Shallow

? Shallow (> 110 + )

Loose, < 1 Weak a/p Thin-blunt No

Loose, < 1 Weak a/p Thin-blunt No

Loose, < 1 Weak a/p Thin-blunt Rather no

Loose, < 1 Weak ?m/l ? No

Loose, < 1 Weak m/l Thin-blunt Blunt constriction

Loose, < 1 Weak ? Thin-blunt No

40–55o 74–92 82.5 (n = 63) 115 (n = 9)

40–58o 75–87 81 (n = 8) 120 (n = 4)

45o

[50–55o] 75.5–87.0 80.5 (n = 7) –

50o 78–81 79.5 (n = 2) –

?

83.5 123.5

45o 68.2–82.4 76 (n = 11) 131 (n = 3)

Small Shallow 130–200 Mean 160 Loose,  1 Weak m/l Thin-blunt Shallow wide furrow 48–55o 76.6–88.8 81.5 (n = 16) 133 (n = 4)

50.0–55.5 54.6–60.0

– –

– –

53.0–56.6 58.2

57.5–60.1 57.2–62.1

– –

– –

– –

70.2 –

NIK: Nikiti 2; RZO: Ravin des Zouaves 5; PXM: Prochoma; VTK: Vathylakkos 1; PER: Perivolaki. Data for Kayadibi, Turkey from Ko¨hler (1987).

species level. Updating previous data (Table 1) it may be assumed that the species shows through time a widening of the skull at the orbit level, elimination of the postcornual fossae, loosening of horn-core torsion, a slight increase of spiraling, an increase of the lateral compression, especially distally (Fig. 10), a clockwise shift of the great basal axis on left, a thinning of the posterior keel in its proximal part at least, and finally initiation of an anterior ‘‘keellike’’ structure. Apart from Greece, N. planicornis is recorded in Turkey (localities Kayadibi, C¸oban Pinar; Ko¨hler, 1987; Geraads and Gu¨lec¸, 1999b), Iran (locality complex of Maragheh; Kostopoulos and Bernor, 2011) and India (type locality of Perim Island; Pilgrim, 1939; Bouvrain, 1979), suggesting a wide late Miocene geographic distribution, comparable to that of gazelles and boselphine-like bovids. By their loose torsion and basally thin posterior keel, the Kayadibi, C¸oban Pinar, Iranian

Fig. 10. Boxplot of horn-core mediolateral compression at the base in different samples of Nisidorcas planicornis. Site abbreviations as in Fig. 8, and VTK: Vathylakkos-1; KDB: Kayadibi, Turkey; MAR: Maragheh, Iran. Data from Ko¨hler (1987), Kostopoulos (2006), and Kostopoulos and Bernor (2011).

and Indian samples of Nisidorcas stand closer to the Vathylakkos and Perivolaki morphotype. Genus Palaeoreas Gaudry, 1861 Type-species: Antilope lindermayeri Wagner, 1848; Pikermi, Greece; late Miocene. Palaeoreas lindermayeri (Wagner, 1848) Fig. 11 1999. cf. Ouzocerus sp. (partim) - Kostopoulos and Koufos, p. 208. Studied material: part of skull with horn-cores, NIK-749; part of skull, NIK1824; frontlets, NIK-1825, 1826, 1828; distal part of left horn-core, NIK-1828; parts of left and right horn-cores, NIK1829; palate, NIK-1844; upper tooth row, NIK1842, 1843; mandibular ramii, NIK-52, 214, 565, 568, NIK-806, 1024, 1082, 1083, 1102,1160, NIK-1830-1841. Measurements: See Tables S15–S17. Description: The skull LGPUT NIK-1824 (Fig. 11(A)) shows a moderately long face; wide and shallow ante-orbital depression; infraorbital foramen above P2; narrow, V-shaped choanae with anterior end at the back lobe of M3; frontals significantly raised between the horn-cores; small supraorbital foramina within large depressions, separated by a wide interfrontal ridge; moderately short braincase; low and broad occipital; relatively long basioccipital; long, straight moderately divergent (508), weakly posteriorly inclined horn-cores inserted behind the orbits, heteronymously twisted on their axis with a strong posterior and a weak ridge-like anterior keel. The P2 is as long as or slightly longer than P3 and its width represents 2/3 of its length; in half of the specimens it appears weakly bilobed lingually (Fig. 11(E)). The paracone rib of the P2 and P3 is strong and placed anteriorly, whereas the parastyle is stronger in P3 than in P2. The central fossete of P3 is usually unequally divided. The P4 is more symmetrical with strong parastyle and paracone rib. All molars have a strong parastyle, mesostyle, and paracone rib. M1 and M2 usually bear a small basal pillar, whereas M2 and M3 show a central islet as well. A weak hypoconal spur is present on M2, vanished with wear. The upper premolars represent 61–69% of the molars. The paraconid is well developed and distinct from the parastylid in 78% of the available p3 and in 87% of the p4 (n = 23). 31% of the p3 (n = 19) show a

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Fig. 11. Palaeoreas lindermayeri from Nikiti 2, Greece. A. Skull NIK-1824 in lateral (A1) and dorsal (A2) views. B. Frontlet NIK-1826 in frontal view. C. Frontlet of a young individual, NIK-1828, in frontal view. D. Right mandibular ramus NIK-1083 in occlusal view. E. Left P2-M3 NIK-1849 in occlusal view. Scale bars: 5 cm (A-C), 2 cm (D, E).

transversally settled front edge. All p3 have a thin and posteriorly directed metaconid that rests independent from the entoconid until advanced wear stage (Fig. 11(D)). The metaconid of the p4 is more transversally settled and larger (Fig. 11(D)). In 17% of the p3 and 8% of the p4 the metaconid fuses early in wear with the entoconid closing the posterior valley of the tooth. Additionally in a single specimen (LGPUT NIK-52) the metaconid turns anteriorly and contacts the paraconid, closing the anterior valley of the tooth. In 50% of the p4 (n = 21), a small tubercle emerges from the base of the anterior valley. All p3 and p4 show a moderate to strong labial groove separating the protoconid from the hypoconid. The lower molars show a weak to moderate goat fold and a thin basal pillar that may, however, be absent in the m3. The third lobe of the m3 is formed by a single tubercle. The lower premolars represent 55.6– 62.7% of the molars. Remarks: This spiral-horned antelope is represented in NIK by seven individuals at least. Kostopoulos and Koufos (1999) mistakenly referred the lower tooth rows LGPUT NIK-52, 214, 565, and 568 as cf. Ouzocerus sp. The new material from NIK clearly shows that this first sample belongs to Palaeoreas, a taxon further documented in the new collection by a partly complete skull (LGPUT NIK-1824; Fig. 11(A)), several frontlets (Fig. 11(B, C)), and

numerous tooth rows. The last forty years, Palaeoreas underwent several important revisions (e.g., Gentry, 1971; Bouvrain, 1980, 1992; Geraads et al., 2003) that make it one of the best-defined late Miocene bovid taxa. Palaeoreas is known by two species: the moderately large P. zouavei Bouvrain, 1980, known exclusively from the Early Turolian faunas of the Axios Valley, Greece (Bouvrain, 1980, 1992), and the smaller P. lindermayeri, known originally from Pikermi (Gaudry, 1861, 1865; Roussiakis, 1996) but also recorded in the Axios Valley, Halmyropotamos, Greece (Melentis, 1968; Bouvrain, 1980), several Bulgarian localities (Geraads et al., 2003, 2011; Hristova et al., 2013), and KemiklitepeD, Turkey (Geraads and Gu¨lec¸, 1999b). The occurrence of P. lindermayeri in Samos and other Balkan and Turkish sites is, however, disputable (e.g., Geraads et al., 2003; Kostopoulos, 2004). The general skull and horn-core features and the overall size of the NIK sample clearly match those of P. lindermayeri as defined by Bouvrain (1980, 1992) and Geraads et al. (2003) (Fig. 12; Tables S15–S17). The single available skull LGPUT NIK-1824 (Fig. 11(A)) is clearly smaller than Palaeoreas lindermayeri from Hadjidimovo, Bulgaria (Fig. 12A), and absolutely and proportionally closer to the Pikermi sample of the species (Figs. 12(A), 13). However, when eliminating

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Fig. 12. Principal Component Analysis of Palaeoreas lindermayeri skulls from several fossil localities. A. PCA based on original raw data. B. PCA based on the same data matrix as previously but after eliminating the size effect using Mosimann’s Log-Shape ratio (i.e., by dividing each value with the specimen’s geometric mean and logarithmizing the results). Bi-co: Bi-condylar width; Bi-ped: Bi-pedicle width; Wbc: Braincase width; Woc: Occipital width; PM: length P2-M3; TDh-c: Horn-core transverse diameter at the base; APDh-c: Horn-core anteroposterior diameter at the base. Data from Geraads et al. (2003).

the effect of size the skull and horn-core shape of all studied samples appears basically the same (Fig. 12(B)). The NIK skull differs from Palaeoreas lindermayeri from Pikermi, Vathylakkos (Greece), Hadjidimovo, and Kalimantsi (Bulgaria) in the less concave frontonasal profile, the less steep craniofacial angle (the frontoparietal angle is usually less than 1008 in Pikermi, and Hadjidimovo, instead of 1208 in LGPUT NIK-1824; the angle between the anterior and posterior parts of the frontal is 90–1008 in Hadjidimovo, 78–878 in Vathylakkos, and 105–1188 in Nikiti 2), the slightly shorter face (the anterior border of the orbit is above the M2/M3 level), the anteriorly narrowing nasals (instead of having parallel sides in Pikermi and Hadjidimovo), the less transversally projected posterior and the much weaker anterior tuberosities of the basioccipital without central groove between them, and the less massive though equally long horn-cores (Fig. 13). A large and deep postcornual fossa is present in LGPUT NIK-1828 (possibly representing a young individual) but this feature is absent in the remaining specimens.

Though morphologically very close to the Pikermian sample (e.g., Gentry, 1971; Roussiakis, 1996), the NIK upper dentition (Table S17) appears slightly longer on average, with longer molar row and shorter premolars compared to the molars (Fig. 14). Dental characters of the lower tooth row from NIK also agree with those given by Gentry (1971), Roussiakis (1996), and Geraads et al. (2003, 2011) for P. lindermayeri (Table S17). The NIK lower dentition shows p/m ratio similar to that from Pikermi and Hadjidimovo, though it is dimensionally intermediate between these two populations and closer to the Strumiani sample (Geraads et al., 2011; Fig. 14). Genus Palaeoryx Gaudry, 1861 Type-species: Antilope pallasi Wagner, 1857; Pikermi, Greece; late Miocene. Palaeoryx cf. pallasi (Wagner, 1857) Fig. 15

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Fig. 13. Comparison of horn-core basal diameters in Palaeoreas lindermayeri from different fossil localities. Data from Geraads et al. (2003, 2011) and Geraads and Gu¨lec¸ (1999b).

Studied material: left maxilla with P2-M3, NIK-1063. Measurements (in mm): LP2-M3 = 115.45; LP2-P4 = 51.7; LM1M3 = 66.3; LP2 = 16.3, WP2 = 13.6; LP3 = 18.2, WP3 = 16.8; LP4 = 15.3, WP4 = 19.2; LM1 = > 18, WM1 = 22.8; LM2 = > 21, WM2 = 26.0; LM3 = 23.9, WM3 = 24.5. Description: The P2 is slightly shorter than the P3 and wider posteriorly than anteriorly (Fig. 15(B)); the parastyle is weak and marginally reaches the labial wall; the paracone rib is moderately developed and shifted anteriorly; the lingual crescent is bilobed. The P3 has a semi-circular occlusal outline, wider posterioly than anteriorly, though the posterolingual third of the tooth is broken; the parastyle is strongly developed and the paracone rib is strong and anteriorly placed and directed. The P4 is wider than long, with equally strong developed parastyle and metastyle, and moderately developed paracone rib placed centrally (Fig. 15(B)). All molars bear strong ribs and styles (with the mesostyle being more pinched

Fig. 15. Palaeoryx cf. pallasi from Nikiti 2, Greece. Left P2-M3 NIK-1063 in labial (A) and occlusal (B) views. Scale bar: 5 cm.

than the other two) and a strong flattened basal pillar attached to the anterior flange of the hypocone (Fig. 15(B)). The fossetes are simple, narrow and deep; the protocone is narrower and more protruding lingually than the hypocone, whereas both of these crescents are slightly constricted lingually. The lobes of the molars fuse slowly together with wear giving rise to a central islet. The occlusal surface of the molars is steep. Remarks: A third large bovid is documented in the NIK locality by a single upper tooth row that appears 15–20% longer than those of the large-sized boselaphines from the same site. Although in advanced stage of wear, the teeth indicate a rather brachyodont bovid with long premolar row (78%) when compared to the molars. The absolute dimensions of the tooth row, the long premolars compared to the molars, the long, lingually bilobed P2, and the complicated labial and lingual relief of the molars are fully compatible with those of Palaeoryx, a large-sized antilopine well known from the Aegean region during the late Miocene (Roussiakis, 1996; Kostopoulos, 2005, 2009c). LGPUT NIK-1063 shows a relatively (compared to the premolars) and absolutely longer molar row than Palaeoryx majori from Samos (Greece), and Akkas¸dag˘i (Turkey) (Kostopoulos, 2005, 2009c), from which it also differs by the absence of a central fold on P3 and P4 and the presence of strong basal pillars on the upper molars. Both features, along with the strong parastyle on P3 and the strong mesostyle on the molars, are similar to those of P. pallasi from Pikermi (Gentry, 1971; Roussiakis, 1996; Kostopoulos, 2009c).

4. Conclusions

Fig. 14. Distribution of lower premolar per molar row length in Palaeoreas lindermayeri from different fossil localities. Data from Geraads et al. (2003, 2011).

The updated and revised artiodactyl list of the Nikiti 2 locality includes three giraffid and seven bovid species: H. duvernoyi, Palaeotagus rouenii, a Palaeotraginae indet., T. amalthea, ?Miotragocerus sp., Gazella pilgrimi, Gazella cf. capricornis, N. planicornis, Palaeoreas lindermayeri, and Palaeoryx cf. pallasi. As a whole, this artiodactyl association clearly foreshadows a Turolian age, and sharply contrasts that from Nikiti 1 (e.g., Koufos, 2006; Koufos et al., 2016a; Table 2), with which it shares only H. duvernoyi. At Nikiti 2, the great abundance of Nisidorcas together with Gazella pilgrimi and boselaphines fits biochronologically better with early to early middle Turolian faunas from continental Greece,

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GEOBIO-736; No. of Pages 16 D.S. Kostopoulos / Geobios xxx (2016) xxx–xxx Table 2 Taxonomic composition of the late Miocene artiodactyl assemblages from Nikiti 1 (NKT), Nikiti 2 (NIK) and Ravin des Zouaves 5 (RZO). : occurrence of a taxon in a site. Artiodactyla Suidae Microstonyx major Propotamochoerus hysudricus Giraffidae Helladotherium duvernoyi Bohlinia attica Palaeotragus sp. Palaeotragus rouenii Palaeotraginae indet. Bovidae Miotragocerus pannoniae Miotragocerus sp. Tragoportax amalthea Prostrepsiceros houtumschindleri Prostrepsiceros axiosi Prostrepsiceros rotundicornis Palaeoreas lindermayeri Palaeoreas zouavei Nisidorcas planicornis Gazella pilgrimi Gazella capricornis Hispanodorcas orientalis

NKT

NIK



  

RZO  





 

 

? 

? 

cf.

      

   sp.

cf.

especially those from the Axios Valley (Table 2). Indeed, Ravin de Zouaves 5 (RZO) and Nikiti 2 (NIK) share six artiodactyl species and one genus (Palaeoreas) in common (Table 2). Nevertheless, cranial and horn-core features of Nisidorcas from Nikiti 2 appear more primitive than those from Ravin de Zouaves 5 and Vathylakkos, suggesting an earlier age. In agreement, Palaeoreas from Nikiti 2 is less advanced than that from Hadjidimovo (Bulgaria) and Pikermi (Greece), of late early and middle Turolian age, respectively. Hence, an early Turolian age, possibly slightly older than Ravin de Zouaves 5 ( 8.2 Ma; Koufos, 2006) is suggested for the Nikiti 2 fauna, in agreement with several other taxa found in this locality (Koufos et al., 2016b). Similarly to mammal assemblages from the western and eastern edges of the late Miocene Eastern Parathethyan domain and in contrast to Anatolia (Kostopoulos and Bernor, 2011; Kostopoulos and Karaku¨tu¨k, 2015), small bovids (< 50 kg) prevail over medium and large ones in the Nikiti 2 bovid association (Fig. 16), and boselaphines exceed significantly caprines (absent in

Fig. 16. Taxonomic composition (%) of the Nikiti 2 artiodactyl assemblage based on the Minimum Number of Individuals (MNI, n = 79) and Number of Identified Specimens (NISP; n = 209).

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NIK). Apart from Nisidorcas, which clearly dominates (> 60%), the composition of the rest of the Nikiti 2 artiodactyl assemblage is rather uniform with all other taxa represented by  10% each (using either MNI or NISP). This biased spectrum towards Nisidorcas cannot, however, be considered as a taphonomic artifact, as similar-sized taxa like gazelles are much less common, and as most taphonomic indications (representation of all skeletal elements, absence of preferential orientation of long bones, high percentage of partly articulated skeletons, etc.) speak for a primary fossil concentration without or with short transportation from the ecocenosis to the taphocenosis. A previous study (Kostopoulos, 2000) showed that Nisidorcas ecomorphology corresponds to a broken cover habitat on hilly terrains, an environment that seems to fit quite well the entire artiodactyl association of the Nikiti 2 mammal assemblage (Merceron et al., 2016). Acknowledgements I am deeply thankful to Denis Geraads (MNHN Paris) and Nikolai Spassov (NHM Sofia) for sharing photos and measurements of Bulgarian material. Thanks are also due to George D. Koufos, Ioanna Sylvestrou, Dora Vlachou, George Konidaris and many students who helped all these years in the field and lab. I am also indebted to Faysal Bibi (Museum fu¨r Naturkunde, Humboldt University Berlin), Nikolai Spassov (NHM Sofia), Zhang Zhaoqun (Chinese Academy of Sciences) and Gilles Escarguel (Editor in chief of Geobios; Universite´ Lyon 1) for useful comments, suggestions and linguistic improvements.

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