Ravin De Research Articles

A review of Palaeogiraffa (Giraffidae, Mammalia) from the Vallesian of the Eastern Mediterranean

Palaeogiraffa is a poorly understood giraffid genus from the Vallesian of Axios Valley (Northern Greece), Turkish Thrace and Central Turkey. The Axios Valley remains come from three non-contemporaneous Vallesian faunas from the localities: Pentalophos, Ravin de la Pluie, and Xirochori, whereas the Turkish remains come from the Vallesian faunas of Middle Sinap, Yulafli, and Küçükçekmece. Remains from Pentalophos and Ravin de la Pluie were described as congeneric with the Iberian Deccenatherium, although they were later attributed to a new, distinct genus known by three different species: Palaeogiraffa macedoniae, Palaeogiraffa pamiri, and Palaeogiraffa major. The type species of the genus, Pg. macedoniae, was based on the most abundant Pentalophos material, while the rest of the species was established mainly on size difference criteria. Here we provide a review of the Palaeogiraffa material in order to (i) test its validity, (ii) re-discuss hypotheses about the genus phylogenetic relationships with other Late Miocene giraffids, and (iii) re-explore intrageneric diversity and number of recognized species based on the available Eastern Mediterranean samples. We confirm Palaeogiraffa as a distinct genus from other Late Miocene giraffids based on its unique proportions, as well as its dental and postcranial morphology. Furthermore, we endorse the likelihood of Palaeogiraffa being phylogenetically linked to the sivatherine lineage, based mainly on its dental and metatarsal morphological features. The generic attribution of specimens from Xirochori, Ravin de la Pluie, Middle Sinap, Yulafli and Küçükçekmece to Palaeogiraffa is adequately supported by the present data. Nevertheless, the recognition of three species is more open to discussion, though they should be retained until more material is available.

Two new Later Stone Age sites from the Final Pleistocene in the Falémé Valley, eastern Senegal.

The understanding of cultural dynamics at work at the end of the Final Pleistocene in West Africa suffers from a significant lack of excavated and dated sites, particularly in the Sahelian and Sudanian ecozones. While the Later Stone Age shows varied behavioral developments in different parts of the continent, the chrono-cultural framework of this period remains largely unknown in West Africa. We report on archaeological, geomorphological, and chronological research on two Final Pleistocene Later Stone Age sites in the Falémé Valley, eastern Senegal. Optically stimulated luminescence ages place the site of Toumboura I-2017 between 17 ± 1 and 16 ± 1 ka and the Ravin de Sansandé site between 13 ± 1 ka and 12 ± 1.1 ka. The excavated lithics show typical Later Stone Age industries, characterized by chaînes opératoires of core reduction mainly producing flakes and bladelets as well as blades and laminar flakes. Segments dominate the toolkits but a few backed bladelets and end-scrapers on flake blanks were recognized. Local raw materials were used, with a preference for chert and quartz, as well as greywacke. These Later Stone Age lithic assemblages are the oldest known in Senegal so far and add to the small number of sites known in West Africa for this period, which are mainly located farther south, in sub-tropical ecozones. The Later Stone Age sites of the Falémé Valley are contemporaneous with typical Middle Stone Age technologies in Senegal dated to at least the Pleistocene/Holocene transition. Our results thus provide new archaeological evidence highlighting the complex cultural processes at work during the Final Pleistocene in West Africa.

The Late Miocene hominoid Ouranopithecus macedoniensis (Bonis, Bouvrain, Geraads & Melentis, 1974): maxillary deciduous dentition and virtual reconstruction of the unerupted permanent teeth

The hominoid Ouranopithecus macedoniensis (Bonis, Bouvrain, Geraads & Melentis, 1974) is known from three Late Miocene localities in Greece. All are correlated with late Vallesian, MN10; more precisely, they are dated between 9.6 and 8.7 Ma. During the last 50 years several fossils of this hominoid have been recovered. The present article describes the first specimen with upper deciduous teeth, recovered from the locality Ravin de la Pluie in Axios Valley (Macedonia, Greece). The upper deciduous dentition of O. macedoniensis is characterised by a small canine relative to the length of the deciduous premolars, with a rounded occlusal crown outline, a trapezoidal dP3 with metacone and a rudimentary hypocone, a sub-squared dP4, and the absence of cingulum. The taxonomic and sexual attribution of the studied material is based on the preserved and virtually reconstructed permanent dentition, using high resolution microcomputed tomography. The morphological and metrical comparisons of the permanent dentition with the available sample of O. macedoniensis, as well as the monospecific character and the strong sexual dimorphism of the Ravin de la Pluie sample, allow its attribution to a female sub-adult individual of this taxon. The upper deciduous teeth of Ouranopithecus macedoniensis preserve some morphological features, that may have a phylogenetic value, but this hypothesis requires further investigation as the available material of all fossil taxa is poor.

The Vallesian large Palaeotragus Gaudry, 1861 (Mammalia: Giraffidae) from Northern Greece

We herein re-discuss the systematics of the Late Miocene representatives of the most common but poorly documented Eurasian giraffid genus Palaeotragus on the occasion of the review and description of new samples from the Vallesian faunas of Northern Greece. Our results detect five Late Miocene Palaeotragus morphotypes, recognizing at least four species. The so called ‘small-sized palaeotrages’ are represented by the type species P. rouenii and the Chinese P. microdon, whereas the validity of P. pavlowae from Grebeniki (Ukraine) is doubted. ‘Large-sized palaeotrages’ are mainly represented by P. coelophrys (synonym of P. expectans, P. borissiaki, P. hoffstetteri, P. quadricornis, and probably P. moldavicus), a species that thrived in the peri- Black Sea territories during Vallesian and survived during Turolian in the Irano-Anatolian domain, likely by adopting a more robust appearance. Large palaeotrages from the Vallesian faunas of Pentalophos and Ravin de la Pluie (Axios Valley, Greece) are identidied as P. coelophrys with certain confidence. The Vallesian P. berislavicus from Berislav (Ukraine) has intermediate morphometric features between P. rouenii and P. coelophrys and it is, therefore, recognized as a most probably valid species. The latest Vallesian Nikiti-1 (Chalkidiki peninsula) large palaeotrage shares many morphometric features with P. berislavicus, suggesting that the species may have invaded Balkans by the end of Vallesian and possibly survived there until middle Turolian. The Late Miocene Palaeotragus asiaticus from Central Asia is a quite problematic species; it appears closely related to the Turolian equivalent P. cf. coelophrys from China and both may be linked to the older Berislav taxon.

The earliest known crown-Testudo tortoise from the late Miocene (Vallesian, 9 Ma) of Greece.

We here report on fossil remains of the earliest known crown-Testudo, an extant clade of Mediterranean testudinid tortoises from the late Miocene (Vallesian, MN 10) from the hominoid locality Ravin de la Pluie (RPl) in Greece. The material studied is a small, nearly complete carapace with a clearly distinct hypo-xiphiplastral hinge. This supports the sensu stricto generic assignment. This new terrestrial testudinid specimen is characterized by a possible tectiform, narrow, elongated shell with a pentagonal pygal and a long, posteriorly elevated, lenticular and rounded dorsal epiplastral lip. These unique features differ from those of other known Mediterranean hinged forms and allow the erection of the new species Testudo hellenica sp. nov. This taxon is phylogenetically close to two Greek species, the extant T. marginata and the fossil T. marmorum (Turolian, around 7.3 Ma). This record provides evidence for the first appearance of the genus Testudo sensu stricto at a minimum age of 9 Ma.

Palaeo- and rock-magnetic investigations across Jurassic-Cretaceous boundary at St Bertrand’s Spring, Drôme, France: applications to magnetostratigraphy

Palaeo- and rock-magnetic investigations of the St Bertrand’s Spring (Le Ravin de Font de St Bertrand) locality in France were carried out in order to contribute to, and improve, the stratigraphy of the Jurassic-Cretaceous boundary interval. Magnetic susceptibility shows slightly diamagnetic behaviour in the lowermost part of the profile and an increase (paramagnetic) towards its middle and upper parts. Rock-magnetic measurements throughout the section show magnetite as the main magnetic fraction, together with traces of hematite. Additionally, thermal demagnetization indicates the presence of goethite. Our magnetostratigraphy indicates three normal/reversed polarity sequences; possibly encompassing the magnetozones M19r to the M17n. This suggests that the St Bertrand section straddles the Tithonian/Berriasian boundary and reaches the middle Berriasian sensu lato.

Lizards and snakes from the late Miocene hominoid locality of Ravin de la Pluie (Axios Valley, Greece)

We here describe lizards and snakes from the late Miocene (MN 10) of Ravin de la Pluie, near Thessaloniki, Greece, a locality widely known for its hominoid primate Ouranopithecus macedoniensis. The new finds comprise two large-sized lizards (a probable anguine and a varanid) and two snakes (an elapid and a small-sized “colubrine”). Even if the material is represented by few specimens, this is the first record of squamates from the late Miocene MN 10 biozone of southeastern Europe and the third only for the whole continent. The importance of the varanid vertebrae for systematic attributions is discussed. The new varanid limb elements described herein rank among the few such specimens in the fossil record of monitor lizards. Judging from the new and previously published varanid appendicular material, we suggest that Neogene monitor lizards from Europe possessed comparatively short and robustly built limbs. Distinctive scars on one of the limb elements are interpreted as bite marks of a predator or scavenger, offering insights on the palaeoecology of the herpetofauna of the locality.

Upper incisor morphology of the Late Miocene hominoid <i>Ouranopithecus macedoniensis</i> from Axios Valley (Macedonia, Greece)

The upper incisor lingual morphology of the late Miocene Greek hominoid Ouranopithecus macedoniensis was almost unknown, as the described earlier maxillary remains preserve only worn incisors. During the most recent excavations in the type locality of Ouranopithecus, Ravin de la Pluie (RPl) of Axios Valley (Macedonia, Greece), four little-worn upper central incisors were recovered. This material and a few additional worn upper incisors, discovered recently, are described and compared in this article. Even though a morphological comparison with the old RPl material, lacking unworn or little worn incisors, is impossible, the metrical comparison and the monospecific character of the RPl hominoid sample suggest that the described incisors can be assigned to Ouranopithecus macedoniensis. The described upper central incisors are separated in two size-groups which in general have similar morphology except for some minor differences such as the presence of a pronounced mesial lingual pillar in the small-sized specimens. The observed significant size difference among the studied incisors is attributed to the strong sexual dimorphism of Ouranopithecus, which is also well expressed in the other teeth. The lingual morphology of the upper incisors of Ouranopithecus are not identical to those of extant great apes, though they have some similarities with those of the African great apes (Gorilla and Pan), while they are clearly different from those of the Asian great ape (Pongo). Even though they have some morphological similarities, the O. macedoniensis central incisors are probably not identical to those of the Eurasian Miocene hominoids; the most similar central incisor is that of Ouranopithecus turkae. Among the known African Miocene hominoids, Nakalipithecus upper central incisor is quite similar in morphology and size to that of Ouranopithecus.

New Material of the Hominoid Ouranopithecus macedoniensis from the Late Miocene of the Axios Valley (Macedonia, Greece) with Some Remarks on Its Sexual Dimorphism

New hominoid teeth from the late Miocene locality Ravin de la Pluie (RPl) of the Axios Valley (Macedonia, Greece) are studied in this article. Their morphology, dimensions and proportions are similar to the hominoid Ouranopithecus macedoniensis, allowing their attribution to this taxon. The studied material provides some new morphological characters for the female P³ (small asymmetry, small mesiobuccal crown projection, paracone higher than protocone) and the lower canine (mesial groove: large in the male and small in the female, distobuccal fovea: large in the female and small in the male). The new material enriches the collection of O. macedoniensis. The estimated degree of sexual dimorphism of the RPl O. macedoniensis, calculated by the multivariate size dimorphism method, is compared with those of extant hominoids (Gorilla, Pan, Pongo) and of the late Miocene Lufengpithecus lufengensis from China, which is considered as more dimorphic than any living hominoid. The results suggest that Ouranopithecus multivariate size dimorphism for the premolar, molar and post-canine row is similar to those of Pongo and Lufengpithecus, slightly higher than that of Gorilla and clearly higher than that of Pan. Therefore, O. macedoniensis is apparently one of the most sexually dimorphic hominoids and the RPl assemblage is monospecific.

Equidae

The locality Küçükçekmece includes a limited hipparion sample mainly consisting of isolated teeth and some postcranial remains. Using various methods, the teeth and postcranials are separated in two different size groups. The first group, characterized by large size, rich enamel plication, deep and thin plis, double-triple pli caballin, oval protocone with flattened lingual border, absence of lingual hypoconal groove, short and robust metapodials, is related to H. sebastopolitanum Borissiak, 1914 from Sebastopol, Ukraine and it is referred to as H. aff. sebastopolitanum. The second group with very large size, very rich enamel plication, very deep plis, double-multiple pli caballin, oval protocone with flattened lingual border, presence of lingual hypoconal groove, plicated or crenulated flexid's borders in the lower teeth, short and robust metapodials larger than those of the first form, is closer to H. giganteum Gromova, 1952 from Grebeniki, Ukraine and it is identified here as H. aff. giganteum. Hipparion sebastopolitanum and H. giganteum are known from the Ukrainian early and late Vallesian localities of Sebastopol and Grebeniki, respectively. The similarities of the first form with H. cf. sebastopolitanum from the Vallesian localities Pentalophos 1 and Ravin de la Pluie in Axios Valley (Macedonia, Greece), and of the second form with H. aff. giganteum from the terminal Vallesian locality of Nikiti 1 (Chalkidiki, Greece) support a Vallesian age for the Küçükçekmece fauna; more precisely an age from upper early Vallesian to the end of Vallesian is quite possible. The limited data of the dental and postcranial morphology of the Küçükçekmece hipparions indicate a probable forest and warm/humid environment.

Synthesis

The study of the new and old collections of the Nikiti vertebrate localities included in this volume provides several new evidences for the taxonomy, composition, chronology and palaeoecology of these late Miocene mammal faunas. The faunal list of both primate bearing localities is enriched and improved by addition of new taxa and revision of older identifications; two new species are recognized in the Nikiti 2 (NIK) fauna (two hipparionine horses) and one subspecies from Nikiti 1 (NKT) is upgraded to the species level. The NKT and NIK faunas include 15 and 19 mammalian taxa, respectively. The mammalian faunas from both sites consist of almost the same families; the absence of some of them in NKT or NIK is most probably artificial. The chronology of the Nikiti mammal assemblages is based on biochronological data only, which allow the correlation of NKT to the terminal Vallesian (between 9.3 and 8.7Ma) and that of NIK to the earliest Turolian (between 8.7 and 8.3Ma). Concerning their age in relation with other neighboring mammal assemblages, NKT is younger than Ravin de la Pluie (Axios Valley, Greece) and isochronous or slightly older than Grebeniki (Ukraine). The NIK assemblage is older than Ravin des Zouaves 5 (Axios Valley) and Sivas (Turkey), dated at ∼8.2Ma and ∼8.3Ma, respectively. The available morphoecological, dental microwear-mesowear, and enamel isotopic analyses of the herbivores, as well as study of the phytolites suggest an open-light cover landscape for both localities. As documented previously in the Axios Valley, the Nikiti mammal fauna exhibits a significant reorganization through the Vallesian/Turolian boundary, including the Ouranopithecus/Mesopithecus replacement. However, this faunistic event is not consistent with the results of independent studies (isotope, dental wear, etc.) that fail to confirm significant climatic or vegetational changes across the same time interval.

Carnivora

The carnivoran sample from Nikiti 2 (NIK) is relatively poor, including a small number of specimens, mainly postcranials. The best-preserved specimen is a skull and mandible, as well as some bones of the common late Miocene hyaenid Adcrocuta eximia. The sample of A. eximia is similar to the subspecies A. eximia eximia from the Turolian of Greece. A second hyaenid species is represented by an upper deciduous dentition; it is smaller than the corresponding dentition of A. eximia from the Greek localities of Pikermi, Perivolaki and Ravin de la Pluie. It could belong to a smaller hyaenid, such as the ictitheres Hyaenictitherium wongii and Ictitherium robustum, or the hyaenid Lycyaena chaeretis; all of these taxa have been already found elsewhere in association with A. eximia. The machairodontid Amphimachairodus giganteus is represented by some postcranial elements. The simultaneous occurrence of A. e. eximia and A. giganteus indicates a Turolian age for the NIK fauna. A. eximia is a common and widespread taxon ranging in Eurasia from the Vallesian to the end of the Turolian, making it useful as a biostratigraphic marker for the beginning of the late Miocene. A. giganteus is known in Europe and eastern Asia during the Turolian.

Hipparion macedonicum revisited: New data on evolution of hipparionine horses from the Late Miocene of Greece

The new expeditions to the Axios Valley (Macedonia, Greece) provided a new set of fossils from the various localities. Among the material collected from the Late Miocene hominoid bearing mammal locality Ravin de la Pluie (RPl) were some remains of Hipparion macedonicum, which was originally described from this locality. The most important is the skull and associated mandible, the first from the type locality. The new material is compared with the previously collected material of the taxon from the Vallesian and Turolian levels of Greece, as well as with corresponding material from Eurasia. The RPl skull is compared with the type skull of H. matthwei, a taxon, which several times has been referred as synonym to H. macedonicum. The comparison indicated several differences which distinguish the two species. Hipparion macedonicum has a continuous stratigraphic range from the Vallesian to middle Turolian and it is possibly present in the late Turolian. The comparison of the chronologically different samples of...

A new global database of δ98Mo in molybdenites: A literature review and new data

Isotopic compositions of Mo in molybdenites were used for deciphering a possible genetic link between isotopic variations and mineralizing processes, based on a worldwide molybdenite databank. We compared the δ98/95Mo (hereafter referred as δ98MoNIST) of 391 molybdenite samples (193 from the literature, 198 for this study) from different localities, different types of occurrences and different ages. The 198 molybdenite samples we analysed represent various types of mineralization in 6 granites, 11 pegmatites, 6 perigranitic veins, 2 greisen, 28 porphyry deposits, 5 skarns, 1 IOCG, and 9 Alpine-type fissure veins, with ages varying from 5Ma to 2.7Ga. The Mo isotopic composition was determined with an MC-ICP-MS Neptune after aqua regia dissolution and adjustment to [Mo]=1μg·ml−1. Mass bias was corrected by using Zr as dopant and standard-sample-standard bracketing. The δ98MoNIST ratios were normalized to NIST3134. External reproducibility is 0.07‰ (2σ). The overall range of the δ98MoNIST ratio in the 391 molybdenite samples varied from −1.62 to 2.27‰, being higher for molybdenite formed in Alpine-type veins, greisen, perigranitic veins and IOCG, than for that in granite, pegmatite, porphyry deposits and skarns. The crystallization temperature can explain some of these differences, as polymetallic Alpine-type fissure veins broadly crystallize at lower temperatures than granite, pegmatite and porphyry deposits. For some occurrences the δ98MoNIST was determined on several molybdenite samples, showing variability at occurrence scale. For example, in the Azegour skarn (Morocco) the δ98MoNIST varies from −0.60 to 0.42‰ (n=29), and in “Ravin de la Ruine” Alpine-type fissure veins (France) the variation is from −0.08 to 0.77‰ (n=3). No correlation is seen between δ98MoNIST and the age of the deposits.

First discovery of postcranial bones of Ouranopithecus macedoniensis (Primates, Hominoidea) from the late Miocene of Macedonia (Greece)

Two well preserved phalanges, one proximal and one intermediate, are identified and described. They constitute the first postcrania ever described of Ouranopithecus macedoniensis (Primates, Hominoidea) from the late Miocene locality of Ravin de la Pluie (RPl), Macedonia (Greece). They are isolated specimens, and the only ones known for their genus. The hypotheses that these specimens derive either from the hand or from the foot were tested. Comparisons with living apes of known positional behaviour were made to assess the functional signal in these specimens. The proximal phalanx, either manual or pedal, closely matches the proximal phalanges of terrestrial quadrupedal primates or the bipedal primate Homo. With respect to the intermediate phalanx we show that it closely matches phalanges of quadrupedal terrestrial primates. The terrestriality of Ouranopithecus, here reconstructed from phalangeal remains, is in agreement with our previous results of dental studies (thick check teeth enamel and microwear pattern of incisors and molars), which indicate that it was a hard object feeder living near the ground.

Comment on: “Exhumation of subcontinental mantle rocks: evidence from ultramafic-bearing clastic deposits nearby the Lherz peridotite body, French Pyrenees” by C. Clerc et al. (Bull. Soc. géol. Fr., 2012, 183, 5, 443–459)

In their recently published paper, C. Clerc et al . [2012] describe the geological setting of the Etang de Lherz peridotite body and give a short review of the associated clastic deposits, proposing a detailed analysis of the breccias cropping out in a limited area east of the lherzolite massif. The cartographic, sedimentological and metamorphic evidence leads the authors to regard these breccias as a shallow marine to lacustrine equivalent of the Albo-Cenomanian “Flysch noir” of the Pyrenees. This interpretation backs up their previous attribution of the breccias to a thick submarine clastic formation, of Albo-Cenomanian age, which incorporates olistoliths derived from the famous Etang de Lherz lherzolite and its neighbouring bodies. Clerc et al. firstly present a somewhat theoretical discussion concerning a 3D reconstruction of the depositional environment of the Lherz breccias, along with a possible scenario for their origin and structural history, which then leads onto three possible scenarios for North-Pyrenean rifting. The authors use this approach to support their previous conclusions, i.e.: the heterometric peridotite fragments contained in the Lherz breccias (from sand-sized debris up to olistoliths) provide clear evidence for an Albo-Cenomanian subaqueous exhumation of the subcontinental mantle at the borders or on the floor of the North-Pyrenean Aulus basin. Our comment focuses on a number of geological details concerning the Lherz breccias and their local or regional context, which the authors fail to take into account, either in their present paper or in earlier studies. Indeed, these details are of crucial interest since they call into question the assumption of a sedimentary origin for the Lherz breccias and, consequently, the exhumation of mantle in the Aulus basin. This map covers an area of 100×200 m located on the western side of the middle part of the so called “Ravin de Paumeres” [or “Ravin de l’Homme-mort” …

Late Miocene Proboscidea (Mammalia) from Macedonia and Samos Island, Greece: preliminary results

The abundant Late Miocene proboscidean remains of Greece have never been studied in detail and compared with those of Eurasia in order to determine their taxonomy and their biostratigraphical and palaeoecological significance. The first results of such study are given in this article. During the past decades, several new proboscidean specimens have been added to the old collections, significantly enriching the available material. The Axios Valley (Macedonia, Greece) proboscidean fossils belong mainly to two species of Choerolophodon: C. anatolicus of early Vallesian age and C. pentelici of late Vallesian–Turolian age. Deinotherium giganteum is rare and recognized only in the late Vallesian locality Ravin de la Pluie of Axios Valley. A zygodont form has also been identified in the Turolian of Axios Valley, attributed to “Mammut” sp. The Late Miocene localities of Nikiti (Macedonia, Greece) revealed several remains of C. pentelici, which are similar to the Turolian ones of Axios Valley. The Samos proboscidean collection includes C. pentelici, “Tetralophodon” atticus, “Mammut” sp. and Deinotherium gigantissimum. The taxonomy of the Late Miocene peri-Mediterranean Choerolophodon is given, and the biostratigraphy and palaeoecology of the Greek Late Miocene proboscideans are discussed.

New material of Carnivora (Mammalia) from the Late Miocene of Axios Valley, Macedonia, Greece

A new collection of carnivores from the Late Miocene deposits of Axios Valley, Macedonia, Greece was collected in 2004–2009 at the localities named Xirochori 1 (XIR) and Ravin de la Pluie (RPl), dated to the Late Vallesian (MN 10), as well as at Ravin des Zouaves 5 (RZO), dated to the Early Turolian (MN 11). The studied material is described and compared morphologically and biometrically with other materials from Greece and its neighboring area. The following species are determined: XIR: Dinocrocuta sp.; RPl: Eomellivora wimani, Adcrocuta eximia leptoryncha, ?Hyaenictis sp., Metailurus parvulus; RZO: Plioviverrops orbignyi, Machairodus giganteus. This collection is interesting because it includes the upper deciduous dentition of A. eximia and Dinocrocuta, which are only known from limited material until now, especially the latter taxon. E. wimani is recognized for the first time in Greece and the Balkans, while M. parvulus is identified for the first time in the Vallesian of Greece. The possible presence of Hyaenictis in RPl is the first indication of its presence in the Vallesian. The guild structure diagrams of the carnivore fauna from the three studied localities provide some indications of their palaeoenvironment, which fits quite well with the results of previous studies on the palaeoenvironments of these localities and of the wider Eastern Mediterranean region.

A new protictithere from the late Miocene hominoid locality Ravin de la Pluie of Axios Valley (Macedonia, Greece)

This article concerns the study of a protictithere from the late Miocene of Axios Valley (Macedonia, Greece). The material was found in the hominoid-bearing mammal locality Ravin de la Pluie (RPl), which is corellated with the late Vallesian, MN 10. The available material, including maxillary and mandibular elements, is described and compared morphologically and metrically with the known Eurasian protictitheres. The small size, the low cusps(-ids) of the teeth, the protocone of the P4, which is in line with the mesial border of the parastyle, the large molars (especially the M2), the small buccal projection of the M1 paracone, the strongly molarized p4, the strong metaconid and large talonid with high entoconid of the m1 separate the RPl protictithere from the known Eurasian protictitheres and allow the erection of a new species, named P. thessalonikensis n. sp.

Contribution to the systematics and phylogeny ofProstrepsiceros vallesiensisBouvrain, 1982 (Mammalia, Bovidae)

ABSTRACT The Vallesian heteronymous spiral-horned antelope Prostrepsiceros vallesiensis is revised according to some new specimens from the type locality, Ravin de la Pluie, Northern Greece. The new frontlet and isolated horn-core are larger, display weaker horn-core compression, and looser spiralling than in the holotype specimen of P. vallesiensis. These morphological differences are interpreted as representing sexual dimorphism, with the holotype being a female individual of the species. Morphological comparisons and a cladistic analysis of several late Miocene spiral horned antelopes allow for recognizing P. vallesiensis as the most likely ancestor of P. vinayaki, P. libycus and Dytikodorcas, while the P. houtumschindleri lineage probably originated from a pre- P. vallesiensis stock.