Elsevier

Annales de Paléontologie

Volume 91, Issue 3, July–September 2005, Pages 227-240
Annales de Paléontologie

Article original
The Late Miocene hominoids Ouranopithecus and Graecopithecus. Implications about their relationships and taxonomyLes hominoïdes du Miocène supérieur Ouranopithecus et Graecopithecus. Implications de leurs relations et taxonomie

https://doi.org/10.1016/j.annpal.2005.05.001Get rights and content

Abstract

Two Late Miocene hominoids are known from Greece. The first, Graecopithecus freybergi, is known by a single mandible with the worn m2 from the locality of Pyrgos Vassilissis, near Athens. The other, Ouranopithecus macedoniensis, is known from Axios Valley and Chalkidiki (Macedonia, Greece) by a partial skull and a set of maxillary and mandibular remains. Some authors consider these two hominoids as synonyms and in the present article a detailed comparison of them is given. The morphology and size of the symphysis, the more robust mandible, the more open dental arcade of Ouranopithecus distinguishes it clearly from Graecopithecus. Moreover, the incompleteness of the mandible of Graecopithecus with the doubtful morphology and size, the limited material and the uncertain geological age of the locality cannot allow precise and clear comparisons with the rest extant and extinct hominoids. Thus in our opinion there are not enough data to support the similarity, and therefore, the synonymy of the two genera. The Pyrgos mandible must remain as a separate and isolated genus with one species, which only includes this sole mandible.

Résumé

Deux genres de primates hominoïdes ont été décrits dans le Miocène supérieur de Grèce. L'un, Graecopithecus freybergi, provient de la localité de Pyrgos Vassilissis, près d'Athènes, dont l'âge n'est pas connu avec précision. Il n'est représenté que par une mandibule portant seulement une m2 complètement usée et les racines d'autres dents. L'autre, Ouranopithecus macedoniensis, provient de gisements situés dans la basse vallée de l'Axios et en Chalcidique (Macédoine, Grèce) qui sont datés du Vallésien supérieur. Il est représenté par une portion de crâne et par tout un ensemble de maxillaires, de mandibules et de dents isolées. Certains auteurs ont estimé que les deux genres étaient synonymes. Le but de cet article est de comparer avec soin le matériel appartenant à ces deux taxons, tout au moins pour les parties présentes sur la mandibule de Pyrgos Vassilisssis. On remarque que la morphologie de la symphyse est différente, que la mandibule d'Ouranopithecus est plus robuste avec une arcade dentaire plus ouverte et que les deux ensembles se distinguent clairement. Étant donné l'état de conservation de la mandibule de Graecopithecus il n'est pas possible de pousser les investigations plus loin. Cependant, il nous semble que les données pour rassembler les deux genres sont insuffisantes et que leur rapprochement par certains auteurs découle surtout de leur position géographique en Grèce bien que le site de Pyrgos-Vassilis soit situé à plusieurs centaines de kilomètres des gisements de Macédoine. Il est donc préférable de conserver Graecopithecus comme un genre séparé contenant une seule espèce fondée sur une unique mandibule incomplète.

Introduction

The hominoid primate Ouranopithecus is widely known from Macedonia, Greece by a quite rich sample of cranial and dental remains. It was originally found in the locality “Ravin de la Pluie” (RPl) of Axios Valley; later it was recognized in the localities “Xirochori-1” (XIR) of Axios Valley too and “Nikiti-1” (NKT), Chalkidiki Peninsula (de Bonis et al., 1990, de Bonis et al., 1974; Koufos, 1993, Koufos, 1995). All Ouranopithecus bearing localities have been dated to Late Vallesian, MN 10 with minor age differences between them. The magnetostratigraphic data suggest an age ~9.6 Ma for XIR and ~9.3 Ma for RPl (de Bonis and Koufos, 1999; Sen et al., 2000). The faunal data from NKT indicate that it is younger than the previous two with an age between 9.3 and 8.7 Ma (Koufos, 2000). The study and analysis of these faunas indicate a relatively more open environment (de Bonis et al., 1992; de Bonis and Koufos, 1999; Koufos, in press). The thick enamel and the dental microwear of Ouranopithecus suggest a hard-food eater (Ungar, 1996).

A hominoid mandible has been found in the Greek locality “Pyrgos Vassilissis”, near Athens. It was originally referred to the cercopithecoid Mesopithecus pentelicus but later it was described under the name Graecopithecus freybergi (von Koenigswald, 1972). The associated faunule includes the following species: Choerolophodon pentelici, Hipparion mediterraneum? “Dicerorhinus orientalis”, Bohlinia attica, Helladotherium duvernoyi? Tragoportax amalthea, Gazella? deperdita, Gazella? cf. gaudryi. This fauna cannot allow a more precise age than Late Miocene (Koufos, 1995; de Bonis and Koufos, 1999). The Pyrgos mandible lacks the external symphysis, the inferior limits of the mandibular bodies are damaged and the anterior dentition is absent. The cheek teeth are almost absent; the right worn m2, the lingual surface of the right p4 and the base of the crown of the right m1 are preserved. The occlusal border of the right m2 is also partially damaged.

The first unearthed mandible from RPl was described as Dryopithecus macedoniensis n. sp., following the nomenclature of that time and distinguishing the RPl mandible from the known hominoid species (de Bonis et al., 1974). The material found during the later years allowed the erection of the new genus Ouranopithecus (de Bonis and Melentis, 1977), a name used by the authors until now. The species Ouranopithecus macedoniensis and Graecopithecus. freybergi have been synonymized and included both to Ankarapithecus meteai; the genus Ankarapithecus was considered as synonym of Sivapithecus and thus all these taxa included to Sivapithecus meteai (Martin and Andrews, 1984). Later on, Ouranopithecus and Graecopithecus were separated from Sivapithecus and referred as G. freybergi (Andrews et al., 1996), while Cameron (1997) distinguished two species: G. freybergi and “G”. macedoniensis. Few years later the name Ankarapithecus became valuable again separated from Sivapithecus (Alpagut et al., 1996; Begun and Gülec, 1998). Thus, there is a puzzle of the generic and specific names of the Miocene hominoids of Greece and Turkey. We believe that the Pyrgos mandible is so incomplete that it cannot provide the available metrical and morphological characters, in order to synonymize it with any of the known hominoids. In fact, the Ouranopithecus material was found in Greece, as Graecopithecus, but this does not mean that any Greek hominoid must belong to the latter genus. The limited morphology, size and age of the Pyrgos mandible cannot allow certain comparisons and results about this. In the present article we shall try to show the limited morphological characters and comparisons of Graecopithecus, as well as to find the differences between the two genera. This article has been presented in the 5th International Congress on Eastern Mediterranean Geology, Thessaloniki, April 2004.

Section snippets

Comparison of Graecopithecus and Ouranopithecus

The Pyrgos mandible is referred as female from the size of its premolars and the roots of the canine (Martin and Andrews, 1984) but it is impossible to be sure from a single specimen. The root of the canines corresponds to a relatively small canine and this is an indication for a female individual. But, this is not clear, as we did not know the size of the male canine. The more complete m2 can probably provide other indications about the sex of the Pyrgos mandible. In the recent gorillas there

Discussion

The systematic position of Ouranopithecus has been discussed since a long time from the middle of 1980s. Martin and Andrews (1984) discussed the phyletic position of Graecopithecus and for the first time synonymized the two genera. Later on, Andrews et al. (1996) and Cameron (1997) discussed again the position of the two genera and proposed synonymy, while the latter author suggested a separate species for the Ouranopithecus sample. At the same time we gave several data, indicating the

Conclusions

The above given comparison of Graecopithecus and Ouranopithecus shows that the two genera cannot be synonymized. The sole known mandible of Graecopithecus is so incomplete that cannot provide enough and reliable data for certain comparisons to the other extinct and extant hominoids. The incompleteness of the mandible provides limited measurements and thus the metric comparisons are not valuable, in several cases. However, some morphological and metrical characters of the Pyrgos mandible are

Aknowledgements

Many thinks to Dr. Peter Andrews for providing us the cast Graecopithecus. Thanks are also due to an anonymous reviewer for his usefull comments. The excavations in Axios valley and Nikiti have been financed by the French CNRS, the Leakey Foundation for Anthroplogical Research, and the National Science Fondation of USA (program RHOI).

References (21)

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