Article originalThe 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
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).
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