George D. Koufos

New remains of Mesopithecus (Primates, Cercopithecoidea) from the late Miocene of Macedonia (Greece), with the description of a new species

ABSTRACT Numerous remains of Mesopithecus have recently been recovered from the late Miocene mammalian localities of Macedonia, northern Greece. Three mandibles from the early Turolian are identified as M. delsoni, sp. nov., probably ancestral to M. pentelicus of Pikermi. Others, dated to the latest Turolian, differ from M. pentelicus, but most of the teeth are so worn that they must be identified as Mesopithecus, aff. M. pentelicus. A last set, coming also from the latest Turolian, could possibly be attributed to Mesopithecus, cf. M. monspessulanus. The evolution of the genus Mesopithecus can be used as a biostratigraphic tool in late Miocene continental deposits.

The Phylogenetic and Functional Implications of Ouranopithecus macedoniensis

The anatomy of fossil mammals is the key to understanding their place among beings, living or fossil. Anatomical characters allow one to both recognize phyletic positions and provide hypotheses on the modes, processes, and even causality of evolution. Most anatomical changes must correspond more or less to a change in adaptation. The new features are on the one hand derived, in a phyletic sense, and on the other hand they are adaptations to a new way of life. These new features help to characterize and to distinguish the younger taxa that emerge from a lineage. For a different way of life, organs must have different functions and, generally, a different shape or a different size.

The Neogene cercopithecids (Mammalia, Primates) of Greece

ABSTRACT The presence of the cercopithecids in the Neogene of Greece is known since the beginning of the 19th century. The excavations of the last 20 years increase their number in Greece. The main taxon of the cercopithecids is Mesopithecus Wagner, 1839 found originally in the middle Turolian (MN 12) locality of Pikermi, near Athens. The Pikermi Mesopithecus sample is rich and belongs to a medium-sized form, M. pentelicus Wagner, 1839. Besides the well-known M. pentelicus, two other species were recognized. The new species M. delsoni Bonis, Bouvrain, Geraads & Koufos, 1990, a large-sized form found in the early Turolian (MN 11) locality Ravin des Zouaves-5 of Axios Valley (Macedonia, Greece) has several differences from the type species. In the middle Turolian (MN 12) localities of Vathylakkos (Axios Valley) and Perivolaki (Thessaly) a large to medium-sized Mesopithecus form with “delsoni” and “pentelicus” characters was found and is referred to as M. delsonil pentelicus. A small-sized form named M. cf. monspessulanus was recognized by a mandibular fragment in the late Turolian (MN 13) locality Dytiko-2 of Axios Valley and indicates the early appearance of the taxon at the end of Miocene. The rest of the material found in the late Turolian localities of Dytiko has some differences from the typical M. pentelicus. Moreover, Mesopithecus was traced in several late Miocene localities, indicating its wide distribution in Greece. Two other cercopithecids were also found in the Pliocene of Greece. Dolichopithecus ruscinensis Depéret, 1889 was recognized in the locality of Megalon Emvolon and in Ptolemais Basin (Macedonia, Greece); both are dated to late Ruscinian (MN 15). The second Pliocene cercopithecid is Paradolichopithecus arvernensis (Depéret, 1929), found in the locality of Vatera (Lesvos Island) and dated to late Pliocene. The stratigraphic distribution and the palaeoenvironment of these cercopithecids are also discussed.

Prodeinotherium bavaricum (Proboscidea, Mammalia) from Lesvos island, Greece; the appearance of deinotheres in the eastern Mediterranean

Abstract The island of Lesvos, eastern Aegean, Greece is well known for the Neogene Petrified Forest of Sigri, situated in the southeastern part of island. The Miocene mammals were hitherto unknown in the island and the deinothere studied herein is the first evidence of their presence. The material was found near the village Gavathas in strongly silicified lacustrine marls. The morphological characters of the stutied teeth and their dimensions indicate that they belong to a primitive form of Prodeinotherium bavaricum. This fact together with the available radiometric ages of the volcanic rocks of the area suggests a minimum age of 18.4 Ma corresponding to the upper part of early Miocene or to late MN 3. The arrival and the dispersion of the deinotheres in Eurasia are also discussed and the Lesvos material represents the first known appearance of deinotheres in Europe. They arrived at the end of MN 3 (18.0–19.0 Ma) and they rapidly dispersed into Europe as their first occurrence in France and Spain is dated at the early MN 4 (18.0). Their first appearance in Asia seems to be earlier as they were traced in the Bugti fauna (Pakistan) dated at 20.5 Ma. This suggests an early connection of Africa and Asia before the final closure of the Tethyan Seaway.

Study of the Turolian Hipparions of thelower Axios Valley (Macedonia, Greece): 1. Locality «Ravin des Zouaves-5 (RZO)

The hipparions of the locality «Ravin des Zouaves-5or RZO of the lower Axios Valley are studied. Three different species are found, the large-sized H. proboscideum Studer,, 1911, the medium-sized H. dietrichi (Wehrli,, 1941) and the small-sized H. macedonicum Koufos,, 1984. The material of each species is described and compared with that from other localities. The teeth of the three species are compared using odontograms and the bones using logarithmic ratio diagrams. The age of the locality is also discussed. According to the fauna and the morphological features of the hipparions it dates back to Latest Vallesian-Early Turolian. Tables with measurements and statistics for the teeth and bones of each species are given. The tables will be in the library of Geobios and anyone can ask these from there or from the author.

A new hipparion (Mammalia, Perissodactyla) from the Vallesian (Late Miocene) of Greece

KurzfassungDas kleinwüchsigeHipparion macedonicum n. sp., das für das Vallesium (unteres Obermiozän) Nordgriechenlands charakteristisch ist, unterscheidet sich von den bekannten anderen kleinwiichsigen Hipparionen Eurasiens durch folgende Merkmale: besonders geringe Körpergröße, kurze Symphyse, lange Schnauze, Starke Faltung des Zahnschmelzes, das gut entwickelte Protostylid, den großen m3 und die verlängerte m1-m3-Reihe.AbstractA small hipparion from the Vallesian (early Late Miocene) of Northern Greece is studied. It is coming from the locality “Ravin de la Pluie” of the lower Axios valley, near Thessaloniki. Its characteristics as the small size, the short symphysis, the long snout, the high enamel plication, the well developed protostylid, the large m3 and the elongated m1-m3 series distinguish it from the other known eurasiatic small hipparions, allowed us to derive a new species,Hipparion macedonicum. It lived during Vallesian and can be used as a stratigraphic species for this period.

Preliminary results about thestratigraphy and the palaeoenvironment of Mygdonia Basin, Macedonia, Greece

Abstract Mygdonia basin (Macedonia, Greece) was formed during early—middle Miocene and then it was filled by a series of Neogene Quaternary deposits, which were divided in two groups, Premygdonian and Mygdonian. Three different formations have been distinguished in Premygdonian Group, while eight mammalian and one molluskan fossiliferous sites have been found. Chrysavgi Fm is the oldest, dated to the end of middle Miocene. Gerakarou Fm, consisted mainly of red-beds, is dated to early Pleistocene (late Villafranchian). Platanochori Fm has been dated to the end of early Pleistocene (latest Villafranchian) and represents the transition from Premygdonian to Mygdonian Group. Since now no fossils have been found in Mygdonian Group, in order to give a precise dating. However, its deposition begun in the middle Pleistocene and it is still continued. During the end of middle Miocene the climatic conditions were dry with seasonal rainfalls. The early Pleistocene is characterized as a dry period, while an increase of wetness at the end of the early Pleistocene is obvious.

Three-dimensional analysis of mandibular dental root morphology in hominoids

Although often preserved in the fossil record, mandibular dental roots are rarely used for evolutionary studies. This study qualitatively and quantitatively characterizes the three-dimensional morphology of hominoid dental roots. The sample comprises extant apes as well as two fossil species, Khoratpithecus piriyai and Ouranopithecus macedoniensis. The morphological differences between extant genera are observed, quantified and tested for their potential in systematics. Dental roots are imaged using X-ray computerized tomography, conventional microtomography and synchrotron microtomography. Resulting data attest to the high association between taxonomy and tooth root morphology, both qualitatively and quantitatively. A cladistic analysis based on the dental root characters resulted in a tree topology congruent with the consensus phylogeny of hominoids, suggesting that tooth roots might provide useful information in reconstructing hominoid phylogeny. Finally, the evolution of the dental root morphology in apes is discussed.

Locomotor evolution of Mesopithecus (Primates: Colobinae) from Greece: evidence from selected astragalar characters

This paper reports our investigations into functional aspects of the astragalus of four samples of the genus Mesopithecus from Greece. More particularly, it aims to infer substrate preferences of M.delsoni/pentelicus from the Middle Turolian site of Perivolaki (central Greece), M. pentelicus from the late Middle Turolian site of Pikermi (southern Greece), and M. cf. pentelicus and M. cf. monspessulanus from the Late Turolian site of Dytiko (northern Greece). For these purposes, selected astragalar functional features, such as trochlea wedging, proximal facet curvature, and head rotation were expressed as linear measurements on both fossil and selected extant colobines. The size-adjusted measurements were used for univariate comparisons as well as a multivariate principal components analysis. Both approaches revealed that the selected characters were able to discriminate between extant arboreal and semiterrestrial colobines, but all fossil forms presented mosaic morphology. Thus, the oldest representative, M.delsoni/pentelicus was reconstructed as mainly semiterrestrial. On the other hand, the astragalus of M. pentelicus appeared to reflect semiterrestrial habits with a moderate adaptation to arboreality. Similar habitus reconstruction was allocated to the more recent M. cf. pentelicus, whereas the sympatric and synchronous M. cf. monspessulanus showed semiterrestrial adaptations with a slight preference of terrestrial substrates. The results mainly conform to paleoenvironmental reconstructions of the fossiliferous localities and denote that Mesopithecus was mainly a semiterrestrial radiation throughout its evolutionary history, with differential rates of use between arboreal and terrestrial substrates. These adaptations could have promoted the dispersal of the genus throughout Eurasia during the latest Miocene and Early Pliocene.

Feeding habits of the first European colobine, Mesopithecus (Mammalia, Primates): evidence from a comparative dental microwear analysis with modern cercopithecids

ABSTRACT This study aims to characterize the feeding habits of the first European colobines, Mesopithecus (late Miocene), through the analysis of its molar microwear pattern. Fifty-seven adult individuals of Mesopithecus (from Greece and Bulgaria) are compared to 162 wild-shot specimens representing nine modern species of African and Asian cercopithecids. Through the combination of a principal component analysis and analyses of variances, Mesopithecus displays significant differences with its extant colobine relatives. Actually, there is no significant difference between the Mesopithecus samples and Chlorocebus aethiops and Papio hamadryas. An intra-specific test dedicated to Mesopithecus pentelicus reveals no difference between the Bulgarian and the Greek samples. To sum up, the species of Mesopithecus were not leaf eaters like their present-day colobine relatives. Considering its wide paleogeographic range, its post-cranial anatomy, the paleoenvironmental data, and the present dietary reconstruction as a whole, the species of Mesopithecus can be depicted as semi-terrestrial monkeys opportunists in term of feeding preferences.