The Neogene cercopithecids (Mammalia, Primates) of Greece

George D. KOUFOS Aristotle University of Thessaloniki, Department of Geology, Laboratory of Geology and Palaeontology, GR-54124 Thessaloniki (Greece) [email protected]

Koufos G. D. 2009. — The Neogene cercopithecids (Mammalia, Primates) of Greece. Geo- diversitas 31 (4) : 817-850.

ABSTRACT Th e presence of the cercopithecids in the Neogene of Greece is known since the beginning of the 19th century. Th e excavations of the last 20 years increase their number in Greece. Th e main taxon of the cercopithecids isMesopithecus Wagner, 1839 found originally in the middle Turolian (MN 12) locality of Pikermi, near Athens. Th e 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. Th e 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 diff erences from the type species. In the middle Turolian (MN 12) localities of Vathylakkos (Axios Valley) and Perivolaki (Th essaly) a large to medium-sized Mesopithecus form with “delsoni” and “pentelicus” characters was found and is referred to as M. delsoni/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. Th e rest of the material found in the late KEY WORDS Turolian localities of Dytiko has some diff erences from the typical M. pentelicus. Mammalia, Moreover, Mesopithecus was traced in several late Miocene localities, indicating Primates, Cercopithecidae, its wide distribution in Greece. Two other cercopithecids were also found in the Mesopithecus, Pliocene of Greece. Dolichopithecus ruscinensis Depéret, 1889 was recognized in Dolichopithecus, Paradolichopithecus, the locality of Megalon Emvolon and in Ptolemais Basin (Macedonia, Greece); Neogene, both are dated to late Ruscinian (MN 15). Th e second Pliocene cercopithecid is Greece, Paradolichopithecus arvernensis (Depéret, 1929), found in the locality of Vatera systematic, biostratigraphy, (Lesvos Island) and dated to late Pliocene. Th e stratigraphic distribution and palaeoecology. the palaeoenvironment of these cercopithecids are also discussed.

GEODIVERSITAS • 2009 • 31 (4) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 817 Koufos G. D.

RÉSUMÉ Les cercopithécidés du Néogène (Mammalia, Primates) de Grèce. La présence des cercopithécoïdes dans le Néogène de Grèce est reconnue depuis le début du xixe siècle. Les fouilles des 20 dernières années ont permis d’enrichir considérablement l’échantillon et de nouveaux taxons ont pu être décrits. Le taxon le plus répandu est Mesopithecus, initialement mis au jour dans la localité Pikermi du Turolien moyen (MN 12), près d’Athènes. L’échantillon de Pikermi est riche et est reconnu comme M. pentelicus Wagner, 1839. Hormis cette espèce particulièrement bien décrite, deux autres taxons ont été décrits. La nouvelle espèce M. delsoni Bonis, Bouvrain, Geraads & Koufos, 1990, est une forme de grande taille trouvée dans la localité Ravin des Zouaves-5, dans des dépôts du Turolien inférieur (MN 11) de la vallée de l’Axios (Macédoine, Grèce). Les localités Vathy- lakkos (vallée de l’Axios) et Perivolaki (Th essalie) du Turolien moyen (MN 12) ont livré une forme de taille moyenne qui partage des caractères avec M. pentelicus et M. delsoni et est ici nommée M. delsoni/pentelicus. Un fragment de mandibule mis au jour dans le Turolien supérieur (MN 13) de Dytiko-2 (vallée de l’Axios) attribué à l’espèce de petite taille M. cf. monspessulanus accrédite une première apparition antérieure au Pliocène pour ce taxon. Le reste du matériel de Dytiko MOTS CLÉS présente quelques diff érences avec M. pentelicus. Le genre Mesopithecus est signalé Mammalia, dans de nombreuses localités du Miocène supérieur confi rmant sa vaste répartition Primates, Cercopithecidae, en Grèce. Deux autres cercopithécoïdes ont été également décrits dans le Pliocène Mesopithecus, de Grèce. Dolichopithecus ruscinensis Depéret, 1889 est reconnu dans la localité Dolichopithecus, Paradolichopithecus, de Megalon Emvolon et dans le bassin de Ptolemais (Macédoine, Grèce) ; deux Néogène, localités datées du Ruscinien récent (MN 15). Le second cercopithécoïde plio- Grèce, cène est Paradolichopithecus arvernensis Depéret, 1929 trouvé dans la localité de systématique, biostratigraphie, Vatera (Pliocène supérieur de Lesbos, Grèce). La distribution stratigraphique et le paléoécologie. paléoenvironnement de ces cercopithécoïdes sont également discutés.

INTRODUCTION found there up to now. Th e genus is also recognized in the neighbouring area, known from the late Th e cercopithecids (Mammalia, Primates) are quite Miocene of Italy, Hungary, FYROM, Bulgaria, common in the Neogene of Greece; the most com- and Ukraine, as well as from the Pliocene of Italy, mon one is Mesopithecus Wagner, 1839, represent- Bulgaria and Romania (Schlosser 1921; Gentilli ing the fi rst evidence of their presence in Greece. et al. 1998; Rook 1999; Kordos 2000; Koufos et It was originally discovered in the late Miocene al. 2003 and references therein; Radulescu et al. locality of Pikermi (Fig. 1) in 1835; during the 2003; Delson et al. 2005; Pradella & Rook 2007; years after several excavators found Mesopithecus NOW 2008) in Pikermi and today its sample is the richest one During the last 20 years new excavations have (Wagner 1839; Roth & Wagner 1854; Gaudry been carried out in various late Miocene Greek 1862-1867; Weithofer 1888; Woodward 1901). localities, providing several remains of Mesop- Later, the genus was recognized in Axios Valley, ithecus. Th e genus was discovered in Axios Valley, Macedonia, Greece (Arambourg & Piveteau 1929). Serres Basin, Th essaly and Chalkidiki (Bonis et al. In the meantime the late Miocene fossiliferous 1990, 1997; Kullmer & Doukas 1995; Koufos et sites of Samos had been discovered (Forsyth Ma- al. 2004; Koufos 2006a; Tsoukala & Bartziokas jor 1888, 1894) and no cercopithecids have been 2008). Th e new ndingsfi enriched our knowledge

818 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

N

7 Serres

11 5 Ptolemais 3 Thessaloniki 4 10 8 9

GREECE Volos 6 12 TURKEY

1 Athens 2

Mediterranean sea

100 km

FIG. 1. — Map indicating the cercopithecid bearing mammal localities of Greece: 1, Pikermi (PIK); 2, Chomateres (CHO); 3, Ravin des Zouaves-5 (RZO); 4, Vathylakkos-2, 3 (VTK, VAT) and Ravin-X (R-X); 5, Dytiko-1, 2, 3 (DTK, DIT, DKO); 6, Perivolaki (PER); 7, Maramena (MAR); 8, Kryopigi (KRY); 9, Nikiti-2 (NIK); 10, Megalon Emvolon (MEV); 11, Ptolemais basin; 12, Vatera (VTR). The map was taken from http://www.shaded-relief.com/. about the systematic and biostratigraphy of this in the Pliocene deposits of Megalon Emvolon and cercopithecid. Moreover, the analysis of the Meso- Ptolemais Basin, Macedonia, Greece (Koufos et al. pithecus associated faunas using new methods al- 1991a; Doukas & de Bruijn 2002). Th e second lowed to determine its palaeoecology. genus is Paradolichopithecus Necrasov, Samson & During the 1990’s two other cercopithecids Radulesco, 1961, discovered in the Pliocene local- were discovered in the Neogene of Greece. Th e ity of Vatera, Lesvos Island (de Vos et al. 2002; genus Dolichopithecus Depéret, 1889 was found Van der Geer & Sondaar 2002).

GEODIVERSITAS • 2009 • 31 (4) 819 Koufos G. D.

Th e present article is an eff ort to summarize R-X Ravin-X, Axios Valley, Macedonia, Greece; all available data about the Greek cercopithecids, RZ1 Ravin des Zouaves-1, Axios Valley, Macedo- nia, Greece; giving information for their localities, stratigraphy, RZO Ravin des Zouaves-5, Axios Valley, Macedo- morphology, age and systematics. Some biostrati- nia, Greece; graphical and palaeoecological remarks for the VAT Vathylakkos-3, Axios Valley, Macedonia, Neogene cercopithecids are also given. Greece; Th is article is written in honour of Prof. Louis de VLO Vathylakkos-1, Axios Valley, Macedonia, Greece; Bonis, University of Poitiers, and really it was a great VTK Vathylakkos-2, Axios Valley, Macedonia, pleasure for me to do it, as Louis is a good friend for a Greece; long time. I met Louis in 1976 when he was excavat- VTR Vatera, Lesvos Island, Greece; ing with Prof. J. K. Melentis in Axios Valley. After that XIR Xirochori-1, Axios Valley, Macedonia, time we have been working together in the area; we Greece. also worked together in Turkey and Chad. All these 35 years we had a nice collaboration and we worked in LATE MIOCENE LOCALITIES a friendly and scientifi cally productive way, trying to solve all small problems by discussion and comprehen- NIKITI sion. Th e writing of this article is a minimum off er to History the long time collaboration, the scientifi c discussions Th e localities of Nikiti were discovered at the be- and the hard work I shared with Louis. ginning of the 1990’s, when the villagers found some fossils during excavations for a new road. ABBREVIATIONS Th ey gave the information to the author who went AMPG Athens Museum of Palaeontology & Geo- logy, University of Athens; there and collected the fossils. Th is fi rst locality was BSPM Bayerische Staatssammlung für Paläontologie named Nikiti-1 (NKT) and it is well known by the und Historische Geologie, München; presence of the hominoid primate Ouranopithecus CHO Chomateres, Attica, Greece; macedoniensis Bonis & Melentis, 1977 (Koufos et DIT Dytiko-2, Axios Valley, Macedonia, Greece; DKO Dytiko-3, Axios Valley, Macedonia, Greece; al. 1991b). Th e prospection of the area gave several DTK Dytiko-1, Axios Valley, Macedonia, Greece; fossiliferous sites; most of them are poor in fossils Fm. Formation; but one named Nikiti-2 (NIK), provided a rich KRY Kryopigi, Chalkidiki, Macedonia, Greece; fauna. Among the collected material from NIK LGPUT Laboratory of Geology and Palaeontology, there are some remains of Mesopithecus, found in University of Th essaloniki; MAR Maramena, Serres Basin, Macedonia, the summer of 2008. Greece; MEV Megalon Emvolon-1, Macedonia, Greece; Stratigraphy and localities MNHN Museum national d’Histoire naturelle, Paris; Th e localities of Nikiti are situated in the Chalkidiki NHML Natural History Museum, London; Peninsula about 120 km east of Th essaloniki city NHMW Naturhistorisches Museum, Vienna; NIK Nikiti-2, Chalkidiki, Macedonia, Greece; (Fig. 1). Th e Neogene deposits of the Nikiti area NKT Nikiti-1, Chalkidiki, Macedonia, Greece; overlie unconformably the granitic basement NOW Neogene Old World database; and they are divided in two formations (Syrides PER Perivolaki, Th essaly, Greece; 1990; Koufos et al. 1991b; Kostopoulos & Kou- PIK Pikermi, Attica, Greece; PNT Pentalophos-1, Axios Valley, Macedonia, fos 1999). Greece; Nikiti Fm. It outcrops around the village of Nikiti PTL Ptolemais, Western Macedonia, Greece; and consists of clastic sediments, mainly sands, PXM Prochoma-1, Axios Valley, Macedonia, gravels with intercalations of pebbles; in the upper Greece; part of the formation there are intercalations or RPl Ravin de la Pluie, Axios Valley, Macedonia, Greece; lenses of red-brown sands, sandstones and sandy R-G Ravin-G or Ravin du Vatilük, Axios Valley, clays. Th e fossils are located in one of these lenses Macedonia, Greece; in the upper part of the Nikiti Fm.

820 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

– Nikolaos Fm. It is a fl uvio-lacustrine formation decades later a monograph on Pikermi Mesopithecus outcropping around the village of Agios Nikolaos. was published by Zapfe (1991), giving more signifi - It consists of marls, marly limestones, clays, sands cance to some undescribed material from AMPG and sandstones. Some macromammals were col- and to the postcranial skeleton. Some Mesopithecus lected from the reddish sands of the lower part remains were also found during the preparation of of the formation, as well as some micromammals several big-blocks from Pikermi, collected earlier from the clays. by various excavators and which remained for many years unprepared in the University of Athens Age (Th eodorou & Roussiakis pers. comm. 2008); this A quite rich mammalian fauna has been collected material is unpublished up to now. from NIK which allows its certain age determina- tion to the beginning of early Turolian (MN 11); Stratigraphy and locality the NIK fauna is slightly younger than the NKT All the old collections originate from the classical one, dated to latest Vallesian (MN 10) and slightly ravine of Pikermi, where the fi rst fossils were found older than that of RZO, dated to early Turolian and all the scientists excavated. Th e fi rst description (MN 11) at ~8.2 Ma (Koufos 2006b and refer- of the deposits and their stratigraphy are given by ences therein). Gaudry (1862-1867: pl. 74, fi g. 1). According to this description, the basement of the Neogene deposits Morphology is the Pentelikon marbles. Th e Neogene sediments Th e available material from NIK is very poor in- consist of alternated red clays and conglomerates- cluding two postcranials which can confi rm the anglomerates with intercalations of red clays. Th e presence of Mesopithecus in its fauna but cannot fossils are concentrated in lenses intercalated in the allow specifi c determination at the moment. Th us, red clays. Th e fossiliferous level, where Gaudry exca- it is referred to as Mesopithecus sp. vated, is situated near the bottom of the ravine and in the red clays. Th is level probably represents the PIKERMI main site where Gaudry excavated. Th e stratigraphy History of the Pikermi Neogene deposits is given by Mari- As it was referred above Mesopithecus was originally nos & Symeonidis (1974). According to them, the recognized in Pikermi; its holotype is a maxilla Neogene deposits overlie the marbles and schists of with M1-M3 sin described by Wagner (1839) as the basement and consist of the following lithology M. pentelicus Wagner, 1839 and fi gured one year from below to upwards. later by Wagner (1840) too. During the 1860’s a – Basal Beds. Th ey are uvialfl deposits consisted of signifi cant number of Mesopithecus remains from loose-cohesive conglomerates, sandstones, marls and Pikermi was collected by A. Gaudry, described clays with all intermediate lithological types; under the name M. pentelici. Gaudry’s collection is – Fossiliferous Beds. Th ey are of fluvial-fl uvio- housed in MNHN and includes several cranial and lacustrine origin, consisting of the typical reddish postcranial remains (Gaudry 1862-1867). Remains clays of Pikermi alternated with conglomerates, of the Pikermi Mesopithecus are also included in sandstones and marshy clays; the latter include Woodward’s collection housed in NHML, Wag- micromammals and lacustrine gastropods. Th e red ner’s collection in BSPM and in the University of clays include the lenses with the mammal fossils; Athens (AMPG collection). Besides these main – Calcareous Beds. Th ey overlie all the above men- collections there are several remains of Mesopithecus tioned beds in the whole area. Th ey consist of white- from Pikermi in various institutes (Stuttgart, Berlin, yellowish to brownish limestones and marls with Florence, Turin, Vienna). Several publications and conglomerates of lacustrine origin as indicated by the reports on this material have been published but remains of Melanopsis Férussac, 1823 and Planorbis the fi rst complete study of the Pikermi sample was (Müller, 1774). All the Neogene deposits are covered given by Delson (1973) in his doctorat thesis. Two by a series of Quaternary conglomerates.

GEODIVERSITAS • 2009 • 31 (4) 821 Koufos G. D.

Although the Pikermi collections are reported is rounded with sagittal sutures originating from the as homogeneous, this cannot be proved, as there orbits and connected in the parietals forming a sagit- are fossiliferous lenses across the ravine in diff erent tal crest (Fig. 2B, C) (the sagittal crest is stronger in horizons (Abel 1927: fi g. 136), while the faunal the males). Th e zygomatic arches are quite strong data would suggest diff erent stratigraphic levels starting above the contact between the M1 and M2. (Th eodorou & Nicolaides 1988). Th e basicranium is fl attened with large and rounded foramen magnum and large occipital condyles. Th e Age palate is elliptical, deep and its posterior end is situ- Th e lack of stratigraphic data and the doubt about ated at the posterior border of the M3 (Fig. 2E, F). the homogeneity of the Pikermi material gave rise Th e mandible has high ascending ramus vertical to to long discussions about its age which, although the mandibular corpus, the inferior border of which there is a general agreement, still continue. Th e is straight. Th e mandibular angle is rounded and available data of Pikermi suggest an age at the end projects strongly in the gonial area. Th e mandibular of middle Turolian (MN 12) at about 7.0 Ma, corpus is shallow and thick; its thickness and depth which is considered as the most possible (Koufos depend upon the sex, being higher in the males 2006b and references therein). Besides all these age (Fig. 3). Th e anterior face of the external symphysis determinations I strongly believe that it is necessary is rounded without symphyseal constriction, while to have a new series of excavations and detailed the internal symphysis has small planum alveolare stratigraphic studies in the classical Pikermi ravine strongly inclined backwards and a small or absent to get certain data which will allow us to determine fossa genioglossa. Th e dentition has the characters of correctly its fauna and age. the colobines with bunodont teeth and four cusps (-ids) in the molars except the m3 which has a clear Morphology hypoconulid (Figs 2; 3). Th e incisors are flattened As it was referred above, the material of Mesopithecus buccolingually, the lower being narrower than the from Pikermi is quite rich, the richest known for the upper ones (the I2 is more triangular-shaped). In the genus. However, its dispersion in several museums upper ones the I1 is larger than I2. In their lingual and institutes makes its study diffi cult. Fortunately, surface there are two grooves running across them. Th e I had the opportunity to study most of the Pikermi canines have a strong sexual dimorphism being re- Mesopithecus sample during the last fi ve years vis- markably larger in the males. Th e upper canine has iting various museums and institutes housing it. triangular transverse section with a relatively deep Th e main characters of the Pikermi Mesopithecus groove running across its mesiolingual surface and are given below. a small distal cingular projection. Th e lower canine It is a colobine monkey having medium size (be- is elliptical, curved backwards in the males, with a tween M. delsoni Bonis, Bouvrain, Geraads & Koufos, large and strong distal angular projection and two 1990 and M. monspessulanus Gervais, 1859). Th e crests running across its mesial and distolingual skull is characterized by strong prognathism giving surface. Th e upper premolars are bicuspid with the to it a primitive feature (Fig. 2A). Th e nasals are nar- buccal cusp larger than the lingual one, which is row and triangular-shaped with its posterior border small and disappears early by the attrition. Th e p3 situated at the middle of the orbits; their anterior is asymmetric, situated laterally to the tooth row axis border form two small processes in the middle of and having one large cuspid extended mesially; it the posterior border of the nasal cavity. Th e nasal has a relatively small honing facet. Th e p4 is more cavity is oval-shaped, large and their posterior bor- symmetrical than the p3 bearing a large buccal and der is situated above the short diastema between a smaller lingual cuspid, as well as, a mesial and a the canine and P3. Th e orbits are large, rounded, larger distal fovea. Th e molars have four cusps (-ids) with large interorbital distance (Fig. 2B-D). Th e and the upper ones are more squarish than the lower supraorbital torus is strong or weak (depending on ones. Th e M3 has more developed distal fovea (talon) the sex), continuous and with glabella. Th e braincase and the m3 a relatively small hypoconulid.

822 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

B A

A-D

C D

E

F

E, F

FIG. 2. — Cranial and maxillary remains of Mesopithecus pentelicus Wagner, 1839 from Pikermi (PIK), Attica, Greece: A, male skull associated with the mandible, NHMW-PIK-A.4714; B, female skull, AMPG-PIK-2; C, male skull, MNHN-PIK-14; D, male facial region, NHML-PIK-M.8947; E, male maxilla, NHML-PIK-M.8946; F, female maxilla, NHML-PIK-M.8948. Scale bars: 1 cm.

GEODIVERSITAS • 2009 • 31 (4) 823 Koufos G. D.

AB

C

DE

F

G

I H

FIG. 3. — Mandibular remains of Mesopithecus pentelicus Wagner, 1839 from Pikermi (PIK), Attica, Greece: A-C, male mandible, AM- PG-PIK-1b; D-F, female mandible, MNHN-PIK-03; G-I, male mandible, NHMW-PIK-1998z-01; A, D, G, occlusal views; B, E, H, right ramus; C, F, I, external symphysis. Scale bar: 1 cm.

824 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

AXIOS VALLEY gravels. According to Mercier (1968) the thickness History of the formation exceeds 1000 m. In the upper Th e late Miocene mammal localities of Axios Valley horizons of the formation four fossiliferous locali- (Macedonia, Greece) are situated across the river ties have been recognized: Pentalophos-1 (PNT), Axios, 25-50 km west of Th essaloniki city (Fig. 1). Xirochori-1 (XIR), Ravin de la Pluie (RPl) and Th e fi rst reference about their presence is that of Ravin des Zouaves-1 (RZ1); Bourcart (1919) who reported some mammal fos- – Vathylakkos Fm. It overlies normally Nea Mes- sils found near the village of Vathylakkos. During simvria Fm., outcropping in the eastern bank of the First World War (1915-1916) the geologist C. Axios River around the villages of Vathylakkos, Arambourg, who arrived in the area as an offi cer Prochoma and Agionerion. It consists of white-grey of the French Army, found some fossils around sands and gravels, alternated with yellowish sandy the village of Vathylakkos. Th en, he investigated clays. Several fossiliferous sites were found in this the area and he found several fossiliferous sites; formation; the majority of Arambourg’s collection thus he made a collection which is now stored in comes from this formation. Th e main fossiliferous MNHN (Arambourg & Piveteau 1929). In 1972 sites are Ravin des Zouaves-5 (RZO), Prochoma-1 the Laboratory of Geology and Palaeontology of (PXM), Vathylakkos-1, 2, 3 (VLO, VTK, VAT), the University of Th essaloniki (Prof. J. K. Melentis Ravin du Vatilük or Ravin-G (R-G) and Ravin-X and from 1976 the author) and the Laboratoire de (R-X). Th e locality R-G corresponds to the new Paléontologie des Vertébrés et Paléontologie humaine one, named VAT; of the University Paris 6 and after 1980 of the Uni- – Dytiko Fm. It outcrops in the western bank of versity of Poitiers (Prof. L. de Bonis) started a series Axios River around the village of Dytiko. It mainly of excavations in Axios Valley which still continue. consists of yellow-yellowish sands, gravels and sandy During these fi eld campaigns new fossiliferous sites clays with intercalations of sandstones, while in the were discovered and a great amount of fossils has top of the formation there are yellowish fresh water been unearthed, while more than 100 articles have marly limestones. Th ree fossiliferous sites Dytiko-1, been published. Th e old Arambourg’s collection 2, 3 (DTK, DIT, DKO) have been discovered in includes some remains of Mesopithecus which come this formation. from the localities R-X and R-G (Arambourg & Th e Mesopithecus bearing localities of Axios Valley Piveteau 1929). It is worth mentioning that Ara- are RZO, VTK, VAT and R-X of Vathylakkos Fm., mbourg’s collection is mixed including fossils from as well as, all the localities of Dytiko Fm. diff erent localities of diff erent age; some fossils have locality indications, some others do not and, thus, Age cannot provide certain data for biochronology and Th e available fauna of the Axios Valley localities comparisons. During the new excavations several allow their certain biochronological dating. Th e remains of Mesopithecus were found in the various locality RZO is dated to early Turolian, MN 11, the localities of Axios Valley, stored in LGPUT (Bonis localities VTK, VAT and R-X to the lower middle et al. 1990, 1997; Koufos et al. 2004). Turolian, MN 12, and the Dytiko localities to late Turolian, MN 13 (Koufos 2006b and references Stratigraphy and localities therein). Th e magnetostratigraphic record suggests Th e late Miocene deposits of Axios Valley are di- an estimated age of ~8.2 Ma for RZO and ~7.3 Ma vided in three diff erent formations (Koufos 1980, for VTK and VAT localities (Sen et al. 2000; Kou- 1990). fos, unpubl. data). – Nea Mesimvria Fm. It outcrops in the southern part of the eastern bank of Axios River. Th e forma- Morphology tion consists of gravels, sands, conglomerates and Locality RZO. Th e available material from RZO red clays; their cohesion varies greatly from very includes only mandibular remains (Fig. 4), described hard and compact conglomerates to loose sands and as a new species M. delsoni (Bonis et al. 1990).

GEODIVERSITAS • 2009 • 31 (4) 825 Koufos G. D.

D A

B

E

C F

G

H

FIG. 4. — Mandibular remains of Mesopithecus delsoni Bonis, Bouvrain, Geraads & Koufos, 1990 from Ravin des Zouaves-5 (RZO), Axios Valley, Macedonia, Greece: A-C, male mandible, LGPUT-RZO-159; D-F, male mandible, LGPUT-RZO-160; G, H, mandibular fragment of a female individual, LGPUT-RZO-327; A, left ramus; B, E, H, occlusal views; C, F, external symphysis; D, right ramus; G, lateral view. Scale bar: 1 cm.

826 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

Th e mandibular corpus is very deep distinguishing M. pentelicus from Pikermi. Th e palate is elliptical the RZO mandibles from those of M. pentelicus. and deep with wide and relatively shallow choanae, Contrary to M. pentelicus, the anterior symphysis the anterior border of which is situated at the level is fl attened and has lateral symphyseal constriction, of the M3. Th e mandible VTK-62 has flattened while the internal symphysis has slightly inclined anterior symphysis with symphyseal constriction. backwards planum alveolare and large fossa genioglossa. Th e internal symphysis has slightly inclined back- Th e inferior transverse torus is thick and prominent. wards planum alveolare and large fossa genioglossa Th e teeth are large, the p3 has large honing facet (Fig. 6E, G). Th e mandibular corpus is not very versus a small one in M. pentelicus and the m3 has deep, having similar depth to that of M. pentelicus a large hypoconulid versus a small one in M. pente- from Pikermi. Th e upper teeth of the Vathylakkos licus. In the distal face of the hypoconulid there is sample have the general morphology of Mesopithecus a deep groove giving to it a bicuspid aspect. Th e and similar dental size to M. pentelicus. Th e upper Principal Component Analysis (PCA) of the RZO premolars have relatively larger lingual cusp (pro- and PIK samples indicates that the RZO sample tocone) than that of M. pentelicus. Th e p3 has large is separated from M. pentelicus having relatively honing facet and the m3 large hypoconulid with larger mandibular and dental dimensions, as well a groove in its distal face, like in M. delsoni. Th e as smaller lower canine (Koufos in press). All the PCA of the upper and lower dental dimensions of above mentioned morphological and metrical dif- the VTK sample with those of M. pentelicus and ferences separate the RZO sample from the typical M. delsoni indicates its separation from both taxa M. pentelicus and allowed Bonis et al. (1990) to having intermediate characters (Koufos in press). erect the new species M. delsoni. Th e VTK Mesopithecus has characters between M. delsoni and M. pentelicus and thus is referred to Locality VTK. Th e Mesopithecus sample from VTK as M. delsoni/pentelicus. In the earlier articles it is is a recent collection and includes some cranial and referred as M. aff . M. pentelicus (Bonis et al. 1997; mandibular remains (Figs 5; 6), housed in LGPUT Koufos et al. 2004) but it is better to refer this as and described earlier (Bonis et al. 1997; Koufos et M. delsoni/pentelicus because of its similarities to al. 2004). Some dental remains stored in MNHN both species; it can be considered as an evolution- and referred to as Braillon’s collection seem to origi- ary stage between the two species. nate from VTK too (Bonis et al. 1997). Th e VTK Mesopithecus is larger in size than the typical M. pen- Localities R-G and R-X. Th e material from these telicus from Pikermi and closer to M. delsoni from two localities includes that of Arambourg’s col- RZO. Th e orbits are more squarish with rounded lection housed in MNHN (Fig. 4). According to corners and they have supraorbital torus which is Aram bourg & Piveteau (1929: 74), Mesopithecus was separated in two parts by a depression in the area found in Ravin du Vatilük or Ravin-G (R-G) and in of the glabella. Th e supraorbital torus is stronger Ravin-X (R-X). However, in the faunal lists given for in the male skull than in the female one (Fig. 5). each locality (Arambourg & Piveteau 1929: 13, 14) A sulcus is present behind the supraorbital torus. Mesopithecus is referred only from R-X. Th e locality Th e interorbital distance is large being 10.8 mm in R-G corresponds to the new one referred as VAT; both VTK-61 and ~10 mm in VTK-56. Th e nasal cavity VAT and R-X are located in Vathylakkos Fm. Th e is deep elliptical-shaped with its posterior border type of fossilization and the colour of the sediments situated above the contact between the P4 and M1. indicate that most specimens of the Arambourg’s Th e lacrymal fossa is well distinguished and it is collection originated from VAT but one specimen extended into the maxilla. Th e zygomatic arches are (MNHN-SLQ-940+941) possibly comes from R-X. strong and their base is situated above the fi rst lobe Th e specimens from VAT have similar morphology of the M2. Th e cranial roof is damaged in VTK-61 to those from VTK and must belong to the same but in the skull VTK-56 the sagittal sutures and morphotype, M. delsoni/pentelicus. Th e specimen crest are weakly developed as in the female skulls of from R-X is a small part of the anterior mandible

GEODIVERSITAS • 2009 • 31 (4) 827 Koufos G. D.

A

B

C

D

FIG. 5. — Cranial remains of Mesopithecus delsoni/pentelicus from Vathylakkos-2 (VTK), Axios Valley, Macedonia, Greece: A, C, male skull, LGPUT-VTK-61; B, D, female skull, LGPUT-VTK-56; A, B, facial views; C, D, occlusal views. Scale bar: 1 cm.

828 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

A B

C D

E G

F

FIG. 6. — Cranial and mandibular remains of Mesopithecus delsoni/pentelicus from Vathylakkos-2 (VTK), Axios Valley, Macedonia, Greece: A, B, young male skull, LGPUT-VTK-78, facial (A) and occlusal (B) views; C, D, young male mandible, LGPUT-VTK-78, oc- clusal (C) and right ramus (D) views; E-G, female mandible, LGPUT-VTK-62, occlusal view (E), right ramus (F) and external symphysis (G). Scale bar: 1 cm.

GEODIVERSITAS • 2009 • 31 (4) 829 Koufos G. D.

A B

D C

E

F

FIG. 7. — Cranial and mandibular remains of Mesopithecus Wagner, 1839 from the localities of Dytiko (DTK, DIT, DKO), Axios Valley, Macedonia, Greece: A, B, M. aff. pentelicus, male skull, LGPUT-DKO-38, occlusal (A) and facial (B) views; C, D, same, partial mandi- ble, LGPUT-DKO-38, right ramus (C) and occlusal views (D); E, F, M. cf. monspessulanus, partial mandible, LGPUT-DIT-22, occlusal view (E) and left ramus (F).

830 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

26

25

24

Length m1-m3 23

22

21 34 36 38 40 42 44 46 48 50 Length c-m3

FIG. 8. — Scatter diagram comparing the length c-m3 versus molar length of M. pentelicus Wagner, 1839, M. monspessulanus Ger- vais, 1859 and M. cf. monspessulanus from DIT. ◆, M. pentelicus, PIK, male; ◇, same, female; xI , M. cf. monspessulanus, DIT-22; ■, M. monspessulanus, Villafranca d’Asti, male (Delson 1973); ●, M. monspessulanus, Montpellier, male (Delson 1973); ○, same, female (Delson 1973).

which has stronger similarities (fl attened external TABLE 1. — Comparative measurements of Mesopithecus Wagner, symphysis with symphyseal constriction, slightly 1839 from Pikermi and Dytiko. inclined planum alveolare, large fossa genioglossa, deep mandibular corpus, strong honing facet in the DKO-231 M. pentelicus Wagner, 1839 p3) to the RZO Mesopithecus and could belong to (Pikermi) M. delsoni; thus it is referred to as Mesopithecus cf. male male female M. delsoni because of the limited material. Orbit length 19.5 20.0-25.5 Orbit breadth 24.0 24.1-30 Dytiko localities. Th e Dytiko material of Meso- Li1 3.2 (3.8) 4.39 (5.4) (3.8) 4.14 (4.4) Bi1 2.7 (2.7) 3.21 (3.6) (2.0) 2.84 (3.4) pithecus is quite rich including both cranial and Li2 4.0 (4.0) 4.60 (5.1) (4.0) 4.40 (4.9) postcranial remains. Th e material from DKO in- Bi2 2.9 (2.7) 3.38 (3.9) (2.6) 3.02 (3.5) cludes a partial male skull associated with the man- dible, DKO-38 (Fig. 7). Although it has slightly smaller size than M. pentelicus (Table 1), a more the facet for the ulna is small in comparison to careful comparison indicates some morphological that of M. pentelicus. Based to these morphological diff erences. Th e orbits are relatively smaller with diff erences of the DKO material from the Pikermi stronger supraorbital torus and the lower incisors M. pentelicus, it is referred to as Mesopithecus aff . are smaller than those of the Pikermi M. pentelicus. M. pentelicus. Th e material from DIT includes a Th e DKO postcranials have also some diff erences strongly deformed skull which is similar both to from those of the typical M. pentelicus (Bonis et al. Pikermi and DKO material and it is also referred 1990). Th e epicondylus medialis of the humerus to as Mesopithecus aff . M. pentelicus (Bonis et al. is less refl ected backwards in comparison to that 1990). Th e material from DTK (Fig. 5) is similar of the Pikermi M. pentelicus. Th e radius has more to Pikermi and it is referred to as Mesopi thecus cf. oval section at the shaft than M. pentelicus, while M. pentelicus (Bonis et al. 1990).

GEODIVERSITAS • 2009 • 31 (4) 831 Koufos G. D.

AB

CD

E

FG

HI

FIG. 9. — Mandibular remains of Mesopithecus Wagner, 1839 from the localities of Chomateres (CHO), Attica, Greece and Perivolaki (PER), Thessaly, Greece: A-D, M. pentelicus Wagner, 1839, male mandible, NHMW-CHO-1623/1a, right ramus (A), left ramus (B), external symphysis (C) and occlusal view (D); E-G, M. delsoni/pentelicus, female mandible, LGPUT-PER-200, occlusal view (E), left ramus (F) and external symphysis (G); H, I, same, left male mandibular ramus, LGPUT-PER-1284, external (H) and occlusal (I) views. Scale bar: 1 cm.

832 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

Among the material of DIT there is a male man- Age dible (DIT-22) which is smaller than the male Based to the preliminary published articles about mandibles of the Pikermi Mesopithecus; its dental Chomateres a faunal list is given by Koufos (2006b); dimensions are remarkably smaller than those of the fauna is similar to that of Pikermi, suggesting the male M. pentelicus of Pikermi ranging between a middle Turolian age, MN 12. In the updating the latter and the male M. monspessulanus (Fig. 8). of the MN biozones (Mein 1990) Chomateres is Th e multivariate comparison of its mandibular and considered as slightly older than Pikermi. A simi- dental dimensions with M. pentelicus clearly confi rms lar age is also proposed by de Bruijn et al. (1992), its smaller size (Koufos in press). Th e small size of Bonis & Koufos (1999) and Koufos (2006b). How- DIT-22 suggests similarities to M. monspessulanus ever, only the study and comparison of the fauna but the limited material cannot allow a certain de- with that of Pikermi and other Greek localities will termination and thus it is referred to as Mesopithecus provide certain data about the precise age of the cf. M. monspessulanus (Bonis et al. 1990). Chomateres fauna.

CHOMATERES Morphology History Th e available Chomateres sample of Mesopithecus In 1971 a new fossiliferous site was discovered includes only two mandibular fragments (Fig. 9A-D). in Attica, named Chomateres or Kisdari (Fig. 1). Th e mandible NHMW-CHO-1613/1a is similar to During the excavating activities in a clay-pit of that of Pikermi but it has smaller tooth rows, possibly the area some mammal fossils were found and due to the very worn dentition; its mandibular dimen- the University of Athens made the fi rst collection sions put it in the group of M. pentelicus (Koufos in (Marinos & Symeonidis 1972). Th e University of press). Th e other mandible (NHMW-CHO-1613- Athens in collaboration with NHMW continued /1b) has mandibular and dental dimensions similar the excavations in Chomateres during 1972-1973, to M. pentelicus but it has some minor morphological while H. de Bruijn washed material and found diff erences. Th e anterior face of the symphysis is some micromammals in the marshy clays of the less convex (more fl attened than in M. pentelicus), area (Marinos & Symeonidis 1974). Th e collected there is a weak symphyseal constriction, a slightly fauna of the large mammals is similar to that of larger fossa genioglossa and a larger honing facet in Pikermi and includes some remains of Mesopithecus the p3. However, both CHO mandibles cannot be (Fig. 7A, B). Th e Chomateres mammal collection separated from M. pentelicus at the moment and they is unpublished except for some general information are referred to as M. pentelicus (Koufos in press). Th e about the locality and fauna and some few taxa, presence of some “delsoni” characters in the CHO like Mesopithecus and Metailurus Zdansky, 1924 sample should be considered as an indication for (Marinos & Symeonidis 1972, 1974; Symeonidis an age slightly older than Pikermi. et al. 1973; Symeonidis 1978; Zapfe 1991). Th e CHO collection is housed in AMPG except for PERIVOLAKI the Mesopithecus sample which is in NHMW, at the History moment. Th e initial description of the Chomateres Th e locality of Perivolaki was discovered in 1996 Mesopithecus is given by Zapfe (1991) who described by some geologists of IGME (Institute of Geologi- a mandibular fragment as a new subspecies, named cal and Mineralogical Exploration, Athens). Th e M. pentelicus microdon. information about the presence of the fossils was given to the author who went there and made a Stratigraphy and locality surface collection saving some fossils. Next year, Th e locality of Chomateres is situated about 2.5 km a team of palaeontologists from the LGPUT, led east of the classical Pikermi ravine. Th e stratigraphy by the author, started a series of excavations which of the Neogene deposits is similar to that given for continued until 2004. During these fi eld trips a great Pikermi (see above). amount of fossils have been unearthed. All Perivolaki

GEODIVERSITAS • 2009 • 31 (4) 833 Koufos G. D.

A B GH

G-J A-C

C IJ

E

D

D-F

F

K

L

FIG. 10. — Mandibular remains of Mesopithecus Wagner, 1839 from Maramena (MAR), Serres Basin Macedonia, Greece, Dolichopithecus Depéret, 1889 from Megalon Emvolon (MEV) and Ptolemais (PTL), Macedonia, Greece and Paradolichopithecus Necrasov, Samson & Radulesco, 1961 from Vatera (VTR), Lesvos Island, Greece: A-C, Mesopithecus sp., AMPG-MAR-040, right female mandibular fragment with i2-p4, labial (A), lingual (B) and occlusal (C) views; D-F, Dolichopithecus ruscinensis Depéret, 1889, female mandible, LGPUT- MEV-1, occlusal view (D), left labial ramus (E) and left lingual ramus (F); G, H, same, M2, AMPG-PTL-nn1, occlusal (G) and labial (H) views; I, J, same, M2, AMPG-PTL-nn2, occlusal (I) and labial (J) views (photos kindly provided by Prof. C. Doukas); K, L, P. arvernensis (Depéret, 1929), two mandibles AMPG-VTR-114F (K) and AMPG-VTR- (L) (photos kindly provided by Dr G. Lyras). Scale bars: 1 cm.

834 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

collection is stored in LGPUT and it was published large hypoconulid in the m3). All these characters in a complete volume of Palaeontographica (Koufos suggest similarities to the Vathylakkos Mesopithecus ed. 2006e). Th e Perivolaki collection includes some and thus it is determined to as Mesopithecus delsoni/ remains of Mesopithecus, which were described by pentelicus (Koufos 2006a, in press). Koufos (2006a). MARAMENA Stratigraphy and locality History Th e locality of Perivolaki is situated near Velestinon Th e locality of Maramena was discovered in an town about 40 km southwest of Volos city, Th essaly, artifi cial outcrop made by the uranium explora- Central Greece (Fig. 1). Th e Neogene stratigraphy tion activities of “Demokritos” (now IGME) in of the Perivolaki wider region is not known but our the 1970’s. A surface collection was made by some investigations in the neighbouring area allow the geologists of IGME and later by H. de Bruijn and C. distinction of four diff erent lithostratigraphic units Doukas. A preliminary list of the Maramena fauna (Koufos et al. 1999; Sylvestrou & Koufos 2006): is given by Karistineos (1984). Th en the University – Unit-A consists mainly of white-grey lacustrine of Athens in collaboration with the Universities of massive marls and marly limestones with a visible Utrecht and Mainz excavated and collected both thickness of 50-60 m; micro- and macro-mammals. Th e micromammals – Unit-B is characterized by interbedded, green- are rich but the macromammals are relatively few brown fi ne to coarse sediments consisting of brown- and include several remains of Mesopithecus, mainly ish-greenish clays alternating with sands, sandstones isolated teeth. Th e Maramena material is housed and marly-sandy limestones with a thickness of in AMPG and was described in a separate volume 60-80 m. Th e fossiliferous site is located in the of Münchner Geowissenschaftliche Abhandlungen upper part of the Unit-B; (Schmidt-Kittler 1995). – Unit-C consists of red-brown conglomerates alternating with yellowish clays and locally with Stratigraphy and locality reddish clays, having a thickness of ~100 m; Th e locality of Maramena is situated in Serres – Unit-D includes the recent deposits consisting Basin, about 10 km north of Serres town (Fig. 1). of terrestrial red clays alternating with clayey sands Th e Neogene deposits of the area can be divided and silts. in the following formations (Armour Brown et al. 1977): Age – Lefkon Fm. which overlies unconformably the Th e biochronological data suggests a middle Turo- basement and has a thickness of ~250 m. It consists lian (MN 12) age for Perivolaki, while the palaeo- of polymict conglomerates and coarse unsorted magnetic record indicates a correlation to Chron arkose with intercalations of fi ne grained sediments C3Br.2r, suggesting an age between 7.3 and 7.1 Ma (arkosic sandstones, siltstones, mudstones, marls and (Koufos et al. 2006). lignites). Th e locality MAR is situated in the upper part of the formation in the ligniferous beds; Morphology – Georgios Fm. which is ~100 m thick and consists Th e Perivolaki Mesopithecus sample consists of man- mainly of carbonate clastic sediments and sandy dibular remains only (Fig. 9E-I). Th e available female limestones interfi ngered with a breccia composed mandible PER-200 is well preserved and character- of granitic blocks. Th e thin sediments include a ized by intermediate size and morphology between marine and brackish fauna, well known in the M. pentelicus (similar dental dimensions and low southern part of the basin; mandibular corpus) and M. delsoni (fl attened anterior – Spilia Fm. It has a thickness of ~300 m and con- symphysis with symphyseal constriction, small and sists mainly of white well-sorted sandstones with roughly inclined backwards planum alveolare, large sporadic lenses of well-rounded pebbles. In the fossa genioglossa, large honing facet in the p3, and center of the area the sandstones interfi nger with

GEODIVERSITAS • 2009 • 31 (4) 835 Koufos G. D.

carbonaceous mudstones, siltstones and fi ne sand- PLIOCENE LOCALITIES stones. Th e Spilia Fm. includes several fossiliferous levels with small mammals. MEGALON EMVOLON History Age Th e locality of Megalon Emvolon was discovered by Th e rich micromammalian fauna of Maramena C. Arambourg at the beginning of the 20th century allowed its age determination to the Turolian/ (see Arambourg & Piveteau 1929, chapter 3); it is Ruscinian boundary, MN 13/14 (Schmidt-Kittler referred as “Falaise de Karabouroun” (Arambourg & et al. 1995). However, the whole fauna has more Piveteau 1929). Th e Arambourg’s collection includes Miocene feature and thus an age to the uppermost few specimens from Megalon Emvolon, housed in Turolian is quite possible (Koufos 2006b). MNHN. During the 1970’s several palaentologists visited the area and collected isolated specimens which Morphology are mainly housed in Athens and Utrecht. In 1989 Th e Mesopithecus sample of Maramena includes the author visited the area and found some small mainly isolated teeth, as well as some mandibular concentrations of fossils dispersed in the sediments fragments (Fig. 10A-C). Th e morphology of the of the area. Among them there were the two branches teeth is similar to Mesopithecus. Th e honing facet of of a cercopithecid mandible (Koufos et al. 1991a). the p3 and the hypoconulid of the m3 are small. In Our eff orts to get more fossils in these sites, where AMPG-MAR-040 the horizontal ramus is broken we found the initial material, were unsuccessful. Th e but its small height below p3 (~18.3 mm) indicates fossils are dispersed in the sediments and there is not similarities with M. pentelicus. Th e material has been any fossiliferous level or large fossiliferous lenses. determined to M. pentelicus (Kullmer & Doukas Th us, we go there from time to time and check if 1995) but after the discoveries in Axios Valley and some fossils have appeared by the erosion. Perivolaki (see above) needs a revision. However, the isolated teeth cannot allow a comparison with Stratigraphy and locality the new material as the distinctive characters are Th e locality of Megalo Emvolon is situated ~20 km mainly recognized in the mandible and skull. Th us, south of Th essaloniki city in a cap, named Megalon it is better to refer this as Mesopithecus sp., at the Emvolon, marking the entrance of Th essaloniki Gulf. moment. Some data about the lithology of the sediments are given by Arambourg & Piveteau (1929); they men- KRYOPIGI tioned reddish marls (~3 m thick) at the bottom Th e locality of Kryopigi is situated in Chalkidiki of the section, followed by grey marls with sands (Macedonia Greece) and it was found during the and gravels (~30 m) and gravels-pebbles at the top 1990’s (Fig. 1). Th e Mesopithecus sample includes a (~15 m). A more detailled lithostratigraphy of the skull associated with the mandible and a maxillary section is given by Koufos et al. (1991a: fi g. 1). fragment. Th e Kryopigi fauna is not studied and – Lower Sands. Th e lower part of the section con- thus the few available data cannot allow a certain sists of fl uvial deposits, mainly sands and gravels age determination except for late Miocene. Although with cross-bedding and with lenses or lenticular the Mesopithecus material is refered to M. pentelicus intercalations of sandstones. (Tsoukala & Bartziokas 2008), detailed descriptions – Red-brown Sands. Th ey overlie the Lower sands and comparisons with other Mesopithecus are neces- and consist of red to brown sands and gravels, red- sary to assign a specifi c level to this sample. Keep- sands with red-clay and intercalations of conglom- ing in mind all the above mentioned, the Kryopigi erates and calcitic concretions. sample must be referred as Mesopithecus sp. at the – Upper Sands. In the upper part of the section there moment, waiting for more precise morphological are fl uvial sands, with cross-bedding, intercalations study, comparisons and dating which will allow a of conglomerates and calcitic concretions. Th e up- certain attribution. per part of the Upper Sands is more clayey.

836 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

As referred above there is not any fossiliferous level PTOLEMAIS BASIN in Megalon Emvolon, but there are small concentra- History tions of fossils dispersed in the outcrop. Based to our Th e cercopithecid material found in the Ptolemais collection there are three diff erent fossiliferous levels. lignitic basin was discovered at the end of the 1990’s Th e Megalon Emvolon 1 (MEV), situated at the bot- by H. de Bruijn and C. Doukas working there for tom of the section (near the sea), the Megalon Em- micromammals. Two cercopithecid teeth were found volon 2 (MEM), situated at the bottom of the Upper in the ligniferous pit, named South Field and they Sands and the Megalon Emvolon 3 (MEL), situated were described by Doukas & de Bruijn (2002). in the clayey part of the Upper Sands. Stratigraphy and locality Age Ptolemais Basin is located in western Macedonia Based to the available faunal data of Megalon Em- (Greece) about 140 km west of Th essaloniki city volon a late Ruscinian (MN 15) age is proposed for (Fig. 1). It is part of a large tectonic depression, it (Koufos 2006b and references therein). fi lled by Neogene-Quaternary deposits including lignites. Th e Neogene deposits of Ptolemais Basin Morphology are divided in three main formations (Koufos & Th e sole known cercopithecid mandible (MEV-1) Pavlides 1988; Steenbrink 2001): belongs to an aged female individual (Fig. 10D-F). – Basal Fm. consisting mainly of gravels, pebbly Th e mandibular corpus is deep and thick with a single breccia and conglomerates. Th ere is no outcrop of mental foramen below the anterior root of the p3. this formation; Th e canine is small, indicating a female individual, – Vegora Fm. consisting of green sands, sand-silts, and characterized by a distal crest beginning from the grey marls and yellowish sandy silts. Th e formation is base to the middle of the canine’s height. Th e p3 has characterized by the presence of lignites (xylitic type). a large protoconid and talonid basin, as well as strong Th e Vegora Fm. includes a rich megafl ora which in- lingual and distal basal cingulum. Th e p4 is narrow dicates a late Miocene age (Kvaček et al. 2002); and characterized by the approximately equal height – Ptolemais Fm. outcroping in the whole Ptole- of the metaconid and protoconid but the latter cuspid mais area and including the main lignitic deposits is wider; this character is quite clear in D. ruscinensis of the area. Th e Ptolemais Fm. consists mainly of (Delson 1973). Th e m2 is larger than m1 and both sandy-silts, silts, sands and clays with lignitic in- have metaconid and entoconid of equal height, while tercalations. Th e formation can be distinguished in they are higher than the protoconid and hypoconid. a lower and upper ligniferous level, separated by Th e distal breadth of the m1 is greater than the mesial an intermediate without lignites part (Koukouzas one. Th e m3 is large with well-developed hypoconulid et al. 1979); situated between the midline of the tooth and the – the Neogene deposits overlied unconformably line connecting the buccal cuspids. In the lingual by Quaternary mainly lacustrine-fl uviolacustrine part of the talonid there is an elevation of the distal deposits. cingulum, indicating the beginning of a sixth cus- pid. Th e mesial breadth of the m3 is larger than the Age distal one and the distal breadth of the m2. Th e size Th e available faunal data of Ptolemais Fm. suggest a of the teeth fi ts quite well to the size of those of D. Ruscinian age (Koufos 2006b and references therein) ruscinensis (Koufos et al. 1991a). Keeping in mind for it; more precisely a late Ruscinian (MN 15) age all the above mentioned, the mandible MEV-1 is is referred for Notio Mb where Dolichopithecus was attributed to D. ruscinensis. Th is is the first evidence found (de Bruijn pers. comm. 2008). of the presence of Dolichopithecus in eastern Mediter- ranean. Recently a new species D. balcanicus has been Morphology described from Bulgaria and the MEV mandible was Th e two available M2 are quite worn and their included to it (Spassov & Geraads 2007). morphology is not clear (Figs 10G-J). However,

GEODIVERSITAS • 2009 • 31 (4) 837 Koufos G. D.

their characters indicate a colobine monkey, while to allow a certain determination of their geologi- their dimensions fall within the range of Dolicho- cal age (de Vos et al. 2002; Lyras & Van der Geer pithecus ruscinensis from Perpignan (Doukas & de 2007). Bruijn 2002). Th us, the authors identifi ed them to this species, despite the material paucity and Morphology limited morphology. Th e sample of cercopithecids from Vatera includes two mandibles, some isolated teeth, including VATERA an adult upper canine, and several postcranials History (Fig. 10K, L) (Van der Geer & Sondaar 2002). Th e fossiliferous site of Vatera is located in Les- According to Van der Geer (pers. comm. 2008) vos Island (Aegean Sea) and was discovered in the lower dentition of the Vatera Paradolichop- 1997 (Fig. 1). Th e fossils were found during the ithecus is typically papionin. Th e anterior teeth construction of a road at an olive yard. Subse- are large and the canine bears a clear sexual quently, the University of Athens and the Natural dimorphism. Th e incisors lack enamel on their History Museum of Leiden collected some fos- lingual surface. Th e p3 is elongated with a rela- sils and the year after they started a systematic tively large distal fovea and rather large honing excavation of the locality. In 2000 the excava- facet. Th e p4 is wide with a relatively larger labial tions were ended due to lack of further fossils. (metaconid) than lingual (protoconid) cuspid. During these fi eld campaigns several fossils had Th e trigonid of the m3 is long with low relief; been collected, among them some cercopithecids between the hypoconulid and the entoconid there remains (Van der Geer & Sondaar 2002; de Vos is a large sixth cuspid. Th e postcranials have been et al. 2002; Sondaar et al. 2006; Lyras & Van studied and they are morphologically and metri- der Greer 2007). cally similar to the type material of P. arvernensis (Depéret, 1929) from Senéze (France) and to that Stratigraphy and locality from Graunceanului, Romania (Van der Geer & Th e locality of Vatera is situated in the southern Sondaar 2002; Sondaar et al. 2006). According part of Lesvos Island (Fig. 1) about 45 km from to Sondaar et al. (2006) the postcranials of the Mytilini town. Th e Neogene deposits of the Vatera Vatera Paradolichopithecus are unique as far as the region consist of a series of lacustrine-brackish degree of terrestriality is considered, indicating deposits (lower part) which are overlied by a series similarities to Australopithecus afarensis Johan- of fl uvial deposits (upper part). Th e lacustrine- son & White, 1978. Th us, the authors proposed brackish deposits consist of marly limestones with that the locomotion of Paradolichopithecus might sandstone and tuffi te’s intercalations. Th e fl uvial be similar to that of Australopithecus Dart, 1925 deposits consist of alternating sandy clays, sandy which, according to them, retained basically a conglomerates, silts and breccia-conglomerates chimpanzee-like locomotion. Chimpanzees can (Drinia et al. 2002). Th e mammal fossils were walk bipedally but also climb to the trees (Sondaar found in the upper fl uvial horizons of the sequence et al. 2006). Th e eruption sequence of the lower and they are dispersed in small pockets. teeth of the Vatera Paradolichopithecus is similar to that of the recent papionins (Van der Geer & Age Dermitzakis in press). Th e Vatera Fm. includes a number of fossiliferous sites (F, DS, E, U, U, T, V) corresponding to diff er- ent fossil pockets. Th e richest fauna including the SYSTEMATICS cercopithecid is F, dated to middle Villafranchian s.l., or MN 17 (late Pliocene), at ~2.0 Ma. Th e fau- Order PRIMATES Linnaeus, 1758 nas from DS and E sites are dated to late Pliocene, Family CERCOPITHECIDAE Gray, 1821 while those of the U, H, T and V sites are too poor Subfamily COLOBINAE, Blyth, 1875

838 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

Genus Mesopithecus Wagner, 1839 LOCALITY. — ?Ravin-X (R-X), Axios Valley, Macedo- nia, Greece.

Mesopithecus pentelicus Wagner, 1839 AGE. — Early-middle Turolian, MN 11-12 (late Mio- cene). HOLOTYPE. — Maxillary fragment with M1-M3 de- scribed by Wagner (1839) and fi gured by Wagner (1840). CHARACTERISTICS. — Characters and size similar to Th e specimen is housed in BSPM numbered as BSPM M. delsoni. Th e sole known specimen does not allow ASII. 11. the certain attribution to this species.

LOCALITIES. — Classical Pikermi ravine (PIK), Attica, Greece (near Athens); Chomateres or Kisdari (CHO), Mesopithecus delsoni/pentelicus Attica, Greece (near Athens). MATERIAL EXAMINED. — Th e studied material includes AGE. — Middle Turolian, MN 12 (late Miocene); more precisely it is referred to the uppermost MN 12 with an that referred in Bonis et al. (1997), Koufos et al. (2004) age of ~7.0 Ma. Th e fauna of Chomateres is considered and Koufos (2006a). slightly older than the Pikermi one. LOCALITIES. — Vathylakkos-2, 3 (VTK, VAT), Axios Valley, Macedonia, Greece; Perivolaki (PER), Th essaly, DIAGNOSIS. — Medium-sized colobine monkey; short, Central Greece. upright face; sexual dimorphism in the skull, canines and postcranials; absent or very small sagittal crest in AGE. — Middle Turolian, MN 12 (late Miocene). More the males; enlarged mandibular angle; shallow man- precisely the Vathylakkos localities are dated to the lower dibular corpus with constant height between p4 and middle Turolian with an estimated age of ~7.3 Ma, while m3; convex anterior symphysis without symphyseal Perivolaki is younger and its magnetostratigraphic study constriction; small and deeply inclined planum al- suggests an age between 7.3-7.1 Ma. veolare; absent or weak fossa genioglossa; small lingual cusp in the P3, 4; small honing facet in the p3; small CHARACTERISTICS. — Size intermediate between M. delsoni hypoconulid in the m3. and M. pentelicus; relatively shallow mandibular corpus, similar to M. pentelicus; high internal cusp (protocone) in the P3,4; fl attened anterior symphysis with symphyseal constric- Mesopithecus delsoni tion; slightly inclined backwards alveolar plane; large fossa Bonis, Bouvrain, Geraads & Koufos, 1990 genioglossa; strong honing facet in the p3; large hypoconulid with distal groove in the m3; similar dental dimensions to HOLOTYPE. — Mandible of male adult individual with M. pentelicus; longer bones than M. pentelicus. both tooth rows, RZO-159. It is housed in the Labo- ratory of Geology and Palaeontology, University of Th essaloniki. Mesopithecus aff . M. pentelicus

LOCALITY. — Ravin des Zouaves-5 (RZO), Axios Valley, MATERIAL EXAMINED. — See Bonis et al. (1990). Macedonia, Greece. LOCALITIES. — Dytiko-2, 3 (DIT, DKO), Axios Valley, AGE. — Early Turolian, MN 11 (late Miocene); estimated Macedonia, Greece. magnetostratigraphic age ~8.2 Ma. AGE. — Late Turolian, MN 13 (late Miocene).

DIAGNOSIS. — Large size; deep mandibular corpus; fl at- CHARACTERISTICS. — Relatively smaller orbits with tened anterior symphysis; strong symphyseal constriction; stronger supraorbital torus than M. pentelicus; longer slightly inclined alveolar plane; large fossa genioglossa; sagittal crest than M. pentelicus; smaller incisors; less re- thick inferior transverse torus; large honing facet in the fl ected backwardly epicondylus medialis in the humerus; p3; well-developed and bicuspid talonid in the m3. oval section of the radius shaft; radius with smaller facet for the ulna. Mesopithecus cf. M. delsoni

MATERIAL EXAMINED. — Mandibular fragment with Mesopithecus cf. M. pentelicus symphysis preserving p3-p4 dex and p4-m1 sin, MNHN- slq-940 + 941. MATERIAL EXAMINED. — See Bonis et al. (1990).

GEODIVERSITAS • 2009 • 31 (4) 839 Koufos G. D.

LOCALITY. — Dytiko-1 (DTK), Axios Valley, Macedo- AGE. — Late Ruscinian, MN 15 (early Pliocene). nia, Greece. DIAGNOSIS. — Colobine monkey of moderate size; con- AGE. — Late Turolian, MN 13 (late Miocene). stant mandibular depth; strong sexual dimorphism in the CHARACTERISTICS. — Similar to M. pentelicus but as canines; strong metaconid and protoconid in the relatively there is few material available and it is badly preserved, narrow p4; the mesial breadth of the m3 is larger than the certain determination is diffi cult. the distal one of the m2; the m3’s hypoconulid is situ- ated between the midline and the line joining the buccal cuspids of the tooth; there is no clear fi fth cuspid in the m3’s talonid but there is a cingular elevation indicating Mesopithecus cf. M. monspessulanus its development (Koufos et al. 1991a).

MATERIAL EXAMINED. — Mandibular fragment with l1-m3 sin, LGPUT-DIT-22. Subfamily CERCOPITHECINAE Gray, 1821 LOCALITY. — Dytiko-2 (DIT), Axios Valley, Macedo- Genus Paradolichopithecus nia, Greece. Necrasov, Samson & Radulesco, 1961 AGE. — Late Turolian, MN 13 (late Miocene).

CHARACTERISTICS. — Mandibular and dental dimen- Paradolichopithecus arvernensis sions smaller than M. pentelicus and closer to M. mons- (Depéret, 1929) pessulanus. LOCALITY. — Vatera (VTR), Lesvos Island, Greece.

AGE. — Middle Villafranchian s.l., MN 17 (late Mesopithecus sp. Pliocene).

MATERIAL EXAMINED. — MCIII, LGPUT-NIK-unum- DIAGNOSIS. — Large-sized; clear sexual dimorphism in bered; MtI, LGPUT-1541. the canine’s complex; large anterior lower teeth; lingual surface of the lower incisors without enamel; p4 with LOCALITY. — Nikiti-2 (NIK), Chalkidiki, Macedonia, larger labial than lingual cuspid; m3 with elongated Greece. trigonid, low relief and a large sixth cuspid; eruption AGE. — Early Turolian, MN 11 (late Miocene); more pattern of lower dentition typically papionin. precisely it is referred to the lowermost part of early Turolian (MN 11) between 8.7 and 8.2 Ma. STRATIGRAPHIC DISTRIBUTION CHARACTERISTICS. — Th e morphology and the size of the bones are close to those of Mesopithecus. Th e known Neogene cercopithecids of Greece be- long to three taxa: Mesopithecus, Dolichopithecus Mesopithecus sp. and Paradolichopithecus. Th ere is also evidence for the presence of Macaca sp. in the early Pleistocene MATERIAL EXAMINED. — See Kullmer & Doukas, 1995. locality Tourkobounia 2 (fi ssure fi lling) near Athens LOCALITY. — Maramena (MAR), Serres Basin, Mace- (Symeonidis & Zapfe 1976). Th e latter two Neogene donia, Greece. genera are relatively rare, and known from one or AGE. — Latest Turolian, MN13/14 (late Miocene). two localities: Dolichopithecus is known from the

CHARACTERISTICS. — Similar to M. pentelicus according localities Megalon Emvolon and Ptolemais, both to Kullmer & Doukas (1995). dated to Pliocene (Fig. 11). Th e Megalon Emvolon fauna includes several species suggesting a late Rus- cinian (MN 15) age (Koufos 2006b and references Dolichopithecus ruscinensis Depéret, 1889 therein). Th e Ptolemais Dolichopithecus (two isolated teeth) comes from Notio Mb of Ptolemais Fm.,

LOCALITY. — Megalon Emvolon (MEV), near Th essalo- dated to late Ruscinian, MN 15 (de Bruijn pers. niki, Macedonia, Greece; Ptolemais Basin (PTL), Notio comm. 2008). Paradolichopithecus is only known Mb of Ptolemais Fm., Macedonia, Greece. from the locality of Vatera (Lesvos Island) dated to

840 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

PARADO- MESOPITHECUS DOLICHO- LICHOPI- Ma PITECHUS

Polarity Elma MN zones TECHUS

C2n 2 C2r

3 C2An VILLAFRANCHIAN

MARAMENA: Mesopithecus sp. PTOLEMAIS: D. ruscinensis

C2Ar 4 DYTIKO-1, 2, 3: Mesopithecus aff. or cf. pentelicus Mesopithecus cf. monspessulanus Paradolichopithecus arvernensis VATERA: C3n RUSCINAN PIKERMI: M. pentelicus 5

C3r CHOMATERES: M. pentelicus

6 PERIVOLAKI: M. delsoni-M. pentelicus C3An

C3Ar

7 C3Bn MEGALON EMVOLON: D. ruscinensis C3Br VATHYLAKKOS-2, 3: M. delsoni-M. pentelicus TUROLIAN C4n ?RAVIN-X: M. cf. delsoni 8 RAVIN DES ZOUAVES-5: M. delsoni MN 11 MN 12 MN 13 MN 14 MN 15 MN 16 MN 17 C4r NIKITI 2: Mesopithecus sp.

C4An Mesopithecus sp. ?KRYOPIGI: 9

FIG. 11. — Stratigraphic distribution of the cercopithecids in the Neogene of Greece. the latest Pliocene, MN 17 (Fig. 8) at ~2.0 Ma (de (NIK), dated biochronologically at the lowermost Vos et al. 2002; Lyras & Van der Geer 2007). early Turolian, MN 11 (Koufos 2006b and refer- On the contrary, Mesopithecus is very common in ences therein). Th e available Mesopithecus material Greece and except Pikermi is known from several from NIK includes some postcranials, which cannot other localities covering whole Turolian (Fig. 11). Its allow specifi c determination at the moment; thus earliest occurrence is traced in the locality Nikiti-2 it is referred to as Mesopithecus sp. Another early

GEODIVERSITAS • 2009 • 31 (4) 841 Koufos G. D.

A 60

55 NIK

50 R-X

PER 45

40 RZO DYTI VATH 35

30 % bovids + giraffids

25 CHO

PIK 20 MAR 15 5 5,5 6 6,5 7 7,5 8 8,5 9 Age (Ma) B 9,40

9,30 PER RZO CHO VATH 9,20 PIK R-X

DYTI 9,10

Ln body mass 9,00

DIT 8,90

M. monspessulanus Europe (Delson 1973) 8,80 4,5 5 5,5 6 6,5 7 7,5 8 8,5 Age (Ma)

FIG. 12. — A, distribution of the bovids + giraffi ds in the Mesopithecus Wagner, 1839 bearing mammal localities of Greece versus their geological age; B, distribution of Mesopithecus body mass in the Mesopithecus bearing mammal localities of Greece versus their geological age. Abbreviations: see text. trace of Mesopithecus comes from the locality RZO Greece by a to medium-sized form having characters of Axios Valley with the large-sized taxon M. delsoni between M. delsoni and M. pentelicus and referred to dated to early Turolian (MN 11); more precisely the as M. delsoni/pentelicus. It is known from the locali- palaeomagnetic record suggests an age of ~8.2 Ma ties VAT and VTK of Axios Valley, dated to middle (Koufos 2006b; Sen et al. 2000). During middle Turolian at ~7.3 Ma (Koufos 2006b, unpubl. data Turolian (MN 12), Mesopithecus is represented in and references therein). Th is form is also known

842 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

TABLE 2. — Faunal composition by taxonomy (% number of species per family or group) of the Mesopithecus Wagner, 1839 bearing mammal localities of Greece. Abbreviations: see text.

NIK RZO VATH R-X PER CHO PIK DYTI MAR MEV VTR Rodentia 044003817050130 Primates 844745283136 Carnivora 15 22 24 14 23 5 36 8 15 13 18 Tubulidentata 0000000400 0 Hyracoidea 0000002000 0 Proboscidea 8747457123012 Equidae 13 14 8 7 13 21 6 38 13 8 6 Rhinocerotidae 0447455430 6 Chalicotheriidae 0480052400 0 Suidae 074745243130 Tragulidae 0040002400 0 Cervidae 0000452430 6 Giraffi dae157878558306 Bovidae 38 30 28 43 38 19 17 28 15 38 41 from the locality PER, dated to middle Turolian too Having in mind all the above mentioned we can but older than Pikermi; the palaeomagnetic record conclude to the following: suggests an age between 7.3-7.1 Ma (Koufos et al. – Mesopithecus appeared in Greece at the beginning 2006). Th e middle-sized M. pentelicus is smaller of Turolian with the large-sized species M. delsoni; than M. delsoni and M. delsoni/pentelicus and ap- – a large-medium size form named M. delsoni/ peared in the localities of Pikermi and Chomateres pentelicus is known from middle Turolian; at the end of middle Turolian (Fig. 11). During – the medium-sized M. pentelicus appeared at the late Turolian (MN 13) Mesopithecus was present by end of middle Turolian and probably exists in late two forms found in the localities of Dytiko, Axios Turolian; Valley (Fig. 11). Th e fi rst form is slightly smaller – the small-sized M. monspessulanus, which is than M. pentelicus with some minor diff erences considered as Ruscinian, seems to appear earlier (see above), while the other is clearly smaller and during the uppermost Turolian; closer to the small-sized M. monspessulanus. Th e – there is a size decrease of Mesopithecus during sole known and small-sized mandibular branch Turolian which can be used as a biostratigraphic from DIT (Fig. 8) indicates an early appearance of tool; M. monspessulanus at the end of Miocene. A similar – Dolichopithecus in Greece is known from late size decrease is also observed in the late Turolian Ruscinian (MN 15) but it is quite possible that it Mesopithecus of Italy (Rook pers. comm. 2008). exists in older levels, since the Ruscinian mammal Th e youngest Greek locality with Mesopithecus is localities are very rare in Greece; Maramena dated to the end of Turolian (Schmidt- – Paradolichopithecus is only known from the end Kittler et al. 1995; Koufos 2006c). Th e majority of Villafranchian (MN 17), but it is possible that of the Maramena sample includes isolated teeth it exists in younger levels. and thus it is diffi cult to compare them with the rest of the Greek material. Th erefore, although it was described as M. pentelicus, it is better to refer PALAEOECOLOGY it as Mesopithecus sp. for the moment. Th e age of the locality Kryopigi in Chalkidiki is not certainly Th e palaeoecology of Mesopithecus has been discussed known, as the material is still undescribed and it quite earlier when Gaudry (1862-1867) described is better to date it to late Miocene for the moment the sample from Pikermi; he proposed a more ter- (Fig. 11). restrial than arboreal way of life for it. Later on,

GEODIVERSITAS • 2009 • 31 (4) 843 Koufos G. D.

TABLE 3. — Modern and fossil faunas used in the Correspondance Factor Analysis of Figure 11.

Locality Abbreviation Country Environment Makokou MAK Gabon Tropical forest La Maboke MAB Republic of Central Africa Tropical forest Sangmelina SNG Cameroon Tropical forest Mont Kivu MKV Zair Tropical forest Mont Nimba MNM Libria, Guinea Tropical forest Transvaal 9 TRA9 South Africa Savannah Amboseli AMB Kenya Savannah Region Gabiro RGA Rwanda Savannah Ihema IHE Rwanda Savannah Transvaal 7 TRA7 South Africa Savannah Zinawe ZNW Mozambuque Savannah Lokori LOK Kenya Savannah Golden Gate GGT South Africa Savannah Eppelsheim EPP Germany Open Hostalets HST Spain Open La Roma LRO Spain Open Los Valles de Fuentinueva LVF Spain Open Can Ponsic PON Spain Open Rudabanya RUD Hungary Open Terassa TRS Spain Open Villadecabals VDC Spain Open several authors studying Mesopithecus, proposed a large groups from one region to another in order semi-terrestrial way of life. Th eir results were based to fi nd food. Th e open character of the landscape mainly to the postcranial proportions of the animal, is strengthened by the high percentage of equids as well as to their comparison with recent cercop- (Fig. 10), the majority of which belongs to running ithecoids (Gabis 1961; Szalay & Delson 1979; Zapfe forms (elongated and slender metapodials). Th e 1991; Youlatos 2003; Koufos et al. 2003). However main rhino of the Mesopithecus bearing localities Escarguel (2005) does not agree that the distal part is “Diceros” neumayri (Osborn, 1900), which also of the hindlimb is discriminative for the way of life characterizes open-dry environments (Giaourtsakis of Mesopithecus. Th e genus Mesopithecus is quite well 2009). Th e suids, tragulids, tapirids and cervids are known from Greece covering the whole Turolian. absent or very rare, confi rming the above mentioned Most of the Mesopithecus bearing localities include open-dry conditions. a rich fauna. Except Pikermi and Ravin-X faunas, Th e comparison of the bovids and giraffi ds per- the others are new collections well correlated to the centage with the geological time suggests a linear stratigraphy and without any admixture. Th e study relation, indicating a decrease of their taxa during and comparison of these faunas can help to the Turolian (Fig. 12A). Th e decrease of the bovids determination of the landscape where Mesopithecus and giraffi ds could indicate a less open and dry pal- was living and consequently of its way of life. aeoenvironment to the end of Miocene. A relatively Th e faunal composition (% taxa per family or more wet with closed spots environment (patchy taxonomic group) of the Mesopithecus bearing lo- environment) has been reported for the late Turolian calities of Greece is given in Table 2. In all these of Eastern Mediterranean region (Koufos 2006c). localities the bovids + giraffi ds dominate; their Th is change of the palaeoenvironment seems to percentage varies between 18 and 58%. Th e domi- agree with the size decrease of Mesopithecus during nance of these two groups suggests a relatively Turolian, leading to the small M. monspessulanus, open environment (wooded or bushed savannah) a form which is more arboreal (Szalay & Delson in which it is possible for these animals to move in 1979). Th is size decrease of Mesopithecus is clear

844 GEODIVERSITAS • 2009 • 31 (4) Neogene cercopithecids (Mammalia, Primates) of Greece

NIK 0.6 GGT PER AMB FOSSIL OPEN RZO R-X FOSSIL CLOSED TRA9 LOK MAR RUV ZNW VATH SER PIK DYTI 0.2 RGA CHO IHE TRA7

-2 -1.6 -1.2 -0.8 -0.4 0.4 0.8 1.2 1.6 LRO

Axis 2 (22.8%) VDC HST -0.2 LVF MNM

TRS -0.4 MAK MAB -0.6 PON MKV EPP SNG -0.8

RUD RECENT OPEN RECENT CLOSED Axis 1 (43.9%)

FIG. 13. — Correspondence Factor Analysis of the Mesopithecus Wagner, 1839 bearing Greek Turolian mammal localities with a set of Vallesian ones from Europe and a set of modern ones from certain environments (Table 2). Abbreviations: see text. in the diagram of Figure 12B in which the linear Eastern Mediterranean faunas has been reported relation between the body mass and the geological in several recent works (Bonis et al. 1992, 1994, age of Mesopithecus is quite clear. Th e body mass of 1999; Merceron et al. 2004, 2005a, b; Koufos et Mesopithecus was calculated using the dimensions of al. 2006, 2009; Kostopoulos, 2009). the m1 according to Legendre (1988: 18, 240). All the available data for the Turolian advocate to Th e open character of the Turolian landscape an open-dry environment where Mesopithecus was is also confi rmed by the Correspondence Factor living. An open environment could be more or less Analysis (CFA) of the Mesopithecus including Greek bushy, bosky with or without gallery forests. In such mammal faunas with modern faunas from certain an environment, a terrestrial rather than arboreal environments and with a set of Vallesian Western way of life is more possible for Mesopithecus. Th e and Central European faunas (Table 3; Fig. 13). Th e open bushy or bosky areas provided enough food to software PAST is used for the multivariate analysis Mesopithecus, especially during the rainfalls period. (Hammer et al. 2001). During Vallesian in Western On the other hand the sparse trees protected it from and Central Europe the conditions were relatively the carnivores and provided a place for passing safely closed and wet (Bonis et al. 1992, 1999; Agustí et the night. Th e dental microwear of M. pentelicus, al. 1999, 2003; Koufos 2006d). Th e analysis indi- M. delsoni and M. delsoni/pentelicus indicates that cates that axis-1 distinguishes clearly the modern their feeding habits are alike, including leaves, seeds, closed and open faunas, while axis-2 separates the fruits or underground vegetal parts (Merceron et modern from the fossil faunas. All the Mesopithecus al. 2009, this volume). Such feeding preferences bearing localities of Greece match together and can are in accordance with their locomotion and the be clearly correlated with the modern open faunas, determined environment. indicating similar palaeoenvironmental conditions. Concerning Ruscinian, the available mammal Th e open-dry character of the Turolian Greek and localities of Greece and Eastern Mediterranean are

GEODIVERSITAS • 2009 • 31 (4) 845 Koufos G. D.

very scarce with limited data. Th e fauna of Megalon (NSF-RHOI). Financial support to visit various Emvolon is very poor to allow certain comparisons museums and institutes was provided to me by the and results. However, there are some indications European Commission’s Research Infrastructure for a signifi cant faunal change at the Miocene/ Action (EU-SYNTHESYS: AT-TAF-702, FR- Pliocene boundary with the extinction of several TAF-3102, GB-TAF-1842) and the NSF-RHOI. Miocene faunal elements. Th is faunal change could I am indebted to N. Symeonidis, M. Dermitzakis, be related to climatic reasons. A change to more G. Th eodorou, C. Doukas, S. Roussiakis, A. Van wet and closed conditions was observed from late der Geer (AMPG), K. Heissig (BSPM), G. Daxner- Turolian (Koufos 2006c) which probably extended Höck, M. Harzhauzer (NHMW), C. Soligo, A. to Ruscinian. Although the faunal elements from Currant, J. Hooker (BMNH), and P. Tassy, S. Sen, Greece are few, the extended Ruscinian lignitic de- C. Sagne, C. Argot (MNHN) for giving me access posits of northern Greece (Koukouzas et al. 1979) to the collections at their disposal and for their and neighbouring countries indicate closed and great hospitality and help during my visit to their wet conditions. Th us one can suppose that Doli- institutes. Many thanks to my collaborators D. S. chopithecus could probably live in a relatively closed Kostopoulos, T. D. Vlachou, G. E. Konidaris and and wet environment. However, such conclusion I. A. Sylvestrou for participating in all expeditions is doubtful and only the discovery of more fossils and helping a lot to the excavations and preparation from that period and their study will provide the of the fossils. Th anks are also due to several col- available data for a certain determination of the leagues, postgraduate and undergraduate students environment. participating in the excavations all these years and Th e sole known occurrence of Paradolichopithecus helping me to create these collections (Axios Valley, is traced in the latest Pliocene locality of Vatera, Nikiti, Perivolaki, Megalon Emvolon). Th anks to Lesvos Island. Th e fauna of Vatera includes elements D. Begun, N. Spassov and A. Ohler for reviewing of open habitats (savannah-like) (Van der Geer & the manuscript. Sondaar 2002). In fact the faunal composition of the known Vatera fauna with a lot of bovids and giraffi ds (Table 3) indicates a relatively open envi- REFERENCES ronment. Th e analysis of the Villafranchian faunas of southern Europe and western Asia with those of ABEL O. 1927. — Lebensbilder aus der Tierwelt der Vorzeit. Verlag von Gustav Fischer, Jena, 679 p. Greece suggests a relatively open/mixed character AGUSTí J., CABRERA L., GARCÉS M., LLENAS M. 1999. — for the late Villafranchian faunas (Kostopoulos & Mammal turnover and global climate change in the Koufos 2000). Considering this interpretation, a late Miocene terrestrial record of the Vallés-Penedés similar environment must be referred for Paradoli- basin (NE Spain), in AGUSTÍ J., ROOK L. & ANDREWS chopithecus. Such an environment fi ts quite well P. (eds), Hominoid Evolution and Climatic Change in Europe vol. I: Th e Evolution of the Neogene Terrestrial with the terrestrial way of life proposed for Para- Ecosystems in Europe. Cambridge University Press, dolichopithecus (Szalay & Delson 1979). New York: 397-412. AGUSTÍ J. A., SANZ DE SIRIA A. & GARCÉS M. 2003. — Explaining the end of the hominoid experiment in Acknowledgements Europe. Journal of Human Evolution 45: 145-153. ARAMBOURG C. & PIVETEAU J. 1929. — Les Vertébrés I would like to thank the editors for their invita- du Pontien de Salonique. Annales de Paléontologie tion to participate in this volume of Geodiversitas 18: 59-138. published in honour of Prof. L. de Bonis. Th e exca- ARMOUR BROWN A., DE BRUIJN H., MANIATI C., SIA- vations in Axios Valley and Nikiti were fi nancially TOS G. & NIESEN P. 1977. — Th e geology of the supported by the Leakey Foundation, the Fonda- Neogene sediments north of Serrai and the use of rodent faunas for biostratigraphic control. Proceed- tion Signer-Pollignac, the University of Th essalo- ings of 6th Colloquium on the geology of Aegean Region niki and the USA National Research Foundation 2: 615-622. project: Research for Human Origins Initiative BONIS L. DE & KOUFOS G. D. 1999. — Th e Miocene

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Submitted on 8 December 2008; accepted on 30 September 2009.

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