George D. Koufos

Aristotle University of Thessaloniki | AUTH · Department of Geology (GGG)

The Late Miocene vertebrate localities in the Axios Valley (Macedonia, Greece) were discovered at the beginning of the last century by the French geologist Camille Arambourg, who made a large fossil collection and carried it back to France. During the last 50 years, extensive fieldwork have been carried out by a Greek–French team, which has yielded...

The hominoid Ouranopithecus macedoniensis (Bonis, Bouvrain, Geraads & Melentis, 1974) is known from three Late Miocene localities in Greece. All are correlated with late Vallesian, MN10; more precisely, they are dated between 9.6 and 8.7 Ma. During the last 50 years several fossils of this hominoid have been recovered. The present article describes...

The present paper deals with the ecomorphological characteristics of two sympatric species of ictitheres: Ictitherium viverrinum Roth & Wagner, 1854 and Hyaenictitherium wongii (Zdansky, 1924) from the Late Miocene of the Eastern Mediterranean. These two species represent different taxonomic groups with allegedly distinct ecological roles. The valu...

The late Miocene is a period of great climatic and ecosystemic changes. The south-eastern Mediterranean region is particularly affected, with increasing aridity and habitat openness. The impact of these changes on rhinocerotids’ ecology has not been studied, although rhinoceroses were a concupious and diverse component of the Miocene mammalian asse...

‘Arambourg collection’ is a collection of mainly mammal fossils from Axios Valley (Macedonia, Greece) made by the French geologist C. Arambourg during 1915–16. It is housed in the Muséum national d’Histoire naturelle of Paris. New excavations in Axios Valley during the last 50 years allowed the collection of many fossils and a new taxonomy of hippa...

In the published manuscript, the study presents diverse geochronological and biochronological data providing age constraints on the site of Tsiotra Vryssi (Mygdonia basin, Greece). One of the methods presented is based on burial ages from cosmogenic radionuclides. Table 2 of this study reports cosmogenic simple burial ages of 1.88 ± 0.16 Myr, 2.10...

Numerous fieldwork campaigns, since 1977, in the Lower Pleistocene fossiliferous deposits of Mygdonia Basin (Greece), led to the discovery of several sites, which are located in distinct stratigraphic horizons and correlated to different parts of the late Villafranchian-Epivillafranchian. Their rich and diversified fauna of different evolutionary s...

The mammal fauna from Libakos was collected by H. Eltgen (Technical University of Clausthal, Germany) during the end of the 1960s and studied by K. Steensma. The locality is situated within the fluviolacustrine-fluvioterrestrial deposits of the Grevena–Neapolis basin (Macedonia, Greece), which are considered as Pliocene-Pleistocene. The carnivoran...

The mephitids are carnivoran family that includes the extant skunks, stink badgers, and the extinct related forms. The family appears in the fossil record at the beginning of the Miocene and today occurs in America (skunks), Indonesia, and Philippines (stink badgers). In the Greek fossil record, the mephitids are represented by the genus Promephiti...

The systematic position of the ailurids was debated for a long time but recent molecular studies indicated that they constitute an independent family within Musteloidea. They include eight extinct taxa and only one extant species, the red panda Ailurus fulgens. In the Greek fossil record, there is only one species, Simocyon primigenius, found in it...

The viverrids are known from the entire Old World but their fossil remains in Greece are rare. They are known from only two fossiliferous sites Aliveri in Euboea Island and Thymiana-B in Chios Island. The new genus and species Euboictis aliverensis was found in Aliveri, dated to the end of the late Miocene. The locality Thymiana-B provided the rela...

The family Percrocutidae was erected to include some Miocene and Pliocene hyaena-like carnivoran taxa of Eurasia and Africa, which were included to the Hyaenidae for a long time. The percrocutids were dispersed from Spain to China and in the Eastern Mediterranean region are traced in Serbia, Bulgaria, and Turkey. In Greece, Percrocutidae are scarce...

The felidae constitutes a large carnivoran family with several living and fossil species, which appeared at the end of the Oligocene and dispersed into the Old and New World. In Greece, the felids are quite common in the Neogene–Quaternary deposits including several species. The earliest occurrence of the family in Greece is dated at the early/midd...

The fossil hyaenids are abundant and diverse, in the Greek fossil record. The hyaenids originate from Viverridae and occur at the beginning of Miocene with some small-sized and dog-like arboreal forms. The earliest hyaenids of Greece appeared in the early/middle Miocene with the genus Protictitherium. During the late Miocene, more than ten species...

The family Canidae includes a variety of carnivoran taxa, which appeared in the late Eocene, about 40.0 Ma ago. It is worldwide distributed, except Antarctica, including several fossil and extant species. In Greece, it is known from Pliocene and Pleistocene; today it is represented by the gray wolf, golden jackal, and red fox. The oldest known cani...

The mustelids constitute a quite rich and diversified family of carnivorans, including a variety of modern taxa (e.g., badgers, otters, minks) and related fossil forms. The family appeared at the end of the Oligocene in the Old World and then dispersed to America. In Greece, the earliest evidence of mustelids is dated at the early/middle Miocene bo...

Family Equidae includes all the modern horses, zebras, donkeys and a great number of related species. The equids appeared with the genus Hyracotherium (= Eohippus) in the Early Eocene and they are still present with the modern wild and domestic representatives of the genus Equus. In Greece, the equids appeared in the early Miocene, but the bulk of...

The hyraxes are a peculiar group of mammals displaying close relationships with the elephants and sirenians. They appeared at the Eocene but later, during the Late Eocene–Oligocene, diversified with several taxa. Their number decreased at the beginning of the Miocene with the increase of the artiodactyls. In Greece, the pliohyracids are rare with a...

Tubulidentata or aardvarks is a group of mammals characterized by the tubular structure of their dentition and their lack of enamel. They also have peculiar external features superficially similar to that of pigs, rabbits, rats, and kangaroos. Their oldest occurrence is in the early Miocene of East Africa, and they are represented today by the sing...

Cercopithecidae or “Old World monkeys” is a group of derived Primates (Catarrhini) recognized in Greece with the genera Mesopithecus, Dolichopithecus, and Paradolichopithecus. The first dominates in the fossil record with the species M. pentelicus, which is present in several late Miocene sites. The large-sized species M. delsoni and the small-size...

The informal group “Miocene hominoids” comprises a primate group of special interest as it is considered including the common ancestor of the modern great apes and humans. The group appeared at the beginning of the Miocene in Africa and then dispersed in the Old World with several taxa. In Greece the Miocene hominoids are known by two taxa. The old...

The sedimentary basins of Greece contain an important record of fossil vertebrates that has been known and studied for nearly two centuries. Here, we present our collective effort to review and summarize this fossil record. A combination of our original research and previously published records permits the complete reassessment of the identified ve...

Objectives: To explore mandibular shape differences between Ouranopithecus macedoniensis and a comparative sample of extant great apes using three-dimensional (3D) geometrics morphometrics. Other objectives are to assess mandibular shape variation and homogeneity within Ouranopithecus, explore the effects of size on mandibular shape, and explore th...

The sedimentary basins of Greece contain an important record of fossil vertebrates that has been known and studied for nearly two centuries. Here, we present our collective effort to review and summarize this fossil record. A combination of our original research and previously published records permits the complete reassessment of the identified ve...

Material belonging to the late Miocene hominoid Ouranopithecus macedoniensis has been poorly analyzed using advanced techniques. This study aims to explore mandibular shape variation between Ouranopithecus macedoniensis and a comparative sample of extant great apes, using three-dimensional (3D) geometric morphometrics. Other objectives are to asses...

The remains of equids are abundant in the Early Pleistocene faunas of Greece. "Apollonia-1" is one of the richest localities from the latest Villafranchian, providing eight skulls, mandibular remains and plenty of postcranial material during several field campaigns. This study focuses mainly on the skulls, mandibular remains and metapodials from th...

The late Miocene ictitheres of Greece have been recognised since the beginning of the 19th Century and include several species. The small-sized Plioviverrops and Protictitherium are well-known, but those of middle-size, Ictitherium, Hyaenictitherium and Lycyaena are poorly understood. The aim of this article is to study a significant sample of mate...

Background and scope: The late Villafranchian large mammal age (~2.0–1.2 Ma) of the Early Pleistocene is a crucial interval of time for mammal/hominin migrations and faunal turnovers in western Eurasia. However, an accurate chronological framework for the Balkans and adjacent territories is still missing, preventing pan-European biogeographic corre...

The study of the bovid assemblage from the primate-bearing site of Xirochori-1 in Axios Valley, Northern Greece, reveals the presence of four taxa, namely Gazella sp., Ouzocerus cf. pentalophosi, Helladorcas sp. and ‘Protoryx’ sp., and allows updating the faunal list of the Xirochori-1 Vallesian mammal fauna, dated at around 9.6 Ma. Gazella sp. fro...

Tragulidae are very rare in the Miocene fossil record of Greece and their known material is quite scarce. Some new remains of them, found in the early/middle Miocene locality Antonios in Chalkidiki Peninsula (Macedonia, Greece), are studied in this article. Based on the size, at least three different tragulids have been recognised, Dorcatherium gun...

This paper aims to contribute to the stratigraphic and geochronological evaluation of the primate bearing Dafnero fossil site of Northern Greece by means of lithostratigraphic, paleomagnetic and paleontological analyses. The 60 m thick fossiliferous deposits of fluviatile origin are recognized as representing a typical braided-river sequence unconf...

The Turolian fossil beds of Greece include a high number of contemporary small carnivoran taxa, such as Mustelidae (Promeles, Parataxidea, Martes, Sinictis, Enhydriodon), Mephitidae (Promephitis) and Hyaenidae (Protictitherium, Plioviverrops). Many of these species are found together in some localities, therefore they must have adapted some differe...

The present study deals with two enigmatic carnivores of the Late Miocene of Greece, Plesiogulo crassa and Simocyon primigenius, from an ecomorphological point of view. These species are compared to extant mustelids and canids respectively, in order to provide some suggestions on their diet. Plesiogulo crassa was a large, wolverine-like mustelid. M...

The homotheres, although known from the Greek fossil record since the beginning of the 1930’s, have only been recorded from fragmentary material, sometimes undescribed and with questionable determinations. In 2017, the discovery of an almost complete cranium and a fairly complete humerus of a machairodontine in Dafnero-3, a lower Pleistocene site f...

We here report on fossil remains of the earliest known crown-Testudo, an extant clade of Mediterranean testudinid tortoises from the late Miocene (Vallesian, MN 10) from the hominoid locality Ravin de la Pluie (RPl) in Greece. The material studied is a small, nearly complete carapace with a clearly distinct hypo-xiphiplastral hinge. This supports t...

In the present article, we study the mammoth remains from the late Villafranchian (Early Pleistocene) locality Apollonia-1 (Mygdonia Basin, Northern Greece). The material consists of a hemi-mandible with the m3, isolated m3/M3 and a maxilla fragment with DP2-DP3. The mandibular and dental features permit their attribution to the southern mammoth Ma...

The reconsideration of the hipparions from the locality Ravin des Zouaves-5 (RZO) of the lower Axios Valley provides a better taxonomy, based on the revision of the old known collection (up to 1987), the inclusion of new discoveries and the bibliographic data of the last 30 years related to the study of the hipparions in Greece and the neighboring...

tThe fossil mammal localities of the Axios Valley (Macedonia, Greece) yielded a rich collec-tion of Mesopithecus remains, which were first described at the beginning of the 1990s. Thelate Turolian Dytiko localities include several specimens of Mesopithecus, which were origi-nally separated in two size-related forms: the relatively larger-sized M. c...

Objectives: This study aims to virtually reconstruct the deformed face (XIR-1) and maxilla (RPl-128) of the Late Miocene hominoid Ouranopithecus macedoniensis from Greece, through the application of mirror-imaging and segmentation. Additionally, analysis was conducted through 3D geometric morphometrics, utilizing a comparative sample of fossil hom...

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

In this article, we present new proboscidean remains from the late Miocene (Turolian) of Samos Island (Greece), which are stored in the old Samos collections of Darmstadt, Frankfurt a.M. (Germany), Lausanne (Switzerland), and Vienna (Austria), and originate from the excavations or fossil collections that took place on the island at the end of the n...

By applying advanced spatial statistical methods, spatial taphonomy complements the traditional taphonomic approach and enhances our understanding of biostratinomic and diagenetic processes. In this study, we elaborate on a specific aspect – spatial anisotropy – of taphonomic processes. We aim to unravel the taphonomic history of the Early Pleistoc...

The Villafranchian ursids of Greece are scanty, represented by a few isolated teeth and bones. During our last field campaigns in the Mygdonia Basin (Macedonia, Greece) we discovered an almost complete cranium, as well as some cranial, dental and postcranial remains, which are presented in this article. The new material originates from the locality...

Monitor lizards (genus Varanus) inhabited Europe at least from the early Miocene to the Pleistocene. Their fossil record is limited to about 40 localities that have provided mostly isolated vertebrae. Due to the poor diagnostic value of these fossils, it was recently claimed that all the European species described prior to the 21st century are not...

List of references consulted to score characters for the phylogenetic analysis. (PDF)

Matrix used in the phylogenetic analyses. (NEX)

Lists of the apomorphies of V. marathonensis, of the synapomorphies of the clade including V. marathonensis and the subgenus Indovaranus, and of Varanus (based on analysis 1B). (PDF)

List of extant comparative specimens of Varanus. (PDF)

New characters added to the original list of Conrad et al. [11]. (PDF)

Results of the phylogenetic analysis 1A. (PDF)

Results of the phylogenetic analysis 2A. (PDF)

Size shifts may be a by-product of alterations in life history traits driven by natural selection. Although this approach has been proposed for islands, it has not yet been explored in continental faunas. The trends towards size decrease experienced by some hipparionins constitute a good case study for the application of a life history framework to...

The Greek fossil record of small and large mammal Local Faunal Assemblages is studied and compared by means of cluster analysis techniques using Jaccard similarity index and unweighted pair-group method. The analysis allow recognizing a good arrangement of the Greek LFAs according to time and a main cluster gap, corresponding to an important faunal...

A new fossil cranium of a large papionin monkey from the Lower Pleistocene site of Dafnero-3 in Western Macedonia, Greece, is described by means of outer and inner morphological and metric traits using high-resolution micro-computed tomography. Comparisons with modern cercopithecids and contemporaneous Eurasian fossil taxa suggest that the new cran...

During the last field campaigns in the mammal fossiliferous site Apollonia 1 (Macedonia, Greece), new carnivoran material has been discovered. The new collection added two new carnivoran taxa, Homotherium latidens and Panthera gombaszögensis. The new canid material and the revision of the old one (a) suggest the presence of two Canis species, C. et...

A great number of carnivoran remains from various late Miocene localities of Turkey, housed in the Natural History Museum of Aegean University, Bornova-Izmir, are described and compared with those from Eurasian localities for their determination. Thirteen different taxa were determined and some of them are recognized for the first time in Turkey or...

We here describe lizards and snakes from the late Miocene (MN 10) of Ravin de la Pluie, near Thessaloniki, Greece, a locality widely known for its hominoid primate Ouranopithecus macedoniensis. The new finds comprise two large-sized lizards (a probable anguine and a varanid) and two snakes (an elapid and a small-sized “colubrine”). Even if the mate...

The upper incisor lingual morphology of the late Miocene Greek hominoid Ouranopithecus macedoniensis was almost unknown, as the described earlier maxillary remains preserve only worn incisors. During the most recent excavations in the type locality of Ouranopithecus, Ravin de la Pluie (RPl) of Axios Valley (Macedonia, Greece), four little-worn uppe...

Objectives: Since the beginning of the 1970's, the chronology and biostratigraphy of the continental deposits, filling the Neogene/Quaternary basins of Greece, restricted to some old information for Pikermi, Samos, " Saloniki " and Megalopolis. The information were scarce, doubtful and in many cases without or limited stratigraphic background. Exte...

Objectives: The hominoid Ouranopithecus macedoniensis is known from the late Miocene deposits of Axios Valley (Macedonia, Greece) in the form of several mandibles, a number of teeth and a partial cranium (XIR-1). The face of XIR-1, which was originated from the fossiliferous site Xirochori 1, is well preserved but slightly distorted as a result of...

We report on the earliest modern Testudo from the late Miocene (Vallesian, MN 10) of the hominoid locality, Ravin de la Pluie (RPl), Greece. The material corresponds to a small nearly complete carapace owning a well distinct hypo-xiphiplastral hinge, which supports the sensu stricto generic assignment. This new terrestrial testudinid specimen is ch...

Most basins of Greece were filled with thick Neogene-Quaternary continental deposits, which include a large number of mammal fossiliferous sites. The investigations of the last 40 years in the various basins of Greece led to the discovery of many new fossiliferous sites. The extensive, long time and continuous excavations in the new fossiliferous s...

The Pleistocene vertebrate localities of the Mygdonia Basin (Macedonia, Greece) are known since the end of the 1970s. Numerous fieldwork campaigns, carried out by the Laboratory of Geology and Palaeontology of the University of Thessaloniki, led to the discovery of several fossiliferous sites from which a great amount of fossils has been unearthed...

Though not yet fully adopted by the European paleontological community, the biochronological term Epivillafranchian, originally introduced by Bourdier (1961) and later by Kahlke (2001), represents a particular late Early Pleistocene time interval (appx. 1.2-0.9 Ma) during which a major re-organization of the European (and partly Eurasian) mammal fa...

The Early Pleistocene (late Villafranchian) fossiliferous locality Tsiotra Vryssi (Mygdonia Basin, Macedonia, Greece) was discovered in 2014 and provided a rather rich vertebrate fauna, including mainly mammals, but also some reptiles and birds (Konidaris et al., 2015). The avian remains collected during the 2014-2016 excavations yielded nearly nin...

The fossiliferous site Tsiotra Vryssi (TSR) is located in Mygdonia Basin (Macedonia, Greece) and provided a rather rich vertebrate fauna, including mainly mammals, but also some reptiles and birds. Based on the mammal biochronology, a late Villafranchian age is proposed for TSR, between 1.8 Ma and 1.2 Ma (Konidaris et al., 2015, this volume). Among...

The aim of this study is to virtually restore symmetry to the deformed face of Ouranopithecus macedoniensis (XIR-1) and to reconstruct its facial anatomy using mirror imaging, a virtual anthropology technique. The most important advantage of such methods, relative to traditional reconstruction techniques, is that they allow digital manipulation of...

The fossiliferous sites of Nikiti were discovered on 1990 during the works for the construction of a country road north of the village. The Laboratory of Geology and Palaeontology, University of Thessaloniki started immediately the collection of the material which was continued till 2009. The extensive field work located several fossiliferous sites...

The Pleistocene vertebrate localities of Mygdonia Basin (Greece) are known since the end of the 1970s. Numerous fieldwork campaigns, carried out by the University of Thessaloniki, led to the discovery of several fossiliferous sites from which a great amount of fossils has been unearthed and studied. During a survey expedition conducted by the Unive...

ABSTRACT The locality Küçükçekmece includes a limited hipparion sample mainly consisting of isolated teeth and some postcranial remains. Using various methods, the teeth and postcranials are separated in two different size groups. The first group, characterized by large size, rich enamel plication, deep and thin plis, double-triple pli caballin, ov...

New hominoid teeth from the late Miocene locality Ravin de la Pluie (RPl) of the Axios Valley (Macedonia, Greece) are studied in this article. Their morphology, dimensions and proportions are similar to the hominoid Ouranopithecus macedoniensis, allowing their attribution to this taxon. The studied material provides some new morphological character...

The locality Küçükçekmece includes a limited hipparion sample mainly consisting of isolated teeth and some postcranial remains. Using various methods, the teeth and postcranials are separated in two different size groups. The first group, characterized by large size, rich enamel plication, deep and thin plis, double-triple pli caballin, oval protoc...

The late Miocene avian localities of Greece are rare and the known material is poor. During the last campaigns of excavations in Nikiti 2 a tarsometatarsus of a large struthionid has been unearthed, enriching the poor known Greek sample of late Miocene birds. Morphological and comparative features suggest similarities with the extant and fossil lar...

The study of the new and old collections of the Nikiti vertebrate localities included in this volume provides several new evidences for the taxonomy, composition, chronology and palaeoecology of these late Miocene mammal faunas. The faunal list of both primate bearing localities is enriched and improved by addition of new taxa and revision of older...

The mammal collection of Nikiti 1 (NKT) is relatively poor; it has been already described in a series of publications. During the last field trips in Nikiti (2004-2009), the unearthed material from NKT remained