- Rümelinstrasse 23,
72070 Tübingen
Germany
- Mammalian Paleontology, Vertebrate Paleontology, Proboscidea, Neogene, Biostratigraphy, Palaeoecology, and 11 moreExcavations, Elephants, Mammoth, Pleistocene Vertebrate, Quaternary palaeontology, Miocene, Vertebrate Palaeontology, Systematics (Taxonomy), Late Miocene, Paleontology, and Vertebrate taphonomyedit
- Senior researcher in the DFG-funded project Early Human Adaptations in Megalopolis (MEGALOPOLIS)edit
Located at the eastern corner of Mediterranean Europe, Greece occupies a critical position for mammal dispersals to/from Europe, Asia, and Africa and constitutes a potential passageway towardsWestern Europe. During recent decades,... more
Located at the eastern corner of Mediterranean Europe, Greece occupies a critical position for mammal dispersals to/from Europe, Asia, and Africa and constitutes a potential passageway towardsWestern Europe. During recent decades, numerous fieldwork campaigns in several Pliocene–
Pleistocene sites have greatly enriched the fossil record and provided valuable taxonomic and biostratigraphic data. However, a fully developed reference biochronological unit scheme for the Greek record that could contribute to correlations at a continental scale is still pending. In this article, we provide the updated Late Pliocene to Middle Pleistocene large mammal succession, and we introduce the Faunal Units (FUs) of Greece. We define eight FUs, the Milia, Dafnero, Gerakarou, Tsiotra Vryssi, Krimni, Apollonia, Marathousa, and Apidima FUs (from the oldest to the youngest),
which are determined by a set of first and last local occurrences. The results form the basis for discussion of already set turnovers, dispersals, and extinction/immigration events and showcase the importance of the local record for the investigation of the European terrestrial ecosystems. By developing the first detailed biochronological scheme for the Pliocene–Pleistocene of Southeastern Europe, this study comprises the basis for an expanded Balkan faunal unit scale and a reference framework for future investigations.
Pleistocene sites have greatly enriched the fossil record and provided valuable taxonomic and biostratigraphic data. However, a fully developed reference biochronological unit scheme for the Greek record that could contribute to correlations at a continental scale is still pending. In this article, we provide the updated Late Pliocene to Middle Pleistocene large mammal succession, and we introduce the Faunal Units (FUs) of Greece. We define eight FUs, the Milia, Dafnero, Gerakarou, Tsiotra Vryssi, Krimni, Apollonia, Marathousa, and Apidima FUs (from the oldest to the youngest),
which are determined by a set of first and last local occurrences. The results form the basis for discussion of already set turnovers, dispersals, and extinction/immigration events and showcase the importance of the local record for the investigation of the European terrestrial ecosystems. By developing the first detailed biochronological scheme for the Pliocene–Pleistocene of Southeastern Europe, this study comprises the basis for an expanded Balkan faunal unit scale and a reference framework for future investigations.
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The Balkans are considered a major glacial refugium where flora and fauna survived glacial periods and repopulated the rest of Europe during interglacials. While it is also thought to have harboured Pleistocene human populations, evidence... more
The Balkans are considered a major glacial refugium where flora and fauna survived glacial periods and repopulated the rest of Europe during interglacials. While it is also thought to have harboured Pleistocene human populations, evidence linking human activity, paleoenvironmental indicators and a secure temporal placement to glacial periods is scant. Here, we present the first intra-tooth multi-isotope analysis for the European straight-tusked elephant Palaeoloxodon antiquus, on an adult male individual excavated in association with lithic artefacts at the MIS 12 site Marathousa 1 (Megalopolis basin, Greece). The studied find also exhibits anthropogenic modifications, providing direct evidence of hominin presence. We employed strontium, carbon and oxygen isotope analysis on enamel bioapatite to investigate its foraging and mobility behaviour, using a sequential sampling strategy along the tooth growth axis of the third upper molar, to assess ecological changes during the last decade of life. We found a geographically restricted range, in a C 3-dominated open woodland environment, and relatively stable conditions over the examined timeframe. Our results show that, despite the severity of the MIS 12 glacial, the Megalopolis basin sustained a mesic habitat, sufficient plant cover and limited seasonal fluctuations in resource availability, pointing to its role as a glacial refugium for both fauna and hominins.
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During the Miocene, proboscideans reached their greatest diversification, and due to their marked evolutionary changes in dental size and morphology, they comprise an important biostratigraphic/biochronological tool. In this article, we... more
During the Miocene, proboscideans reached their greatest diversification, and due to their marked evolutionary changes in dental size and morphology, they comprise an important biostratigraphic/biochronological tool. In this article, we study the proboscideans from the Late Miocene hominid locality Hammerschmiede (Germany), whose fossiliferous layers HAM 6, HAM 4 and HAM 5 are dated to 11.42, 11.44 and 11.62 Ma, respectively. The studied material consists of mandibular, tusk and cheek tooth specimens, which are attributed to the deinothere Deinotherium levius and the tetralophodont gomphothere Tetralophodon longirostris. An almost complete juvenile mandible of D. levius was CT-scanned and revealed that the erupting lower tusks represent the permanent ones. The mandible is most possibly associated with a lower deciduous tusk, and therefore these specimens capture the rare, and short in duration, moment of transition between deciduous and permanent lower tusks in fossil proboscideans and represent the first such example in deinotheres. The chronologically well-constrained proboscidean fauna from Hammerschmiede and the examination of other assemblages from European localities indicate that the coexistence of D. levius and T. longirostris characterizes the late Astaracian-earliest Vallesian, while Hammerschmiede may showcase the transition from the Middle Miocene trilophodont (Gomphotherium)-dominated faunas of central Europe to the Late Miocene tetralophodont-dominated ones. Finally, in order to decipher the dietary preferences of the Hammerschmiede Tetralophodon we performed dental mesowear angle analysis, which revealed a mixed-feeding diet with an important browsing component, significantly different from the heavily browsing one of Deinotherium known from other localities. Such distinct feeding habits between the taxa indicate niche partitioning, which allowed their sympatry.
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Mammutidae comprise a proboscidean family that originated in Africa during the late Oligocene, dispersed across the Holarctic during the Miocene, and survived in North America until the end of the Pleistocene. Despite their long... more
Mammutidae comprise a proboscidean family that originated in Africa during the late Oligocene, dispersed across the Holarctic during the Miocene, and survived in North America until the end of the Pleistocene. Despite their long evolutionary history and wide geographic distribution mammutids are particularly scarce in the Miocene of Eurasia. Here, we present a new mammutid specimen (an upper deciduous premolar) from the Upper Miocene locality of Sazak in southwestern Turkey. Morphological and metric traits of the tooth, in particular the well-expressed zygodonty, are distinct from the more basal Zygolophodon and permit its assignment to the more derived “Mammut.” Due to the absence of more diagnostic specimens, a specific attribution is not possible; however, considering the Turolian age of the associated fauna an attribution to the Late Miocene representative of the genus, “Mammut” obliquelophus, is possible. Turolian mammutids are rare in the fossil record and therefore our knowledge remains only fragmentary. Despite the existence of a single specimen, the presence of this genus in Sazak corresponds to its first report in the Upper Miocene of Turkey, as well as the first one in western Asia. The presence of “Mammut” in the Upper Miocene of China was recently confirmed, and therefore the record of “Mammut” at Sazak, i.e., at the western margin of Asia, not only adds to the scanty record of the genus in the Upper Miocene of Eurasia but also provides another line of evidence of the paleozoogeographic link enabling Europe–East Asia proboscidean interchanges during the Late Miocene.
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In this article we present the new open-air Middle Pleistocene locality Marathousa 2, which was discovered during a double intensive and targeted field survey in the lignite mines of the Megalopolis Basin (Greece). The locality is... more
In this article we present the new open-air Middle Pleistocene locality Marathousa 2, which was discovered during a double intensive and targeted field survey in the lignite mines of the Megalopolis Basin (Greece). The locality is situated just below the Lignite Seam III of the Marathousa Member (Choremi Formation), and its similar stratigraphic position to the chronologically well-constrained locality Marathousa 1 (dated between 500 and 400 ka and correlated to Marine Isotope Stage 12) indicates a comparable age for Marathousa 2. The locality most
notably yielded dental and postcranial remains of Hippopotamus, with those found at the locality’s Area A most likely belonging to a single individual. The dimensions of the dental elements support an attribution to Hippopotamus antiquus, a large-sized taxon, almost twice as large as the recent Hippopotamus amphibius, and with strong aquatic habits. Cut marks identified on a thoracic vertebra indicate hominin butchering of the carcass. Preliminary paleoenvironmental remarks are based on the mammal and mollusk fauna. The latter is represented solely by freshwater taxa and is dominated by Valvatidae (Valvata cristata, Valvata studeri) and Bithyniidae (Bithynia candiota), but includes also, in lower percentages, Sphaeriidae, Lymnaeidae, Planorbidae and Unionidae. The available
paleoenvironmental evidence indicates that hominin activities took place at the margins of or near, a likely cold, freshwater body, but with temperatures that allowed its persistence throughout the year, and perhaps under drier conditions compared to preceding and succeeding periods in the basin. In addition to the exploitation of elephants in Marathousa 1, Marathousa 2 provides further evidence of megafauna exploitation during the Middle Pleistocene in the Megalopolis Basin and one of the few examples of hippopotamus carcass processing during the
Lower Paleolithic in Europe. Therefore, it advances our knowledge of the food acquisition strategies and subsistence behavior of Pleistocene Homo, and highlights the need for further research in the basin, where megafauna are well recorded.
notably yielded dental and postcranial remains of Hippopotamus, with those found at the locality’s Area A most likely belonging to a single individual. The dimensions of the dental elements support an attribution to Hippopotamus antiquus, a large-sized taxon, almost twice as large as the recent Hippopotamus amphibius, and with strong aquatic habits. Cut marks identified on a thoracic vertebra indicate hominin butchering of the carcass. Preliminary paleoenvironmental remarks are based on the mammal and mollusk fauna. The latter is represented solely by freshwater taxa and is dominated by Valvatidae (Valvata cristata, Valvata studeri) and Bithyniidae (Bithynia candiota), but includes also, in lower percentages, Sphaeriidae, Lymnaeidae, Planorbidae and Unionidae. The available
paleoenvironmental evidence indicates that hominin activities took place at the margins of or near, a likely cold, freshwater body, but with temperatures that allowed its persistence throughout the year, and perhaps under drier conditions compared to preceding and succeeding periods in the basin. In addition to the exploitation of elephants in Marathousa 1, Marathousa 2 provides further evidence of megafauna exploitation during the Middle Pleistocene in the Megalopolis Basin and one of the few examples of hippopotamus carcass processing during the
Lower Paleolithic in Europe. Therefore, it advances our knowledge of the food acquisition strategies and subsistence behavior of Pleistocene Homo, and highlights the need for further research in the basin, where megafauna are well recorded.
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The Early Pleistocene mammal communities of Europe are characterised by a great diversity of large carnivorans. Among them, the largest ever hyaenid, Pachycrocuta brevirostris, a fierce predator with great bone-cracking adaptations that... more
The Early Pleistocene mammal communities of Europe are characterised by a great diversity of large carnivorans. Among them, the largest ever hyaenid, Pachycrocuta brevirostris, a fierce predator with great bone-cracking adaptations that has left its taphonomic signature on several fossiliferous sites. Here, we perform a rigorous taphonomic analysis focusing on bone surface modifications and damage patterns on large-sized ungulate bones from the site Tsiotra Vryssi (1.78 to ~1.5 Ma; Mygdonia Basin, Greece), aiming to identify the main biotic agent responsible for the modifications. Results reveal significant carnivore ravaging of the assemblage, and selective consumption of bones/bone portions related to nutrient value. Comparisons with modifications on similar-sized ungulate carcasses produced by extant and extinct predators, and similarities with Pachycrocuta-modified assemblages, indicate that Pachycrocuta was the principal agent of modification. Overall, this study not only provides taphonomic evidence for the interpretation of Tsiotra Vryssi, but also offers insights into the palaeobiology, and particularly carcass consumption behaviour
of the giant hyaena. Hence, it advances our knowledge on carnivoran guild dynamics and prey–predator relationships during this epoch and has important implications for the investigations of the subsistence behaviour of the meat-eating hominins, who entered Eurasia at ~1.8 Ma, roughly synchronously with Pachycrocuta.
of the giant hyaena. Hence, it advances our knowledge on carnivoran guild dynamics and prey–predator relationships during this epoch and has important implications for the investigations of the subsistence behaviour of the meat-eating hominins, who entered Eurasia at ~1.8 Ma, roughly synchronously with Pachycrocuta.
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In this paper, we present the new Lower Pleistocene vertebrate site Krimni-3 (KMN) in the Mygdonia Basin (Central Macedonia, Greece), one of the most important and richest fossiliferous basins in southeastern Europe. The ongoing since... more
In this paper, we present the new Lower Pleistocene vertebrate site Krimni-3 (KMN) in the Mygdonia Basin (Central Macedonia, Greece), one of the most important and richest fossiliferous basins in southeastern Europe. The ongoing since 2019 research at KMN revealed a rather rich and diversified fauna, and the first results on its stratigraphy, faunal composition, taphonomic context and chronology are provided herein.
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This study investigates the community structure, dynamics and evolution of the guilds of large carnivorans during the Pleistocene of Europe. Emphasis is given to important renewals, the composition of the guilds in terms of dietary... more
This study investigates the community structure, dynamics and evolution of the guilds of large carnivorans during the Pleistocene of Europe. Emphasis is given to important renewals, the composition of the guilds in terms of dietary preferences and foraging strategies, and to intraguild competition for access to food resources. For this purpose, cluster, principal component, and guild structure analyses are performed combining four ecological/behavioural parameters —body mass, diet, prey acquisition strategy, sociality— of large carnivorans that practice hunting and/or scavenging on large prey. Results show only minimal niche overlap, indicating that large predators may have reduced/avoided competition by almost exclusively occupying different niches, i.e. they did not compete for the same resources and/or employed different foraging strategy. Such niche partitioning and competition avoidance may have reduced the occurrences of potential trophic conflict and could explain the cooccurrence of a high diversity of large predators within the same broad feeding guild. Furthermore, the major predator guild remodeling took place close to the Early/Middle Pleistocene transition, when previously
dominant carnivorans (e.g. Pachycrocuta, Megantereon, Acinonyx, Panthera gombaszoegensis) went extinct and new immigrants arrived (e.g. Panthera spelaea, Panthera pardus, Crocuta crocuta), forming the Galerian to Late Pleistocene guilds. Finally, the inclusion of the meat-eating Homo in the carnivore guild is discussed, including its possible
impact to the demise of carnivoran diversity and accordingly of the several large carnivoran niches towards the end of the Pleistocene.
dominant carnivorans (e.g. Pachycrocuta, Megantereon, Acinonyx, Panthera gombaszoegensis) went extinct and new immigrants arrived (e.g. Panthera spelaea, Panthera pardus, Crocuta crocuta), forming the Galerian to Late Pleistocene guilds. Finally, the inclusion of the meat-eating Homo in the carnivore guild is discussed, including its possible
impact to the demise of carnivoran diversity and accordingly of the several large carnivoran niches towards the end of the Pleistocene.
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Proboscideans (Mammalia: Proboscidea) originated during the Eocene (perhaps already during the Paleocene) in Africa. Their fossil record narrates an amazing evolutionary history, ranging from the Paleogene to the Quaternary. Proboscideans... more
Proboscideans (Mammalia: Proboscidea) originated during the Eocene (perhaps already during the Paleocene) in Africa. Their fossil record narrates an amazing evolutionary history, ranging from the Paleogene to the Quaternary. Proboscideans experienced in the past a great diversification and wide distribution in Africa, Europe, Asia, and the Americas. They persist until today with only two genera, Loxodonta and Elephas, geographically confined in regions of Africa and Asia, respectively. The review of the fossil record of the Neogene proboscideans (excluding the members of Elephantidae that are treated elsewhere) in Greece revealed the presence of deinotheres (Deinotheriidae), mammutids (Mammutidae), choerolophodonts (Choerolophodontidae), amebelodonts (Amebelodontidae), tetralophodont gomphotheres (Gomphotheriidae), and stegodonts (Stegodontidae) in more than fifty localities, ranging from the early Miocene to the Early Pleistocene. Fourteen taxa are here considered valid, three of them (Choerolophodon chioticus, C. pentelici, and Konobelodon atticus) erected from type localities in Greece. The most diverse localities are Pikermi and Samos, where at least four proboscidean species have been recorded. The peak in taxonomic diversity occurred during the Turolian (late Miocene). The Greek proboscidean fossil record contains several highlights. The earliest appearance of the family Deinotheriidae in Europe is documented in Gavathas of Lesvos Island, and it is the proboscidean family with the widest temporal distribution in Greece. A deinotheriid skull from Samos Island is so far the most complete juvenile one known from Eurasia and Africa. Choerolophodon presents the widest temporal distribution among the genera in Greece, and where present, it is the dominant genus in terms of abundance. The rich choerolophodont sample allows the distinction into evolutionary stages and renders the genus as biostatigraphically important for Southeastern Europe. The late Miocene Anancus from Chomateri represents the first appearance of the genus in Greece and one of the earliest occurences in Europe. The sample of the late Pliocene Mammut borsoni from Milia, Grevena, is the richest one of this species, including partial skeletons, the longest upper tusks ever recorded in the world and the most complete mandible in Europe. During the Pliocene–Early Pleistocene, the most frequent and widespread proboscidean is the last European gomphothere Anancus arvernensis. Finally, the Siatista Stegodon is the first evidence of the presence of stegodontids in Europe.
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In this article, we describe an almost complete macaque mandible from the Middle Pleistocene locality Marathousa 1 in the Megalopolis Basin of southern Greece. The mandible belonged to a male individual of advanced ontogenetic age and of... more
In this article, we describe an almost complete macaque mandible from the Middle Pleistocene locality Marathousa 1 in the Megalopolis Basin of southern Greece. The mandible belonged to a male individual of advanced ontogenetic age and of estimated body mass ~13 kg. Comparative metric analysis of its teeth permits its attribution to the Barbary macaque Macaca sylvanus, a species that was geographically widely distributed in Western Eurasia during the Plio-Pleistocene. The dental dimensions of the Marathousa 1 macaque fit better within the variation of the Early Pleistocene M. s. florentina and the Middle to Late Pleistocene M. s. pliocena rather than with the extant representative M. s. sylvanus. Moreover, principal component analysis reveals a better match with M. s. pliocena. However, because no clear-cut diagnostic criteria have been defined to differentiate these European fossil subspecies, we attribute the Marathousa 1 specimen to M. s. cf. pliocena, in agreement with the chronology of the locality. Previously known only from the Early Pleistocene of Greece by some isolated teeth, this is the first record of Macaca in the Middle Pleistocene of the country and one of very few in the eastern sector of the peri-Mediterranean region. We discuss the presence of macaques in the paleolake environment of Marathousa 1, as well as their predation risks from both carnivores and hominins present at the locality.
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Lithics and cut-marked mammal bones, excavated from the paleo-lake Marathousa 1 (MAR-1) sediments in the Megalopolis Basin, southern Greece, indicate traces of hominin activity occurring along a paleo-shoreline ca. 444,000 years (444 ka)... more
Lithics and cut-marked mammal bones, excavated from the paleo-lake Marathousa 1 (MAR-1) sediments in the Megalopolis Basin, southern Greece, indicate traces of hominin activity occurring along a paleo-shoreline ca. 444,000 years (444 ka) ago. However, the local environment and climatic conditions promoting hominin activity in the area during the MIS12 glacial remain largely unknown. In order to reconstruct the paleo-environment including paleo-lake levels and governing paleo-climatic factors on a high temporal resolution, we
analyzed a 6-meter-long sediment sequence from the archeological site MAR-1 and a Bayesian age model was computed for a better age constrain of the different sedimentary units. A multiproxy approach was applied using ostracods, sponge spicules, diatoms, grain sizes, total organic carbon, total inorganic carbon and conventional X-ray fluorescence analysis. The results from the site represent a protected region surrounded by high mountains under the constant influence of water, either as a shallow partly anoxic water body surrounded by reed belts (>463–457 ka, <434–427 ka), a riverine-lake deltaic system
(∼457–448 ka), a floodplain (∼448–444 ka) or a seasonal freshwater pond (∼444–436 ka). The local changes of water levels resemble large trends and rhythms of regional records from the Mediterranean and appear to directly respond to sea surface temperature (SST)
changes of the North Atlantic. In particular, when the SSTs are high, more moisture reaches the study area and vice versa. Additional water reaches MAR-1 through melting of the surrounding glaciers after brief warm phases during MIS12 in the Mediterranean realm, which leads to the formation of smaller fresh water ponds, where also the horizon of the excavated remains is placed. Such ponds, rich in ostracods and other microorganisms, provided mammals and humans valuable resources, such as potable water, a wide range of plant species and hunting opportunities. These deposits therefore bear a high archeological potential. The results from our study suggest that the Megalopolis Basin could have served as a refugium for hominins and other organisms due to its capacity to retain freshwater bodies during glacial and interglacial periods.
analyzed a 6-meter-long sediment sequence from the archeological site MAR-1 and a Bayesian age model was computed for a better age constrain of the different sedimentary units. A multiproxy approach was applied using ostracods, sponge spicules, diatoms, grain sizes, total organic carbon, total inorganic carbon and conventional X-ray fluorescence analysis. The results from the site represent a protected region surrounded by high mountains under the constant influence of water, either as a shallow partly anoxic water body surrounded by reed belts (>463–457 ka, <434–427 ka), a riverine-lake deltaic system
(∼457–448 ka), a floodplain (∼448–444 ka) or a seasonal freshwater pond (∼444–436 ka). The local changes of water levels resemble large trends and rhythms of regional records from the Mediterranean and appear to directly respond to sea surface temperature (SST)
changes of the North Atlantic. In particular, when the SSTs are high, more moisture reaches the study area and vice versa. Additional water reaches MAR-1 through melting of the surrounding glaciers after brief warm phases during MIS12 in the Mediterranean realm, which leads to the formation of smaller fresh water ponds, where also the horizon of the excavated remains is placed. Such ponds, rich in ostracods and other microorganisms, provided mammals and humans valuable resources, such as potable water, a wide range of plant species and hunting opportunities. These deposits therefore bear a high archeological potential. The results from our study suggest that the Megalopolis Basin could have served as a refugium for hominins and other organisms due to its capacity to retain freshwater bodies during glacial and interglacial periods.
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In recent years, a significant number of Pleistocene localities with evidence of proboscidean exploitation by humans has been discovered, substantially enriching our knowledge on Homo subsistence strategies and megafauna acquisition. In... more
In recent years, a significant number of Pleistocene localities with evidence of proboscidean exploitation by humans has been discovered, substantially enriching our knowledge on Homo subsistence strategies and megafauna acquisition. In this study, we provide a synthesis of the evidence for Proboscidea-Homo interactions in Early and Middle Pleistocene open-air sites of western Eurasia with direct (e.g., presence of cut marks, proboscidean bone artifacts, fractures for marrow extraction) and indirect (e.g., association and refitting of lithic artifacts, use-wear analysis) evidence of exploitation. Sex and ontogenetic age of butchered proboscideans are recorded, in order to assess possible human preferences. Furthermore, we investigate the role of large carnivores focusing on important renewals in the carnivore guilds, and their significance in terms of carrion availability for scavenging and human-carnivore competition for access to food resources. By applying an ecomorphological/behavioral approach, we examine the large carnivore community structure and dynamics, with emphasis in the hunting strategies of large predators. Additionally, we aim to infer their possible role in the changes of early human subsistence strategies focusing on proboscidean acquisition, and to explore the role of humans within the predatory guild. The ecological adaptations of the two common Middle Pleistocene proboscideans in Europe, the European straight-tusked elephant Palaeoloxodon antiquus and the steppe mammoth Mammuthus trogontherii, are also evaluated. Finally, we discuss various aspects of the Homo bio-cultural evolution during the period under study, including developments in material culture and relevant inferences about human social behavior.
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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... more
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 correlations and schemes. In this article, we report the first detailed chronological scheme for the late Villafranchian of southeastern Europe through a comprehensive and multidisciplinary dating approach (biochronology, magnetostratigraphy, and cosmogenic radionuclides) of the recently discovered Lower Pleistocene
vertebrate site Tsiotra Vryssi (TSR) in the Mygdonia Basin, Greece. Results: The minimum burial ages (1.88 ± 0.16 Ma, 2.10 ± 0.18 Ma, and 1.98 ± 0.18 Ma) provided by the method of cosmogenic radionuclides indicate that the normal magnetic polarity identified below the fossiliferous layer correlates to the Olduvai subchron (1.95–1.78 Ma; C2n). Therefore, an age younger than 1.78 Ma is indicated for the fossiliferous layer, which was deposited during reverse polarity chron C1r. These results are in agreement with the biochronological data, which further point to an upper age limit at ~1.5 Ma. Overall, an age between 1.78 and ~1.5 Ma (i.e., within the first part of the late Villafranchian) is proposed for the TSR fauna. Conclusions: Our results not only provide age constraints for the local mammal faunal succession, thus allowing for a better understanding of faunal changes within the same sedimentary basin, but also contribute to improving correlations on a broader scale, leading to more accurate biogeographic, palaeoecological, and taphonomic interpretations.
vertebrate site Tsiotra Vryssi (TSR) in the Mygdonia Basin, Greece. Results: The minimum burial ages (1.88 ± 0.16 Ma, 2.10 ± 0.18 Ma, and 1.98 ± 0.18 Ma) provided by the method of cosmogenic radionuclides indicate that the normal magnetic polarity identified below the fossiliferous layer correlates to the Olduvai subchron (1.95–1.78 Ma; C2n). Therefore, an age younger than 1.78 Ma is indicated for the fossiliferous layer, which was deposited during reverse polarity chron C1r. These results are in agreement with the biochronological data, which further point to an upper age limit at ~1.5 Ma. Overall, an age between 1.78 and ~1.5 Ma (i.e., within the first part of the late Villafranchian) is proposed for the TSR fauna. Conclusions: Our results not only provide age constraints for the local mammal faunal succession, thus allowing for a better understanding of faunal changes within the same sedimentary basin, but also contribute to improving correlations on a broader scale, leading to more accurate biogeographic, palaeoecological, and taphonomic interpretations.
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Recent excavations at the Middle Pleistocene open-air site of Marathousa 1 have unearthed in one of the two investigated areas (Area A) a partial skeleton of a single individual of Palaeoloxodon antiquus and other faunal remains in... more
Recent excavations at the Middle Pleistocene open-air site of Marathousa 1 have unearthed in one of the two investigated areas (Area A) a partial skeleton of a single individual of Palaeoloxodon antiquus and other faunal remains in spatial and stratigraphic association with lithic artefacts. In Area B, a much higher number of lithic artefacts was collected, spatially and stratigraphically associated also with faunal remains. The two areas are stratigraphically correlated, the main fossiliferous layers representing an en mass depositional process in a lake margin context. Evidence of butchering (cut-marks) has been identified on bones of the elephant skeleton, as well on elephant and other mammal bones from Area B. However, due to the secondary deposition of the main find-bearing units, it is of primary importance to evaluate the degree and reliability of the spatial association of the lithic artefacts with the faunal remains. Indeed, spatial association does not necessarily imply causation, since natural syn-and post-depositional processes may equally produce spatial association. Assessing the degree and extent of post-depositional reworking processes is crucial to fully comprehend the archaeological record, and therefore to reliably interpret past human behaviours. The present study uses a comprehensive set of spatial statistics in order to disentangle the depositional processes behind the distribution of the archaeological and palaeontological record at Marathousa 1. Preliminary results of our analyses suggest that a high-energy erosional process, attributed to a hyperconcentrated flow deposited at the margin of a swamp, reworked an autochthonous, exposed or slightly buried, scatter of lithic artefacts and faunal remains. Minor reworking and substantial spatial association of the lithic and faunal assemblages support the current interpretation of Marathousa 1 as a butchering site.
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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... more
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 Mammuthus meridionalis (Nesti, 1825). Yet, the dental features indicate an evolutionary stage somewhat more derived than the Upper Valdarno mammoth (beginning of late Villafranchian) in terms of the higher number of plates in the DP3 and the marginally higher hypsodonty index in the M3. This is also revealed by the application of a Principal Component Analysis, where several dental features are combined. Moreover, the relatively deep mandibular corpus is a derived feature. In these aspects, the Apollonia-1 sample fits better with corresponding specimens from European localities correlated to the second part of the late Villafranchian and the Epivillafranchian. From this period, two subspecies have been proposed from Western Europe: M. m. vestinus (Azzaroli in Ambrosetti, Azzaroli, Bonadonna & Follieri, 1972) from Italian localities correlated to the Farneta Faunal Unit and M. m. depereti Coppens & Beden, 1982 from Saint-Prest (France). We propose that M. m. vestinus can be regarded a valid subspecies of the southern mammoth and that M. m. depereti is possibly a junior synonym. The results of this study are in agreement with the existing biochronological correlation of the Apollonia-1 fauna and further support the potential biostratigraphic significance of M. meridionalis within the Early Pleistocene, albeit conclusions based on limited/fragmentary sample should always be taken with caution.
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In the present article, we study the proboscidean remains from three upper Miocene localities of Northern Greece: Thermopigi (Serres), Neokaisareia (Pieria) and Platania (Drama). The material from the Turolian locality of Thermopigi... more
In the present article, we study the proboscidean remains from three upper Miocene localities of Northern Greece: Thermopigi (Serres), Neokaisareia (Pieria) and Platania (Drama). The material from the Turolian locality of Thermopigi includes only postcranial specimens. The morphological features of the scapula indicate the presence of the deinotheriid Deinotherium sp., whereas the rest of the specimens are morphologically distinct from Deinotherium and can be referred to Elephantimorpha indet. The material from Neokaisareia consists of a partial skeleton of a single individual and is attributed to the mammutid Mammut sp. (M. obliquelophus?). This taxon is known in Greece from the early–middle Turolian. The Platania proboscidean belongs to the tetralophodont amebelodontid Konobelodon cf. atticus. The genus Konobelodon was already present during the Vallesian of the wider area, but the lower tusk of the Platania shovel-tusker presents some morphological and metrical differences from the Vallesian representative, yet it has also smaller dimensions in its deciduous dentition than the morphologically similar Turolian specimens. The type locality of K. atticus is Pikermi (Attica, Greece), correlated to the middle Turolian, but the known biostratigraphic range of this species covers the entire Turolian. Platania is possibly correlated close to the Vallesian/Turolian boundary and the possible record of this species could document one of its earliest occurrences.
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In this article, we report on two proboscidean upper tusks from a new locality in Iran, Kivi-1 (KIV). The tusks are oval to circular in cross-section, lack an enamel band, are strongly curved and double-twisted, and turn inwards at the... more
In this article, we report on two proboscidean upper tusks from a new locality in Iran, Kivi-1 (KIV). The tusks are oval to circular in cross-section, lack an enamel band, are strongly curved and double-twisted, and turn inwards at the tip. This morphology permits the attribution to Choerolophodon, an elephantimorph proboscidean well distributed during the Miocene in the Old World and already known from the late Miocene of Iran. Although the known upper tusks of Choerolophodon are few, the current data indicate that the curvature of the upper tusks may have biostratigraphic significance: in earlier and more primitive representatives from the early and middle Miocene of Eurasia, the curvature is less pronounced, whereas in the later and more derived ones from the late Miocene, it is strong. Based on this feature, we propose that the horizons from which these tusks derive date to the upper Miocene. During this epoch, two choerolophodont species were present in the Greco-Iranian palaeobiogeographic province, C. anatolicus and C. pentelici, the latter one well represented in the nearby Maragheh faunas. Because more diagnostic specimens are so far lacking, we attribute the KIV specimens to Choerolophodon sp. indet. “derived morph”. Based on the presence of either C. anatolicus or C. pentelici in KIV, a relatively open palaeoenvironment is assumed, in agreement with previous palaeoecological reconstructions from the wider region during this period.
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In this article, we present the proboscideans from the late Miocene (Turolian) locality Chomateri, Greece, near the classical locality Pikermi. The material consists of juvenile teeth, whose morphological features, such as the dislocation... more
In this article, we present the proboscideans from the late Miocene (Turolian) locality Chomateri, Greece, near the classical locality Pikermi. The material consists of juvenile teeth, whose morphological features, such as the dislocation of the half-loph(id)s and the resultant alternate arrangement of the successive loph(id)s (anancoidy), permit assignment to the tetralophodont gomphothere Anancus. However, the anancoidy is rather weak and the occlusal morphology simple, both regarded as primitive features within anancines. Reexamination of the late Miocene anancines from Europe reveals that they all share primitive molar features (weak anancoidy, simple morphology, thick enamel) with the material from Hohenwarth, Austria, showing further primitive skull features, such as the longer mandibular symphysis compared with other anancines. The proper name to refer to the late Miocene anancines from Europe is Anancus lehmanni (Gaziry, 1997), with type locality Dorn Dürkheim 1 (Turolian; Germany). The presence of Anancus in Chomateri constitutes the first late Miocene record of the genus in Greece, and the first faunal element that clearly indicates that Chomateri postdates the classical Pikermi. Finally, we discuss the biostratigraphy and the biogeography of the late Miocene anancines of the Old World. Anancus originated possibly at ∼9.0–8.5 Ma in Asia and entered Europe during the second half of the Turolian, at ∼7.2 Ma (Tortonian-Messinian boundary). The arrival of Anancus in Europe coincides with a faunal turnover in both the eastern and western sectors of the European Mediterranean region, and in the southern Balkans in particular, with the decline of the ‘Pikermian’ large-mammal fauna.
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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... more
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 nineteenth century and the beginning of the twentieth century. The specimens belong to juvenile individuals of deinotheres, choerolophodonts and amebelodonts. The deinothere material is attributed to the last European huge-sized deinothere, Deinotherium proavum. The described skull from Samos is currently the most complete specimen of all known Miocene juvenile deinotheres from Eurasia and Africa. The majority of the Samos choerolophodont specimens belong to the advanced morph of Choerolophodon pentelici, whereas one shows more archaic features and belongs to the primitive evolutionary stage of this species. This more primitive morph could originate from the lower fossiliferous horizons of Samos, which are dated to the early Turolian. The third proboscidean is attributed to the tetralophodont shovel-tusker Konobelodon atticus, a rare taxon in the Samos fauna. Together with the previously described zygodont Mammut from Samos, these four proboscideans are typical of the Turolian proboscidean fauna of southeastern Europe. We discuss the biostratigraphy of the Samos proboscideans with the aim of unraveling some aspects of the chronological range of the late Miocene proboscideans, focusing in particular on the Southern Balkans and Turkey.
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A B S T R A C T The technological systems and subsistence strategies of Middle Pleistocene hominins in SouthEast Europe are insufficiently understood due to the scarcity of well-preserved, excavated assemblages. In this paper, we present... more
A B S T R A C T The technological systems and subsistence strategies of Middle Pleistocene hominins in SouthEast Europe are insufficiently understood due to the scarcity of well-preserved, excavated assemblages. In this paper, we present first results from the study of the lithic and bone artifacts unearthed at the Lower Palaeolithic site Marathousa 1 (MAR-1), Megalopolis, Greece. The context of the site represents a depositional environment close to a lake-shore, where rapid burial in a very fine-grained matrix ensured extraordinary conditions for preservation. Lithic artifacts occur in spatial and stratigraphic association with remains of the elephant Palaeoloxodon antiquus as well as other mammals. Bones, including those of elephants, show clear anthropogenic flaking scars, cut-marks and fracture patterns indicating deliberate breakage and modification by early humans. The MAR-1 lithic assemblage is composed of small-sized debitage, retouched tools, a few small and exhausted cores, as well as a large number of debris and retouch products, such as chips and resharpening flakes. Currently, there are no indications of Acheulean bifacial debitage, large cutting tools are missing, and a key aspect of the material refers to its 'microlithic' character. The scarcity of cores and primary flakes indicates a fragmented reduction sequence and complex discard patterns that require further investigation. On the basis of the ongoing analysis of lithic material from three field seasons, we discuss aspects of assemblage composition and the role of raw material types, the main technological and typological traits of the industry, as well as the potential contribution of the MAR-1 assemblage in broader discussions about Middle Pleistocene lithic techno-complexes and subsistence strategies in Eurasia. Finally, we briefly present a small sample of bone artifacts, which suggest that hominin exploitation of the animal carcasses was not restricted to marrow extraction and bone processing for nutritional needs, but included also the knapping of bones, potentially with the aim of using the knapped products as tools. The MAR-1 archaeological record compares well with other important Eurasian sites yielding 'small tool as-semblages', such as Ficoncella, La Polledrara, Bilzingsleben, Schöningen and Vértesszőlős, some of which, like MAR-1, have provided evidence of elephant or other mega-fauna exploitation.
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Marathousa 1 is a Lower Palaeolithic open-air site located in the Megalopolis basin, an area in Southern Greece known for its fossiliferous sediments. Mining activities in the basin uncovered a thick sequence of Middle Pleistocene... more
Marathousa 1 is a Lower Palaeolithic open-air site located in the Megalopolis basin, an area in Southern Greece known for its fossiliferous sediments. Mining activities in the basin uncovered a thick sequence of Middle Pleistocene lacustrine deposits representing the environment of a palaeolake. Marathousa 1 was discovered in 2013 during a targeted palaeoanthropological survey and excavated subsequently by an interdisciplinary team from the Ephoreia of Palaeoanthropology–Speleology of Greece and the University of Tübingen, Germany. This article presents results from the ongoing investigation and reviews the state of knowledge about the site. Systematic excavations during five field seasons have exposed a total of 72 m 2 and revealed a partial skeleton of the elephant Palaeoloxodon antiquus and remains of other large mammals in spatial and stratigraphic association with a " small tool " lithic assemblage. Faunal and taphonomic studies indicate the presence of cut-marks and percussion damage on elephant and other large mammal bones. The study of site formation processes, together with taphonomic and geostatistic spatial analyses confirm the association of fossil and hominin activity remains and the stratigraphic integrity of the site. Radiometric dating, geological and biostratigraphical evidence suggest that hominin activity at the site occurred between 0.5 and 0.4 Ma. Marathousa 1 is the oldest currently known archaeological site in Greece and the only Lower Palaeolithic butchering site in the Southern Balkans. It is also a key site for documenting high resolution palaeoclimatic, palaeoenvironmental and cultural records of a geographical area that potentially acted as a refugium during the successive waves of hominin colonization of Europe.
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The technological systems and subsistence strategies of Middle Pleistocene hominins in SouthEast Europe are insufficiently understood due to the scarcity of well-preserved, excavated assemblages. In this paper, we present first results... more
The technological systems and subsistence strategies of Middle Pleistocene hominins in SouthEast Europe are insufficiently understood due to the scarcity of well-preserved, excavated assemblages. In this paper, we present first results from the study of the lithic and bone artifacts unearthed at the Lower Palaeolithic site Marathousa 1 (MAR-1), Megalopolis, Greece. The context of the site represents a depositional environment close to a lake-shore, where rapid burial in a very fine-grained matrix ensured extraordinary conditions for preservation. Lithic artifacts occur in spatial and stratigraphic association with remains of the elephant Palaeoloxodon antiquus as well as other mammals. Bones, including those of elephants, show clear anthropogenic flaking scars, cut-marks and fracture patterns indicating deliberate breakage and modification by early humans. The MAR-1 lithic assemblage is composed of small-sized debitage, retouched tools, a few small and exhausted cores, as well as a large number of debris and retouch products, such as chips and resharpening flakes. Currently, there are no indications of Acheulean bifacial debitage, large cutting tools are missing, and a key aspect of the material refers to its 'microlithic' character. The scarcity of cores and primary flakes indicates a fragmented reduction sequence and complex discard patterns that require further investigation. On the basis of the ongoing analysis of lithic material from three field seasons, we discuss aspects of assemblage composition and the role of raw material types, the main technological and typological traits of the industry, as well as the potential contribution of the MAR-1 assemblage in broader discussions about Middle Pleistocene lithic techno-complexes and subsistence strategies in Eurasia. Finally, we briefly present a small sample of bone artifacts, which suggest that hominin exploitation of the animal carcasses was not restricted to marrow extraction and bone processing for nutritional needs, but included also the knapping of bones, potentially with the aim of using the knapped products as tools. The MAR-1 archaeological record compares well with other important Eurasian sites yielding 'small tool as-semblages', such as Ficoncella, La Polledrara, Bilzingsleben, Schöningen and Vértesszőlős, some of which, like MAR-1, have provided evidence of elephant or other mega-fauna exploitation.
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In this article, we present the first results on the large mammal fauna from the new open-air Lower Palaeolithic locality Marathousa 1 (MAR-1) (Megalopolis Basin, Peloponnesus, Greece). MAR-1 belongs to the Marathousa Member of the... more
In this article, we present the first results on the large mammal fauna from the new open-air Lower Palaeolithic locality Marathousa 1 (MAR-1) (Megalopolis Basin, Peloponnesus, Greece). MAR-1 belongs to the Marathousa Member of the Choremi Formation and its large mammal faunal list (collection 2013-2016) includes the castorid Castor fiber, the mustelids Mustela sp. and Lutra simplicidens, the felid Felis sp., the canids Vulpes sp. and Canis sp., the elephantid Palaeoloxodon antiquus, the hippopotamid Hippopotamus antiquus, the bovid Bison sp., and the cervids Dama sp. and Cervus elaphus. This faunal association is common in the Galerian (Middle Pleistocene) mammal communities of Europe (ca. 0.9-0.4 Ma). The MAR-1 fauna is consistent with a temperate climate and is indicative of a landscape with substantial woodland components with more open areas, close to permanent and large freshwater bodies. Of particular interest are an elephant cranium and numerous postcranial elements, which were found in close anatomical association and are attributed to a single individual of the straight-tusked elephant Palaeoloxodon antiquus. The skeleton belonged to a male individual in its late adulthood close to or in its sixties, with live skeletal height around 3.7 m at the shoulder and body mass around 9.0 tonnes. The good state of preservation of the MAR-1 bones allows the identification of taphonomic modifications. Cut marks on the elephant skeleton, and on other elephant and mammal bones, indicate human exploitation by means of butchering activities, in accordance with the traits of the lithic assemblage and its spatial association with the bones. Carnivore activity is also recorded on some elephant and cervid bones. Marathousa 1 is among the oldest elephant butchering sites in Europe and the only one known in Southeastern Europe.
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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,... more
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 Tsiotra Vryssi (TSR), dated to the late Villafranchian (1.8-1.2 Ma). The specimens are described and compared with ursids from various European localities; their morphological characters and dimensions place it with Ursus etruscus Cuvier, 1823, while the dental features are most similar to the specimens from Pietrafitta (Italy). The TSR cranium was CT-scanned in order to create a 3D virtual model and study its internal anatomy. Its observed endocranial traits exhibit primitive conditions, confirming the basal position of U. etruscus among the Pleistocene European ursids. Most of these features are shared with U. arctos and clearly differ from U. deningeri and U. spelaeus. Finally, we provide a revision of the known Greek material of U. etruscus and the biostratigraphic distribution of the MioceneePleistocene ursids of Greece. The earliest secure appearance of Ursus etruscus in Greece is found in the locality Dafnero 1 (Western Macedonia), dated to the end of the middle Villafranchian at ~2.0 Ma, whereas its probable last occurrence is known from the locality Apollonia 1 of the Mygdonia Basin, dated to the latest Villafranchian at ~1.2 Ma.
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In this article, we present the proboscidean assemblage from the upper Miocene vertebrate locality Nikiti 2 (Chalkidiki Peninsula, Macedonia, Greece). Based on the dental remains, the material belongs exclusively to the genus... more
In this article, we present the proboscidean assemblage from the upper Miocene vertebrate locality Nikiti 2 (Chalkidiki Peninsula, Macedonia, Greece). Based on the dental remains, the material belongs exclusively to the genus Choerolophodon. The morphology of the dp3/DP3 and the dimensions of the deciduous dentition clearly distinguish the Nikiti 2 choerolophodont from the early Vallesian Choerolophodon anatolicus and permit its assignment to C. pentelici. C. pentelici is represented in SE Europe–SW Asia by a primitive morph, known from the late Vallesian–earliest Turolian, and an advanced morph Turolian in age. The morphometric features of the Nikiti 2 specimens and their comparison with the choerolophodont material from upper Miocene localities of the wider region indicate that the Nikiti 2 C. pentelici belongs to the advanced morph of the species, suggestive for a Turolian age for the locality. These results agree with the rest of the fauna, which is correlated to the early Turolian (MN 11).
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Presented here are the deinotheriid dental and postcranial remains from the Late Miocene localities Pikermi and Halmyropotamos (Greece). The study and comparison of the available juvenile dental material from Pikermi with other relevant... more
Presented here are the deinotheriid dental and postcranial remains from the Late Miocene localities Pikermi and Halmyropotamos (Greece). The study and comparison of the available juvenile dental material from Pikermi with other relevant specimens from Europe showed that it belongs to the huge-sized Deinotherium proavum (¼ D. gigantissimum). Additionally, several postcranial specimens from Pikermi, as well as from Halmyropotamos, present deinotheriid features, which distinguish them from elephantoids, and permit their attribution also to D. proavum. This species is known from the Turolian and represents the terminal evolutionary stage of the European deinotheres. Its presence in Pikermi and Halmyr-opotamos is compatible with the middle Turolian (MN 12) age of the localities. Additionally, the tax-onomy of European deinotheres is discussed, as well as the biostratigraphical and geographical distribution of D. proavum.
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We here report the first results from a systematic research project in Mani (Southern Greece), which includes survey and test excavations. Forty-six caves, rockshelters and open-air sites in lowland settings were surveyed.... more
We here report the first results from a systematic research project in Mani (Southern Greece), which includes
survey and test excavations. Forty-six caves, rockshelters and open-air sites in lowland settings were
surveyed. Geomorphological data were collected in order to assess how geological processes affect the
preservation of sites and bias site distribution patterns. Artifacts manufactured from non-local rock indicate
potential raw material transfers and suggest links among the different regions of Mani, related to mobility
patterns. Our research in the Mani has nearly doubled the number of known Middle Palaeolithic sites from
the region and confirmed that the peninsula has the strongest ‘Neanderthal signal’ identified to date in
Greece. Almost all sites are located at coastal areas. Despite the influence of Pleistocene landscape
dynamics, this distribution emerges as a persistent pattern, perhaps indicating a preference for coastal
locations. The Neanderthal occupation of Mani can illuminate important aspects of Middle Palaeolithic
adaptation in one of the southernmost coastal regions of Europe.
survey and test excavations. Forty-six caves, rockshelters and open-air sites in lowland settings were
surveyed. Geomorphological data were collected in order to assess how geological processes affect the
preservation of sites and bias site distribution patterns. Artifacts manufactured from non-local rock indicate
potential raw material transfers and suggest links among the different regions of Mani, related to mobility
patterns. Our research in the Mani has nearly doubled the number of known Middle Palaeolithic sites from
the region and confirmed that the peninsula has the strongest ‘Neanderthal signal’ identified to date in
Greece. Almost all sites are located at coastal areas. Despite the influence of Pleistocene landscape
dynamics, this distribution emerges as a persistent pattern, perhaps indicating a preference for coastal
locations. The Neanderthal occupation of Mani can illuminate important aspects of Middle Palaeolithic
adaptation in one of the southernmost coastal regions of Europe.
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As one of the oldest known Eurasian fossil vertebrate localities, Samos late Miocene fauna attracted the interest of specialists by its richness and overall importance. Nevertheless, crucial taxonomical questions and chronological... more
As one of the oldest known Eurasian fossil vertebrate localities, Samos late Miocene fauna attracted the interest of specialists by its richness and
overall importance. Nevertheless, crucial taxonomical questions and chronological problems obscured its value. The detailed study of the local
stratigraphy, the collection of new fossil material and its study, the revision of the old collections and the updated magneto-chronology of the
fossiliferous deposits permited to re-discuss most of the problems in a special volume edited in 2009 by Koufos and Nagel and to provide a clearer
and more precise idea about the Samos fauna and its age. A synopsis of this work is given here. The systematic study of the new collection (1200
identified specimens) allows the determination of 42 species from three fossil horizons, ranging from the upper part of early Turolian (MN11) to the
end of middle Turolian (MN12). Taxonomic novelties are the presence of the carnivore genus Protictitherium found for the first time in Samos, the
establishment of the new name Skoufotragus for Pachytragus Schlosser with the new species Skoufotragus zemalisorum, and the amended
morphology of Pseudomeriones and Urmiatherium. Additionally six Hipparion and four Gazella species were recognized and a better
morphometric distinction between Samotherium boissieri and Samotherium major was performed. This study also improved the correlation
of the old fossiliferous sites with the new ones and with the local stratigraphy of the Mytilinii Basin, while precise ages have been obtained for the
mammal localities. The new data together with the old collections indicate the presence in Samos of four chronologically successive mammal
assemblages reflecting a ‘‘four stages-of-evolution’’ scheme. The Turolian palaeoenvironment of Samos is determined as an open bushland with
thick grassy-floor of C3 graminoids with possible increase of the open and dry character from the beginning to the end of Middle Turolian. The
Samos mammal faunas are palaeobiogeographically closer to the Asian ones than to those from the Greek mainland.
overall importance. Nevertheless, crucial taxonomical questions and chronological problems obscured its value. The detailed study of the local
stratigraphy, the collection of new fossil material and its study, the revision of the old collections and the updated magneto-chronology of the
fossiliferous deposits permited to re-discuss most of the problems in a special volume edited in 2009 by Koufos and Nagel and to provide a clearer
and more precise idea about the Samos fauna and its age. A synopsis of this work is given here. The systematic study of the new collection (1200
identified specimens) allows the determination of 42 species from three fossil horizons, ranging from the upper part of early Turolian (MN11) to the
end of middle Turolian (MN12). Taxonomic novelties are the presence of the carnivore genus Protictitherium found for the first time in Samos, the
establishment of the new name Skoufotragus for Pachytragus Schlosser with the new species Skoufotragus zemalisorum, and the amended
morphology of Pseudomeriones and Urmiatherium. Additionally six Hipparion and four Gazella species were recognized and a better
morphometric distinction between Samotherium boissieri and Samotherium major was performed. This study also improved the correlation
of the old fossiliferous sites with the new ones and with the local stratigraphy of the Mytilinii Basin, while precise ages have been obtained for the
mammal localities. The new data together with the old collections indicate the presence in Samos of four chronologically successive mammal
assemblages reflecting a ‘‘four stages-of-evolution’’ scheme. The Turolian palaeoenvironment of Samos is determined as an open bushland with
thick grassy-floor of C3 graminoids with possible increase of the open and dry character from the beginning to the end of Middle Turolian. The
Samos mammal faunas are palaeobiogeographically closer to the Asian ones than to those from the Greek mainland.
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Two new vertebrate fossiliferous localities from the Pleistocene of the Mygdonia Basin (Macedonia, Greece), Tsiotra Vryssi (TSR) and Platanochori-1 (PLN), are presented in this article. TSR belongs to the Gerakarou Formation and its... more
Two new vertebrate fossiliferous localities from the Pleistocene of the Mygdonia Basin (Macedonia, Greece), Tsiotra Vryssi (TSR) and Platanochori-1 (PLN), are presented in this article. TSR belongs to the Gerakarou Formation and its preliminary faunal list includes the corvid Corvus pliocaenus, the hyaenid Pachycrocuta brevirostris, the canid Canis etruscus, the rhinoceros Stephanorhinus sp., two species of Equus, the bovid Bison sp., the cervid Metacervocerus rhenanus and the giraffid Palaeotragus sp. PLN belongs to the Platanochori Formation (overlying Gerakarou Formation) and its fauna includes the following taxa: Stephanorhinus hundsheimensis, Equus apolloniensis, Bison sp. and Pontoceros ambiguus. The preliminary comparison of these faunal assemblages with the already known localities from the Mygdonia Basin and the wider region, combined with the geological data, suggest a Late Villafranchian age for TSR, chronologically intermediate between the localities Gerakarou-1 and Apollonia-1, while the fauna from PLN is indicative of a Latest Villafranchian age, similar to that of Apollonia-1. These new localities will enhance our understanding of the crucial time period of the earliest dispersal of hominins into Europe.
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The Miocene elephantoid Choerolophodon from SE Europe-SW Asia is studied, based on the Greek localities of Thymiana, Axios Valley, Pikermi, Samos and Nikiti-2. Although this genus is well documented in Greece, there is no recent taxonomic... more
The Miocene elephantoid Choerolophodon from SE Europe-SW Asia is studied, based on the Greek localities of Thymiana, Axios Valley, Pikermi, Samos and Nikiti-2. Although this genus is well documented in Greece, there is no recent taxonomic study and its biostratigraphy and palaeoecology had never been examined in detail. Three choerolophodont species are recognized in SE Europe–SW Asia: the Middle Miocene C. chioticus (late Orleanian, MN 5), and the Late Miocene C. anatolicus (early Vallesian, MN 9) and C. pentelici (late Vallesian–Turolian, MN 10–MN 13). The latter species is divided into a primitive morph, dated to the late Vallesian and possibly earliest Turolian, and an advanced morph from the Turolian. At the end of the Miocene Choerolophodon disappeared. With respect to the geographical distribution of Choerolophodon, the genus is very well documented in SE Europe–SW Asia but it is unknown from Central, Western and Northern Europe, a fact which is attributed to ecological factors. The phylogenetic relationships of all known choerolophodont species from the Miocene of the Old World are examined using a cladistic analysis, and their proposed biogeography is discussed. Finally, the palaeoecology of Late Miocene Choerolophodon from Northern Greece is studied through a dental microwear analysis, revealing that Choerolophodon foraged mainly on grasses, a type of vegetation widespread in open environments. These palaeoecological results are in agreement with previous environmental reconstructions for the Late Miocene of Greece.
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Presented here are the tetralophodont proboscidean remains from the late Miocene locality Pikermi (Attica, Greece). The study of the Pikermi tetralophodons and their comparison with other relevant specimens revealed that the late Miocene... more
Presented here are the tetralophodont proboscidean remains from the late Miocene locality Pikermi (Attica, Greece). The study of the Pikermi tetralophodons and their comparison with other relevant specimens revealed that the late Miocene species ‘Tetralophodon’ atticus and ‘Mastodon’ grandincisivus are synonymous. The morphological characters of the Pikermi specimens, as well as of those attributed to ‘Mastodon’ grandincisivus, are consistent with the North American shovel-tusker Amebelodon (Konobelodon). However, generic rank revision of Konobelodon is deemed appropriate. The observed differences between the type species Konobelodon britti and the Pikermi tetralophodon indicate that the latter belongs to a distinct species, Konobelodon atticus. This species encompasses the Turolian tetralophodont shovel-tuskers from eastern Europe–western Asia. The paleogeographic distribution and biostratigraphy of Konobelodon atticus are discussed, as well as the evolution and the possible migrational scenarios of Konobelodon.
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The abundant Late Miocene proboscidean remains of Greece have never been studied in detail and compared with those of Eurasia in order to determine their taxonomy and their biostratigraphical and palaeoecological significance. The first... more
The abundant Late Miocene proboscidean remains of Greece have never been studied in detail and compared with those of Eurasia in order to determine their taxonomy and their biostratigraphical and palaeoecological significance. The first results of such study are given in this article. During the past decades, several new proboscidean specimens have been added to the old collections, significantly enriching the available material. The Axios Valley (Macedonia, Greece) proboscidean fossils belong mainly to two species of Choerolophodon: C. anatolicus of early Vallesian age and C. pentelici of late Vallesian–Turolian age. Deinotherium giganteum is rare and recognized only in the late Vallesian locality Ravin de la Pluie of Axios Valley. A zygodont form has also been identified in the Turolian of Axios Valley, attributed to “Mammut” sp. The Late Miocene localities of Nikiti (Macedonia, Greece) revealed several remains of C. pentelici, which are similar to the Turolian ones of Axios Valley. The Samos proboscidean collection includes C. pentelici, “Tetralophodon” atticus, “Mammut” sp. and Deinotherium gigantissimum. The taxonomy of the Late Miocene peri-Mediterranean Choerolophodon is given, and the biostratigraphy and palaeoecology of the Greek Late Miocene proboscideans are discussed. Die obermiozänen Proboscidier aus Griechenland, obwohl sie ziemlich verbreitet sind, wurden niemals im Detail untersucht und mit anderen aus Eurasien verglichen, um ihre Taxonomie, Biostratigraphie und Paläoökologie zu bestimmen. Die ersten Ergebnisse dieser Studie sind in diesem Artikel präsentiert. In den letzten Jahrzehnten kamen viele neue Proboscidier-Funde hinzu und bereichern erheblich das Material der alten Sammlungen. Der Großteil der Proboscidea-Fauna aus dem Axios Tal (Mazedonien, Griechenland) wurde der Gattung Choerolophodon mit zwei Arten zugewiesen: C. anatolicus aus dem unteren Vallesium und C. pentelici aus dem oberen Vallesium-Turolium. Die Art Deinotherium giganteum ist selten und wurde nur in der oberen Vallesium Fundstelle Ravin de la Pluie des Axios Tals nachgewiesen. Eine zygodonte Form ist aus dem Turolium des Axios Tals bekannt und wurde der Gattung “Mammut” zugewiesen. Aus den obermiozänen Fundstellen Nikiti (Mazedonien, Griechenland) stammen zahlreiche Reste von C. pentelici, die denjenigen aus dem Turolium des Axios Tals ähneln. Die Proboscidier-Fauna aus Samos enthält die Arten C. pentelici, “Tetralophodon” atticus, “Mammut” sp. und Deinotherium gigantissimum. Die Taxonomie der obermiozänen Arten von Choerolophodon um den Mittelmeerraum wird vorgestellt und die Biostratigraphie und Paläoökologie der griechischen Proboscidier aus dem Oberen Miozän werden besprochen.
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A micromammalian fauna recovered about 15 years ago from an argillaceous bed on the south-western coast of Lake Vistonis (Xanthi-Komotini Basin, Thrace, NE Greece) and partially determined in Syrides et al. (Comptes Rendus Acad Sci Paris... more
A micromammalian fauna recovered about 15 years ago from an argillaceous bed on the south-western coast of Lake Vistonis (Xanthi-Komotini Basin, Thrace, NE Greece) and partially determined in Syrides et al. (Comptes Rendus Acad Sci Paris 324 (2):427-433, 1997) has been thoroughly re-examined. The small (number of identified specimens = 112) but rather diverse (at least in terms of rodents) collection of isolated teeth has revealed the presence of Asoriculus gibberodon, Deinsdorfia kerkhoffi, Soricidae indet., Leporidae indet., Prolagus michauxi, Pliopetaurista dehneli, Myomimus maritsensis, Murinae indet., Occitanomys adroveri, Apodemus cf. dominans, Apodemus gorafensis, Rhagapodemus primaevus, Micromys steffensi, Mesocricetus cf. primitivus, Pliospalax sp. and Pseudomeriones cf. rhodius. The composition of the fauna and comparisons with Turolian and Ruscinian mammalian faunas from the east Mediterranean area suggest a latest Turolian-earliest Ruscinian (MN13/14) age. The taxa favouring dry conditions represent 51.3% in the total MNI (20 of 39 individuals), those favouring wet conditions only 23.1% (9 of 39 individuals). In this overall dry land, the presence of a small water body is indicated by the presence of two (or three) soricid species and Micromys, whereas the record of Pliopetaurista dehneli indicates the presence of trees in the area.
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Marathousa 1 (MAR-1), located in the Megalopolis palaeolake basin, southern Greece, preserves an exceptionally wellpreserved archaeological and palaeontological assemblage. Radiometric dating, magnetostratigraphy, and geological and... more
Marathousa 1 (MAR-1), located in the Megalopolis palaeolake basin, southern Greece, preserves an exceptionally wellpreserved archaeological and palaeontological assemblage. Radiometric dating, magnetostratigraphy, and geological and biochronological analyses indicate an age of 500-400 ka, and place the locality within the Marine Isotope Stage (MIS) 12 [1 and references therein], making it the oldest radiometrically dated archaeological locality currently known in Greece. The locality has yielded lithic and bone artefacts in stratigraphic and spatial association with micro-and macro-fauna (e.g., insects, ostracods, molluscs, fishes, amphibians, reptiles, birds, mammals), as well as micro-and macro-flora [1]. Notably, it preserves the skeletal remains of at least two individuals of the European straight-tusked elephant Palaeoloxodon antiquus, some of which preserve evidence of anthropogenic modifications. In addition to P. antiquus, the locality's large mammal faunal assemblage includes the beaver Castor fiber, the weasel Mustela sp., the otter Lutra simplicidens, the wildcat Felis sp., the fox Vulpes sp., the wolf-sized Canis sp., the hippo Hippopotamus antiquus, the large bovid Bison sp., and the deer Dama sp. and Cervus elaphus [2]. During the excavation campaign of 2019, a cercopithecine primate, represented by an almost complete mandible (MAR-1-9B), was discovered. The specimen preserves both rami and corpora, as well as the most of its dentition. Dental metric comparative analysis permits its attribution to the Barbary macaque Macaca sylvanus, a species that was geographically widely distributed in western Eurasia during the Plio-Pleistocene [3]. Three (chrono) subspecies [4] of European macaques from this period are generally recognized: M. s. prisca from the Pliocene, M. s. florentina from the Early Pleistocene, and M. s. pliocena from the Middle until the Late Pleistocene. The dental dimensions of MAR-1-9B fit better within the variation of M. s. florentina and M. s. pliocena rather than with the extant representative M. s. sylvanus. Moreover, a principal component analysis reveals a better match with M. s. pliocena. Since no clear-cut diagnostic criteria have been defined to differentiate these European fossil subspecies, we attribute MAR-1-9B to M. s. cf. pliocena, in agreement also with the chronology of the locality. MAR-1-9B constitutes one of the very few known records of Macaca in the eastern sector of the peri-Mediterranean region and its first occurrence in the Middle Pleistocene of Greece. It therefore extends the chronological range of cercopithecids and documents, for the first time, the coexistence of macaques and hominins in the country. The palaeolake environment at Marathousa 1 would have offered macaques the protection and resources of the nearby woodlands, as well as access to freshwater and a variety of feeding opportunities throughout the year. Although MAR-1-9B does not show any evidence of anthropogenic or carnivore modifications, prey-predator relationships and interactions among hominins, carnivores and macaques at Marathousa 1 are likely: lithic artefacts and bones with anthropogenic modifications recovered at the site are spatially and stratigraphically associated with carnivore remains and carnivore-modified bones of their prey. In Europe, several Middle Pleistocene localities document the coexistence of macaques and hominins [3], including a few which, like Marathousa 1, have also yielded evidence of elephant butchering: e.g., Fontana Ranuccio, La Polledrara (Italy), Bilzingsleben (Germany), Ebbsfleet (United Kingdom) and Ambrona (Spain) [5]. MAR-1-9B marks the first documented such record in South-East Europe.
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The Pleistocene terrestrial ecosystems of Europe were dominated by powerful carnivores, such as large canids (e.g., Canis, Lycaon), hyenas (e.g., Chasmaporthetes, Pliocrocuta, Pachycrocuta, Crocuta) and large felids (e.g., Megantereon,... more
The Pleistocene terrestrial ecosystems of Europe were dominated by powerful carnivores, such as large canids (e.g., Canis, Lycaon), hyenas (e.g., Chasmaporthetes, Pliocrocuta, Pachycrocuta, Crocuta) and large felids (e.g., Megantereon, Homotherium, Panthera), which were the top predators of this epoch, and competed constantly for the acquisition and exploitation of animal (food) resources. Additionally, the arrival of Homo, having a meat-eating behavior, in western Eurasia at the late Early Pleistocene had a profound impact on the carnivore community dynamics.
By employing an ecomorphological/behavioral approach, the present study contributes to the investigation of the community structure, dynamics, and evolution of the Pleistocene large carnivore guilds of Europe. Emphasis is given to important renewals, the ecological niches, the composition of the guilds in terms of dietary preferences and foraging strategies, and to the intraguild competition, including hominin-carnivore interactions, for access to food resources. Combined are four ecomorphological/behavioral parameters (body mass, diet, hunting strategy, sociality) of large carnivores that practice hunting and/or scavenging on large prey. Four chronofaunas are separated: 2.5-2.0 Ma (middle Villafranchian, i.e., before the arrival of Homo), 1.8-1.0 Ma (within the late Villafranchian and Epivillafranchian, i.e., during the first human colonization of western Eurasia), 500-300 ka (late Galerian-early Aurelian, i.e., when human presence is well recorded almost throughout Europe), and 120-10 ka (Late Pleistocene, i.e., comprising the last diversified European large carnivore guild). 3D guild structure diagrams are analyzed and compared, including comparisons with the modern savanna-ecosystem of Serengeti in Tanzania, and the results are discussed and interpreted.
By employing an ecomorphological/behavioral approach, the present study contributes to the investigation of the community structure, dynamics, and evolution of the Pleistocene large carnivore guilds of Europe. Emphasis is given to important renewals, the ecological niches, the composition of the guilds in terms of dietary preferences and foraging strategies, and to the intraguild competition, including hominin-carnivore interactions, for access to food resources. Combined are four ecomorphological/behavioral parameters (body mass, diet, hunting strategy, sociality) of large carnivores that practice hunting and/or scavenging on large prey. Four chronofaunas are separated: 2.5-2.0 Ma (middle Villafranchian, i.e., before the arrival of Homo), 1.8-1.0 Ma (within the late Villafranchian and Epivillafranchian, i.e., during the first human colonization of western Eurasia), 500-300 ka (late Galerian-early Aurelian, i.e., when human presence is well recorded almost throughout Europe), and 120-10 ka (Late Pleistocene, i.e., comprising the last diversified European large carnivore guild). 3D guild structure diagrams are analyzed and compared, including comparisons with the modern savanna-ecosystem of Serengeti in Tanzania, and the results are discussed and interpreted.
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Archaic humans (early Homo) and carnivores inhabited the Early and Middle Pleistocene landscapes of Europe, and shared ecosystems for more than 1 million years. Indeed, many archaeo-palaeontological sites evidence the co-existence of... more
Archaic humans (early Homo) and carnivores inhabited the Early and Middle Pleistocene landscapes of Europe, and shared ecosystems for more than 1 million years. Indeed, many archaeo-palaeontological sites evidence the co-existence of humans and carnivores, and demonstrate a certain degree of human-carnivore competition for acquisition and exploitation of animal (meat/bone) resources. We investigate here the role of large carnivores in the evolution of hominin subsistence strategies during the Early and Middle Pleistocene of Europe, focusing on important renewals in the carnivore guilds, and their significance in terms of carrion availability for scavenging and human-carnivore competition for access to food resources. Based on a previous ecomorphological approach of carnivore guild analysis [1], a modified version was recently employed [2] and is presented herein, combining four ecomorphological/behavioral parameters of large carnivores (body mass-BM, diet, hunting strategy, sociality) that practice hunting and/or scavenging on large prey. 3D guild structure diagrams were constructed and analyzed aiming to: 1) examine the community structure and dynamics of the predatory guilds, 2) infer the possible role of carnivores in the changes of early Homo subsistence strategies (passive/active scavenging and hunting), and 3) assess the role of hominins within the guilds.
The late Villafranchian–Epivillafranchian (Early Pleistocene) carnivore guild was dominated by large-sized, hypercarnivorous and ambush-hunting felids (e.g., the saber-toothed cats Megantereon and Homotherium), and by the large-sized, bone-cracking and scavenging hyaenid Pachycrocuta. Τhe latter in particular was the most direct competitor of Homo for scavenging food resources (leftovers) left behind mainly by the saber-toothed cats [3]. As a member of the predatory guild (evident from the presence of cut and percussion marks on mammal bones), Homo would occupy the ecological space that was “available” for a predator with a 30–100 kg BM and a (mostly?) scavenging behavior, perhaps with a hypocarnivorous/carnivorous diet according to ecological circumstances and geographic setting. Τhe disappearance of most of the Early Pleistocene carnivore components (including Pachycrocuta and Megantereon) towards the end of this period, and their replacement by the Galerian (Middle Pleistocene) to modern hyenas and felids, resulted in the change of the structure and dynamics of the guild. Most notably, this reorganization included the
decrease of carrion providers (hunters), and the higher representation of species with scavenging, bone-cracking and pack-hunting behavior. In this Middle Pleistocene guild, Homo would occupy the niche that was previously held by Megantereon, in the group of predators with 30–100 kg BM. Similar to Megantereon, humans could have a carnivorous to hypercarnivorous diet, but unlike the solitary and “ambush-and-slash” felid, the biological, technological, cultural and social developments would have allowed humans to employ a modified hunting strategy: the cooperative “ambush-and-spear” strategy (in accordance with the use of hunting spears during this period). The incorporation of such hunting behavior made humans fairly independent of erratic food sources
from scavenging carnivore kills and allowed the provisioning of animal resources on a more regular basis. Moreover, even though the carnivore diversity slightly increased during this period, carnivore representation in the archaeo-palaeontological localities is rather low in both species and specimens number. This is possibly an anthropogenic effect on the ecosystem due to: 1) the firmer establishment of the hominin niche, including anti-predator strategies and expulsion of large carnivores from the region of human influence; and 2) the reduction of food quantity through human confrontational scavenging or decrease in prey availability through human hunting (see also [4] and [5]).
The late Villafranchian–Epivillafranchian (Early Pleistocene) carnivore guild was dominated by large-sized, hypercarnivorous and ambush-hunting felids (e.g., the saber-toothed cats Megantereon and Homotherium), and by the large-sized, bone-cracking and scavenging hyaenid Pachycrocuta. Τhe latter in particular was the most direct competitor of Homo for scavenging food resources (leftovers) left behind mainly by the saber-toothed cats [3]. As a member of the predatory guild (evident from the presence of cut and percussion marks on mammal bones), Homo would occupy the ecological space that was “available” for a predator with a 30–100 kg BM and a (mostly?) scavenging behavior, perhaps with a hypocarnivorous/carnivorous diet according to ecological circumstances and geographic setting. Τhe disappearance of most of the Early Pleistocene carnivore components (including Pachycrocuta and Megantereon) towards the end of this period, and their replacement by the Galerian (Middle Pleistocene) to modern hyenas and felids, resulted in the change of the structure and dynamics of the guild. Most notably, this reorganization included the
decrease of carrion providers (hunters), and the higher representation of species with scavenging, bone-cracking and pack-hunting behavior. In this Middle Pleistocene guild, Homo would occupy the niche that was previously held by Megantereon, in the group of predators with 30–100 kg BM. Similar to Megantereon, humans could have a carnivorous to hypercarnivorous diet, but unlike the solitary and “ambush-and-slash” felid, the biological, technological, cultural and social developments would have allowed humans to employ a modified hunting strategy: the cooperative “ambush-and-spear” strategy (in accordance with the use of hunting spears during this period). The incorporation of such hunting behavior made humans fairly independent of erratic food sources
from scavenging carnivore kills and allowed the provisioning of animal resources on a more regular basis. Moreover, even though the carnivore diversity slightly increased during this period, carnivore representation in the archaeo-palaeontological localities is rather low in both species and specimens number. This is possibly an anthropogenic effect on the ecosystem due to: 1) the firmer establishment of the hominin niche, including anti-predator strategies and expulsion of large carnivores from the region of human influence; and 2) the reduction of food quantity through human confrontational scavenging or decrease in prey availability through human hunting (see also [4] and [5]).
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In recent years, a significant number of Pleistocene localities with evidence of proboscidean exploitation by humans have been discovered, substantially enriching our knowledge on Homo subsistence strategies and interactions with... more
In recent years, a significant number of Pleistocene localities with evidence of proboscidean exploitation by humans have been discovered, substantially enriching our knowledge on Homo subsistence strategies and interactions with megaherbivores. However, the human engagement in proboscidean assemblages, as well as the degree of interaction is not always straightforward. First, the stratigraphic association of proboscidean remains and artefacts does not in itself necessarily imply anthropogenic processing of the fauna and the verification of their functional relation requires a taphonomical analysis (e.g., Giusti et al., 2018). Second, hominin exploitation of carcasses can be extremely difficult to demonstrate, because bone surface modifications can result from other (non-human) agents, human-induced cut marks are only rarely preserved (e.g., Haynes and Klimowicz, 2015) and possible subsequent weathering may delete direct evidences on the bone surfaces .
In this study we provide a synthesis of the Proboscidea-Homo record in Early and Middle Pleistocene open-air localities of western Eurasia, by documenting any direct (e.g., presence of cut marks, elephant bone tools, fractures for marrow extraction) and indirect (e.g., association and refitting of lithic artefacts, use-wear analysis) evidence of proboscidean carcass exploitation. Sex and ontogenetic age of butchered proboscideans are recorded, so as to assess possible preferences by humans. Furthermore, we investigate the role of the large carnivores (hyaenids, felids, large canids). We focus on important renewals in the carnivore guilds, and their significance in terms of availability of carrion for scavenging and the hominin-carnivore competition for food resources. The ecological adaptations of the two Middle Pleistocene elephantids in Europe, the straight-tusked elephant Palaeoloxodon antiquus and the steppe mammoth Mammuthus trogontherii, are also evaluated. Finally, we discuss various aspects of the evolution of Pleistocene Homo, including technological advances in material culture, important developments in cognition and relevant inferences about human social behavior. The objective of this study is to re-address the key issues in the Homo-Proboscidea research agenda, assess emerging patterns between ecological, ethological, environmental and cultural parameters, and identify potential biases that obstruct nuanced interpretations of the record.
In this study we provide a synthesis of the Proboscidea-Homo record in Early and Middle Pleistocene open-air localities of western Eurasia, by documenting any direct (e.g., presence of cut marks, elephant bone tools, fractures for marrow extraction) and indirect (e.g., association and refitting of lithic artefacts, use-wear analysis) evidence of proboscidean carcass exploitation. Sex and ontogenetic age of butchered proboscideans are recorded, so as to assess possible preferences by humans. Furthermore, we investigate the role of the large carnivores (hyaenids, felids, large canids). We focus on important renewals in the carnivore guilds, and their significance in terms of availability of carrion for scavenging and the hominin-carnivore competition for food resources. The ecological adaptations of the two Middle Pleistocene elephantids in Europe, the straight-tusked elephant Palaeoloxodon antiquus and the steppe mammoth Mammuthus trogontherii, are also evaluated. Finally, we discuss various aspects of the evolution of Pleistocene Homo, including technological advances in material culture, important developments in cognition and relevant inferences about human social behavior. The objective of this study is to re-address the key issues in the Homo-Proboscidea research agenda, assess emerging patterns between ecological, ethological, environmental and cultural parameters, and identify potential biases that obstruct nuanced interpretations of the record.
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The technological sophistication and behavior of early hominins in SouthEast Europe is poorly understood due to the scarcity of well-preserved, excavated assemblages. In this paper we present preliminary results from the study of the... more
The technological sophistication and behavior of early hominins in SouthEast Europe is poorly understood due to the scarcity of well-preserved, excavated assemblages. In this paper we present preliminary results from the study of the cultural material unearthed at the Lower Paleolithic site of Marathousa 1 (MAR-1), Megalopolis, Greece. The MAR-1 lithic assemblage is composed of small-sized flakes and flake fragments, retouched tools, cores that are commonly small and exhausted, as well as a large number of debris and retouch products, such as chips and resharpening flakes. So far, there are no indications of bifacial debitage and a key aspect of the material refers to its 'microlithic' character. The MAR-1 industry fits well in a group of important Eurasian sites with flake-based, small-tool, non-handaxe assemblages, such as Isernia, Ficoncella, Bilzingsleben, Schöningen, Vértesszölös and La Polledrara, many of which, like MAR-1, have yielded evidence of elephant or other mega-fauna exploitation. On the basis of the ongoing analysis of lithic material from three field seasons, we discuss aspects of assemblage composition and the role of raw material types, the main technological and typological traits of the industry, as well as the potential contribution of the MAR-1 evidence in broader discussions about Middle Pleistocene lithic techno-complexes and subsistence strategies in Europe. Finally, we briefly present faunal material with flake scars and bone flakes, which suggest that hominin exploitation of the carcasses was not restricted only to marrow extraction and bone processing for nutritional needs, but included also the knapping of bones, potentially with the aim of using the knapped products as tools.
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The Megalopolis Basin (Peloponnesus, Greece) has long been known for its Middle Pleistocene mammal fossils (see [1] and references therein). In 2013 a palaeolithic/palaeoanthropological survey, conducted by a joint team from the Ephorate... more
The Megalopolis Basin (Peloponnesus, Greece) has long been known for its Middle Pleistocene mammal fossils (see [1] and references therein). In 2013 a palaeolithic/palaeoanthropological survey, conducted by a joint team from the Ephorate of Palaeoanthropology Speleology of the Greek Ministry of Culture and the University of Tübingen, led to the discovery of a new open-air locality, Marathousa-1 (MAR-1), when stratified bones and lithic artefacts were identified in a section of the Marathousa Member, Choremi Formation [2]. The preliminary ESR analysis dates the locality at 0.5-0.4 Ma [3]. The large mammal faunal material collected so far (2013-2016) includes the castorid Castor fiber, the mustelids Lutra simplicidens and Mustela sp., the felid Felis sp., the canids Vulpes sp. and Canis sp., the elephantid Elephas (Palaeoloxodon) antiquus, the hippopotamid Hippopotamus antiquus, the bovid Bison sp., and the cervids Dama sp. and Cervus elaphus. This faunal association is common in the Galerian mammal communities of Europe (ca. 0.9-0.4 Ma). Furthermore, it is consistent with a temperate climate, and is indicative of a landscape with substantial woodland components and more open areas, close to permanent and large freshwater bodies. Of particular interest are an elephant cranium and numerous postcranial elements, which were found in close anatomical association and are attributed to a single individual of the straight-tusked elephant Elephas (P.) antiquus. The skeleton belonged to an old male in its sixties, with live skeletal height around 3.7 meters and body mass around 9.0 tones. The good state of preservation of the MAR-1 bones allows the identification of taphonomic modifications. Three-dimensional virtual reconstructions of marks with the use of a confocal microscope enabled the identification of cut marks and their micromorphological characteristics. Cut marks and percussion damage indicate hominin exploitation of the elephant carcass and other mammal bones by means of butchering activities, which is in accordance
with the lithic assemblage and its spatial association with the bones. Carnivore gnawing is also evident on some specimens, suggesting a certain degree of carnivore competition with humans for early access to the animal carcasses. Marathousa-1 is among the oldest elephant butchering sites in Europe and the only one known in Southeastern Europe.
with the lithic assemblage and its spatial association with the bones. Carnivore gnawing is also evident on some specimens, suggesting a certain degree of carnivore competition with humans for early access to the animal carcasses. Marathousa-1 is among the oldest elephant butchering sites in Europe and the only one known in Southeastern Europe.
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Introduction: The fossiliferous locality Chomateri (Attica, Greece), near the classical late Miocene site Pikermi, was discovered in 1971 and excavated from 1972 until 1980. The large mammal assemblage from Chomateri is not studied in... more
Introduction: The fossiliferous locality Chomateri (Attica, Greece), near the classical late Miocene site Pikermi, was discovered in 1971 and excavated from 1972 until 1980. The large mammal assemblage from Chomateri is not studied in detail as a whole, so its faunal context and correlation with Pikermi (end of MN 12), as well as with other Turolian localities of Greece and the wider region, are not yet entirely clear. Concerning the proboscideans, in the rich vertebrate collections of Pikermi four taxa are included –Deinotherium proavum, Mammut sp., Choerolophodon pentelici, Konobelodon atticus–, whereas in the by far smaller Chomateri collections (Athens, Vienna), the only two known proboscidean specimens (maxilla with DP2–DP4, isolated dp4) belong to another taxon.
Results: The dislocation of the half-loph(id)s and the resultant alternate arrangement of the successive loph(id)s (anancoidy), are diagnostic features of Anancus, which are present in the dp4/DP4 from Chomateri and allow the attribution to this genus. However, the anancoidy is rather weak and the occlusal morphology simple, both regarded as primitive features within anancines. This could indicate an early evolutionary stage for the Chomateri anancine, in agreement with the Turolian age of the locality. Nonetheless, recent studies reveal the high morphological variability within Anancus populations, which might display a mixture of primitive/derived dental characters. Thus, any biochronological conclusion based on isolated specimens should be always taken with caution. The limited comparative material and the problematic taxonomy of the European late Miocene anancines do not permit a safe specific allocation of the Chomateri specimens, and they are attributed for the moment to Anancus sp. The presence of this genus in Chomateri consists the first faunal evidence that the locality post-dates Pikermi. This age difference is considered to be relatively small due to their overall faunal similarity and their faunal differences with the Dytiko localities (MN 13, Axios Valley, Greece).
Conclusions: Previously known only from the Plio-Pleistocene, this is the first report of Anancus in the late Miocene of Greece, marking its earliest occurrence in the Greek fossil record. Based on the current data, the migration of Anancus to Southern Europe (e.g., Spain, Southern Balkans) took place during the end of the middle Turolian, at ~7.2–7.1 Ma, a period that roughly coincides with the decline of the “Pikermian” large mammal fauna. Late Miocene anancines were probably inhabitants of mosaic or even more wooded environments. This further reinforces the existing palaeoecological reconstructions of Greece and the wider region, which indicate a gradual transition to a more closed environment towards the end of the Turolian.
Results: The dislocation of the half-loph(id)s and the resultant alternate arrangement of the successive loph(id)s (anancoidy), are diagnostic features of Anancus, which are present in the dp4/DP4 from Chomateri and allow the attribution to this genus. However, the anancoidy is rather weak and the occlusal morphology simple, both regarded as primitive features within anancines. This could indicate an early evolutionary stage for the Chomateri anancine, in agreement with the Turolian age of the locality. Nonetheless, recent studies reveal the high morphological variability within Anancus populations, which might display a mixture of primitive/derived dental characters. Thus, any biochronological conclusion based on isolated specimens should be always taken with caution. The limited comparative material and the problematic taxonomy of the European late Miocene anancines do not permit a safe specific allocation of the Chomateri specimens, and they are attributed for the moment to Anancus sp. The presence of this genus in Chomateri consists the first faunal evidence that the locality post-dates Pikermi. This age difference is considered to be relatively small due to their overall faunal similarity and their faunal differences with the Dytiko localities (MN 13, Axios Valley, Greece).
Conclusions: Previously known only from the Plio-Pleistocene, this is the first report of Anancus in the late Miocene of Greece, marking its earliest occurrence in the Greek fossil record. Based on the current data, the migration of Anancus to Southern Europe (e.g., Spain, Southern Balkans) took place during the end of the middle Turolian, at ~7.2–7.1 Ma, a period that roughly coincides with the decline of the “Pikermian” large mammal fauna. Late Miocene anancines were probably inhabitants of mosaic or even more wooded environments. This further reinforces the existing palaeoecological reconstructions of Greece and the wider region, which indicate a gradual transition to a more closed environment towards the end of the Turolian.
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The Megalopolis Basin (Peloponnesus, Greece) has long been known for its Middle Pleistocene mammal fossils (see Melentis, 1961). In 2013 a palaeolithic/palaeoanthropological survey, conducted by a joint team of the Ephorate of... more
The Megalopolis Basin (Peloponnesus, Greece) has long been known for its Middle Pleistocene mammal fossils (see Melentis, 1961). In 2013 a palaeolithic/palaeoanthropological survey, conducted by a joint team of the Ephorate of Palaeoanthropology-Speleology of the Greek Ministry of Culture and the University of Tübingen in the frame of the ERC project PaGE, led to the discovery of a new locality –Marathousa 1 (MAR)–, when stratified bones and lithic artefacts were identified in a section of the Marathousa Member, Choremi Formation (Panagopoulou et al., 2015). The large mammal faunal material collected so far (2013–2016) includes the castorid Castor fiber, the mustelids Lutra simplicidens and Mustela sp., the felid Felis sp., the canid Canis sp., the elephantid Elephas (Palaeoloxodon) antiquus, the hippopotamid Hippopotamus antiquus, the bovid Bison sp., and the cervids Dama sp. and Cervus elaphus. This faunal association is common in the Galerian mammal communities of Europe (ca. 0.9–0.4 Ma). Furthermore, it is consistent with a temperate climate, and is indicative of a landscape with substantial woodland components and more open areas, close to permanent and large freshwater bodies.
Of particular interest are an elephant cranium and several postcranial elements, which were found in close anatomical association and are attributed to a single individual of the straight-tusked elephant Elephas (P.) antiquus. The good state of preservation of the MAR bones allows to identify taphonomic modifications. Cut marks and percussion damage indicate hominin exploitation of elephant and other mammal bones by means of butchering activities. Carnivore gnawing is also evident on some specimens, suggesting a certain degree of carnivore competition with humans for early access to the animal carcasses.
Of particular interest are an elephant cranium and several postcranial elements, which were found in close anatomical association and are attributed to a single individual of the straight-tusked elephant Elephas (P.) antiquus. The good state of preservation of the MAR bones allows to identify taphonomic modifications. Cut marks and percussion damage indicate hominin exploitation of elephant and other mammal bones by means of butchering activities. Carnivore gnawing is also evident on some specimens, suggesting a certain degree of carnivore competition with humans for early access to the animal carcasses.
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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... more
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 University of Thessaloniki and the University of Tübingen in 2014 a new locality –named Tsiotra Vryssi (TSR)– was discovered. The collected material includes the corvid Corvus pliocaenus (Aves), the hyaenid Pachycrocuta brevirostris, the canid Canis etruscus, the rhinocerotid Stephanorhinus sp., two species of Equus, the bovid Bison sp., the cervid Metacervocerus rhenanus and the giraffid Palaeotragus sp. The locality was further excavated in 2015 and the fauna was significantly enriched by the mammoth Mammuthus meridionalis, the bovid Leptobos sp., the antelope Pontoceros ambiguus and the giant cervid Praemegaceros sp. The presence of Pachycrocuta brevirostris clearly indicates a late Villafranchian age for TSR, while the presence of two Equus species, Praemegaceros sp., Pontoceros ambiguus, and the co-occurrence of Leptobos and Bison, suggest that TSR is chronologically intermediate between Gerakarou-1 and Apollonia-1 (Mygdonia Basin), and therefore it can be preliminary dated to 1.8–1.2 Ma. The TSR fauna will provide additional palaeogeographical and palaeoecological information about a crucial time interval for mammal migrations and turnovers, as well as for the first hominin dispersals from Africa to Europe.
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Research Interests: Proboscidea and Miocene
The late Miocene proboscideans of Greece, although they are known since the middle of the 19th Century, they have never been studied in detail. Their collection includes numerous specimens and increased remarkably during the last decades... more
The late Miocene proboscideans of Greece, although they are known since the middle of the 19th Century, they have never been studied in detail. Their collection includes numerous specimens and increased remarkably during the last decades by the excavations in the Axios Valley and Nikiti (Macedonia, Greece), as well as in Samos Island (Central-Eastern Aegean Sea). The detailed study of this material provided significant results concerning their taxonomy, evolution, biochronology and palaeoecology.
The deinotheriids are represented by two species, Deinotherium giganteum and Deinotherium gigantissimum (= D. proavum). The former is recognized in the late Vallesian of Axios Valley and the latter in the Turolian of Samos, Pikermi (Attica), Halmyropotamos (Euboea) and Perivolaki (Thessaly). The mammutids, present in the Turolian of Axios Valley, Samos, Pikermi and Halmyropotamos, belong to “Mammut” sp. The choerolophodont material belongs to the genus Choerolophodon, which dominates in all late Miocene localities of Greece. Two species and three evolutionary stages of Choerolophodon have been recognized in the late Miocene of the peri-Mediterranean region. The more primitive species C. anatolicus, recognized in the early Vallesian of Axios Valley, corresponds to the first evolutionary stage. The advanced species C. pentelici, known from Pikermi, Samos, Axios Valley and Nikiti, can be distinguished in two stages: a primitive one recognized in the late Vallesian and an advanced one in the Turolian. The cladistic analysis of the Miocene Old World Choerolophodon taxa proposes that C. anatolicus is a sister group of C. corrugatus and C. pentelici. The study of the Pikermi tetralophodont material indicates that two late Miocene species, “Tetralophodon” atticus and “Mastodon” grandincisivus, are synonyms. The morphology of the Pikermi tetralophodons allowed their attribution to the tetralophodont shovel-tusker Konobelodon atticus. This species encompasses all Turolian tetralophodont shovel-tuskers from Eastern Europe-Western Asia. In Greece it is also present at Samos and Kerassia (Euboea). The Chomateri (Attica, Turolian) material belongs to Anancus sp., representing the first evidence of the genus in the late Miocene of Greece.
Concerning biochronology, the genus Deinotherium is known from the Vallesian with the species D. giganteum and from the Turolian with D. gigantissimum. “Mammut” sp. is known from the early and middle Turolian. Choerolophodon anatolicus is present in the early Vallesian (MN 9), whereas C. pentelici has a longer stratigraphic distribution; its primitive form is known from the late Vallesian (MN 10), while the most advanced one covers the entire Turolian (MN 11-13). Konobelodon atticus is known from the middle Turolian of Pikermi, whereas its presence at Samos cannot be dated because it lacks locality information.
The dental microwear analysis of Choerolophodon from Axios Valley and Nikiti indicates an open environment. These results fit well with previous palaeoecological studies of the late Miocene faunas of Greece. Moreover, the absence of Tetralophodon from the Vallesian deposits of Axios Valley and the wider area is probably due to the different palaeoecological conditions between Central and Southeastern Europe. This is probably the reason why Choerolophodon could not migrate to Central Europe, although it was widespread in the Greco-Iranian Province. During the Turolian, the coexistence of Deinotherium gigantissimum, “Mammut” sp., Choerolophodon pentelici and Konobelodon atticus is probably attributed to the different ecological niches that they occupied. The appearance of Anancus in the second half of Turolian coincides with the gradual transition to a more closed environment and more humid conditions towards the end of Turolian.
The deinotheriids are represented by two species, Deinotherium giganteum and Deinotherium gigantissimum (= D. proavum). The former is recognized in the late Vallesian of Axios Valley and the latter in the Turolian of Samos, Pikermi (Attica), Halmyropotamos (Euboea) and Perivolaki (Thessaly). The mammutids, present in the Turolian of Axios Valley, Samos, Pikermi and Halmyropotamos, belong to “Mammut” sp. The choerolophodont material belongs to the genus Choerolophodon, which dominates in all late Miocene localities of Greece. Two species and three evolutionary stages of Choerolophodon have been recognized in the late Miocene of the peri-Mediterranean region. The more primitive species C. anatolicus, recognized in the early Vallesian of Axios Valley, corresponds to the first evolutionary stage. The advanced species C. pentelici, known from Pikermi, Samos, Axios Valley and Nikiti, can be distinguished in two stages: a primitive one recognized in the late Vallesian and an advanced one in the Turolian. The cladistic analysis of the Miocene Old World Choerolophodon taxa proposes that C. anatolicus is a sister group of C. corrugatus and C. pentelici. The study of the Pikermi tetralophodont material indicates that two late Miocene species, “Tetralophodon” atticus and “Mastodon” grandincisivus, are synonyms. The morphology of the Pikermi tetralophodons allowed their attribution to the tetralophodont shovel-tusker Konobelodon atticus. This species encompasses all Turolian tetralophodont shovel-tuskers from Eastern Europe-Western Asia. In Greece it is also present at Samos and Kerassia (Euboea). The Chomateri (Attica, Turolian) material belongs to Anancus sp., representing the first evidence of the genus in the late Miocene of Greece.
Concerning biochronology, the genus Deinotherium is known from the Vallesian with the species D. giganteum and from the Turolian with D. gigantissimum. “Mammut” sp. is known from the early and middle Turolian. Choerolophodon anatolicus is present in the early Vallesian (MN 9), whereas C. pentelici has a longer stratigraphic distribution; its primitive form is known from the late Vallesian (MN 10), while the most advanced one covers the entire Turolian (MN 11-13). Konobelodon atticus is known from the middle Turolian of Pikermi, whereas its presence at Samos cannot be dated because it lacks locality information.
The dental microwear analysis of Choerolophodon from Axios Valley and Nikiti indicates an open environment. These results fit well with previous palaeoecological studies of the late Miocene faunas of Greece. Moreover, the absence of Tetralophodon from the Vallesian deposits of Axios Valley and the wider area is probably due to the different palaeoecological conditions between Central and Southeastern Europe. This is probably the reason why Choerolophodon could not migrate to Central Europe, although it was widespread in the Greco-Iranian Province. During the Turolian, the coexistence of Deinotherium gigantissimum, “Mammut” sp., Choerolophodon pentelici and Konobelodon atticus is probably attributed to the different ecological niches that they occupied. The appearance of Anancus in the second half of Turolian coincides with the gradual transition to a more closed environment and more humid conditions towards the end of Turolian.
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In recent decades, a significant number of Pleistocene (ca. 2.6 million years–10,000 years ago) open-air and cave sites yielding elephant or mammoth bones in direct association with hominin remains and/or lithic artifacts have been... more
In recent decades, a significant number of Pleistocene (ca. 2.6 million years–10,000 years ago) open-air and cave sites yielding elephant or mammoth bones in direct association with hominin remains and/or lithic artifacts have been discovered in Eurasia, Africa and America. Many of them show strong evidence of acquisition and processing of proboscidean carcasses by early humans, leading scientists to interpret them as “elephant butchering sites”. Indeed, proboscidean exploitation by early Homo has been proposed to have been critical for Palaeolithic human lifeways, influencing not only their subsistence, but also other aspects of early human evolution and adaptations. The nature and degree of interactions between humans and elephants comprises an important field in palaeoanthropological studies since decades, but many questions remain still unanswered or partially explored. By bringing together research papers from the fields of Palaeolithic Archaeology, Palaeoanthropology, Palaeontology, Zooarchaeology, Geology, Ethnography and Nutrition Studies, the book systematically covers a diverse array of perspectives on elephant-human interactions across the world from the Pleistocene times until today. The volume includes 19 contributions and is organized into four thematic sections: 1) The Palaeolithic record, 2) A view of the evidence, 3) Elephants in past human nutrition, and 4) Ethnography – Human-elephant interactions in recent Africa. Collectively, the volume not only showcases the current state of knowledge, but also intends to provoke renewed interest for current and further research, and build an interdisciplinary and synthetic understanding of the significance of proboscideans throughout human evolution.