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The Role of the Central Balkans in the Peopling of Europe: Paleoanthropological Evidence

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Abstract

The paucity of fossil human remains from the Central Balkans represents a very serious lacuna in our understanding of human evolution in the Pleistocene of Europe, which is—as a result—strongly influenced by the material from the better researched parts of the continent further to the west of the Balkans. The scant fossil record from the Central Balkans suffers from a lack of archaeological/geological context, and with the exception of the Balanica hominin (BH-1) has no associated chronological data. In this chapter, I present all of the purported Pleistocene specimens currently known from the area and discuss their possible affinities.

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... In other European MP specimens, such as those of Ceprano (Italy), Aroeira (Portugal), Mala Balanica (Serbia) and Arago (France), the Neanderthal affinities are less clear (e.g. Bermúdez de Castro et al. 2019;Daura et al. 2017;Manzi, 2016;Roksandic 2016;Roksandic et al. 2011;Skinner et al. 2016) attesting for the possible coexistence of at least two hominin lineages in the European MP (e.g. Bermúdez de Castro et al. 2016;Dennell et al. 2011;MacDonald et al. 2012). ...
... One group would cluster specimens that are lacking Neanderthal apomorphies in their dentitions such as Mala Balanica (BH-1), Mauer or Arago (e.g. Bailey 2002;Bermúdez de Castro et al. 2019;Gómez-Robles et al. 2007, 2011Roksandic 2016;Skinner et al. 2016). While, the second group would be characterised by closer morphological dental affinities with the classic Neanderthals, including the Atapuerca-SH, Pontnewydd, Fontana Ranuccio, Visogliano, Steinheim and Montmaurin hominins (e.g. ...
... As such, based on dental analyses, two groups are recognised; the first one clusters specimens characterised by a more primitive morphology with less to none Neanderthal affinities such as Mala Balanica (BH-1), Mauer or Arago (e.g. Bailey 2002;Bermúdez de Castro et al. 2019;Gómez-Robles et al. 2007, 2011Roksandic, 2016;Skinner et al. 2016). The second group includes those hominins exhibiting most (if not all) the dental features that are considered typical of the Neanderthal species, including the Atapuerca-SH, Pontnewydd, Fontana Ranuccio, Visogliano, Steinheim, Montmaurin and BSV hominins (e.g. ...
Article
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The study of dental morphology can be a very useful tool to understand the origin and evolution of Neanderthals in Europe during the Middle Pleistocene (MP). At present, the earliest evidence, ca. 430 ka, of a pre-Neanderthal population in Europe is the hominin sample from Atapuerca-Sima de los Huesos (SH) that present clear dental affinities with Neanderthals while other penecontemporaneous populations, such as Arago or Mala Balanica, exhibit less Neanderthal traits. We present the morphometric study of the external and internal dental structures of eleven hominin dental remains recovered from the MP, ca. 240 ka, French site of Biache-Saint-Vaast (BSV). Our analyses place the BSV hominins within the MP group, together with SH, Fontana Ranuccio, Visogliano, Steinheim or Montmaurin, that show greater morphological affinities with Neander- thals. Moreover, we identified interpopulation variability in the expression of the enamel thickness trait, with BSV hominins sharing the unique combination of thin and thick pattern in the premolars and molars with the SH population. These results further support the coexistence of two or more populations in Europe during the MP that reflect the population and settlement of human groups suggested by the Central Area of Dispersals of Eurasia (CADE) and sink and source model.
... The Central Balkans are situated at the center of an important geographical, biological, and cultural crossroads (Roksandic, 2016). The major migration corridors of the Danube (EasteWest) and Great Morava (NortheSouth) Rivers cut through the mountainous terrain of the Balkan Peninsula, and are likely routes for the earliest migrations of archaic and modern humans into Europe (Conard and Bolus, 2003;Mihailovi c, 2009;Roksandic et al., 2018). ...
... Apart from several important Neanderthal sites in Croatia (Jankovi c et al., 2016), Neanderthal and early modern human remains in Greece (Harvati, 2016;Harvati et al., 2019), and Late Pleistocene early modern humans in Romania and Bulgaria (Harvati and Roksandic, 2016;Hublin et al., 2020), human fossils from the rest of the peninsula are rare. In the Central Balkans, comprising parts of current Serbia, Bosnia and Herzegovina, Bulgaria, and Northern Macedonia (Roksandic, 2016), nonmodern human fossils are known from only three sites. An infant radius from Kozarnika Cave, Bulgaria (Tillier et al., 2017) and a non-adult radius (Pes-2) from Pe sturina Cave, southeast Serbia (Lindal et al., 2020) have been tentatively assigned to Neanderthals. ...
... Hominins from each of these sites present morphological particularities that are difficult to reconcile with the prevalent understanding of human evolution in Europe as a mosaic progression from the common ancestor towards Neanderthals (Roksandic et al., accepted). Dated to older than 400 ka, the Mala Balanica mandible shows no Neanderthal traits Roksandic, 2016); the Petralona cranium shows a mix of African Bodo/ Kabwe-like traits (e.g., Stringer, 1974;Rightmire, 2008;Freidline et al., 2012), the persistence of primitive H. erectus-like traits (e.g., Rightmire, 1990;Cartmill and Smith, 2009), and a Neanderthal-like mid-facial morphology (e.g., Hublin, 1998;Dean et al., 1998;Freidline et al., 2012). However, the Petralona M 1 crown is absolutely large (134.5 mm 2 ), with a relatively large hypocone (23.0%) and a relative cusp size sequence of PR > PA > HY > ME (reviewer 1, personal comm.), ...
Article
Neanderthals are Eurasian fossil hominins whose distinctive morphology developed in the southwestern corner of Europe and later spread throughout the continent, reaching Southwest Asia before the Late Pleistocene and spreading into Central Asia by 59–49 ka. The timing, tempo, and route of the Neanderthal movements eastward are poorly documented. The earliest probable evidence of Neanderthals in Asia comes from Karain E Cave (Anatolia, Turkey), dated to 250–200 ka. We present four Chibanian (Middle Pleistocene) hominin specimens, representing at least two individuals, from Velika Balanica Cave (Serbia): a permanent upper third molar (BH-2), a deciduous upper fourth premolar (BH-3) refitted to a poorly preserved maxillary fragment with the permanent first molar in the alveolus (BH-4), and a permanent upper central incisor (BH-5). We provide descriptions of the teeth, as well as a comparative analysis of the well-preserved M1 (BH-4), including assessments of cusp angles, relative occlusal polygon area, relative cusp base areas, two- and three-dimensional enamel thickness, and taurodontism. Morphology of both the occlusal surface and the enamel dentine junction of the M1 indicates that the maxillary fragment and associated dP4 belonged to an early Neanderthal child. The heavily worn I1 and M3 are consistent with the Neanderthal morphology, although they are less distinct taxonomically. These Chibanian remains with provenance from layer 3a are constrained by two thermoluminescence dates: 285 ± 34 ka and 295 ± 74 ka. They represent the earliest current evidence of Neanderthal spread into the Eastern Mediterranean Area. We discuss these findings in light of recent direct evidence for cultural connections between Southwestern Asia and Southeast Europe in the Chibanian.
... A left semi-mandible, BH-1, from Mala Balanica, Serbia, represents the only Middle Pleistocene hominin specimen from the Central Balkans (Roksandic, 2016). Mala Balanica cave, which together with Velika Balanica forms the Balanica cave complex in the Si cevo Gorge, has been the focus of systematic research since 2004 (Mihailovi c, 2009;Mihailovi c and Bogi cevi c, 2016). ...
... The excavations are ongoing and the bedrock has not been reached in either of the caves. The details of the morphology of the BH-1 mandible are described elsewhere (Roksandic et al., 2011;Skinner et al., 2016;Roksandic, 2016). Here we will briefly outline its most salient aspects, which show that the mandible, as well as its preserved dentition, are characterized by a complete lack of derived Neanderthal traits. ...
Article
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The hominin mandible BH-1 from the Middle Pleistocene cave of Mala Balanica suggested the possibility that human populations in this part of the continent were not subject to the process of Neanderthalization observed in the west. We review the paleoanthropological evidence from the Central Balkans in the context of the Eastern Mediterranean geographic entity. The current hominin fossil record of the early Middle Pleistocene in the region suggests that Europe was inhabited by two different populations: a population in the west of the continent with derived Neanderthal morphology; and a more variable population in the east characterized by a combination of plesiomorphous and synapomorphous traits. We suggest that - in order to continue using the nomenclature of Homo heidelbergensis - the current hypodigm needs to be revised to include only the specimens from the latter group.
... D. Refugial character: Palaeofaunal evidence and pollen data suggest that the Balkans and the wider E-NE Mediterranean provided refugia (Hewitt, 1999;Tzedakis, 2004;Palombo et al., 2006;Stewart et al., 2010;Dennell et al., 2011) sustaining populations during the glacial periods that were able to repopulate previously abandoned areas to the north and the west when climatic conditions ameliorated. Moreover, these conditions permitted undisrupted movement and communication, via multiple ice-free routes between SE Europe and SW Asia throughout the Pleistocene (Roksandic, 2016;Dobos and Iovita, 2016;Ivanova, 2016;Strait et al., 2016). Recent comparative analyses of faunal evidence from Eurasian sites with an early hominin presence suggest that the Peri-Pontic routes (around the Black Sea) and the Bosporus passage (via W. Anatolia) could be associated with the initial peopling of Europe at-or slightly prior to 1 Mya (Spassov, 2016;Koufos and Kostopoulos, 2016). ...
... Changes in palaeogeography are progressively acknowledged, and considered a significant component in the history of hominin movements during the Early and Middle Pleistocene (e.g. Galanidou et al., 2016;Roksandic, 2016;Strait et al., 2016). In the last decade, several mentions have been made of potential trans-Aegean passages (Tourloukis and Karkanas, 2012;Strait et al., 2016;Carter et al., 2019) but they are quite generalised. ...
Article
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The Balkan Peninsula lies on a key geographical location between Africa and Eurasia. The southern part of the Balkan Peninsula, referred to as the Aegean region was a passageway for migrations of Eurasian fauna throughout the Pleistocene. Recent advances in the Lower Palaeolithic archaeology of Greece and in the research of the submerged landscapes of the Aegean region prompt a reconsideration of the biogeographical role of this area in Middle Pleistocene hominin settlement and migration. In this paper, we articulate and elaborate on a working hypothesis, namely that the Aegean region was not a barrier during the Middle Pleistocene but instead it offered attractive lands for occupation and viable pathways for dispersal. Several methodological challenges emerge associated with the dynamic nature of the Aegean context. We propose an interdisciplinary approach to address the challenges. Using available archaeological and palaeogeographical evidence within a GIS-based framework, we seek to explore the ‘hominin factor’ in relation to the changing Aegean landscapes and geographical affordances. Shifting away from the ‘terrestrial Eurocentric’ point of view in the discussion of the patterns of early colonisation of Europe and the dispersal routes followed by hominins, we propose a revised understanding of early European settlement and its eastern gateways.
... The biogeographic importance of the Balkan Peninsula in general, and the Central Balkans in particular, stands in stark contrast to the evidence from the area, with very few multiple occupation sites and a very restricted number of Middle Paleolithic sites in general (Mihailovi c, 2014a;Harvati and Roksandic, 2016). A high degree of competition with carnivores could have limited the ability of hominins to settle in the area (Churchill, 2014); however, paucity of research could be a more parsimonious explanation (Roksandic, 2016). ...
... The intensification of surveys and excavations of Paleolithic sites in Serbia over the last two decades (Mihailovi c, 2008;Mihailovi c, 2014a and references therein;Marín-Arroyo and Mihailovi c, 2017) has resulted in the discovery of 15 cave and around 10 open-air Paleolithic sites in the region. Hominin fossil remains from two of these sites enabled us to propose a different role for the Balkans in the Middle Pleistocene Roksandic, 2016;Skinner et al., 2016). However, Pe sturina Cave, near Ni s, represents the only Middle Paleolithic site from the Late Pleistocene in the territory of southern Serbia to date. ...
Article
Neanderthals were the only human group in Europe throughout the Late Pleistocene until the arrival of modern humans, and while their presence has been confirmed in the surrounding regions, no Neanderthal fossils are known to date from the Central Balkans. Systematic excavations of Pešturina Cave (Serbia) resulted in the discovery of a permanent right M1 (Pes-3). The specimen was recovered from stratigraphic Layer 4b with an estimated age of 102.4 ± 3.2 ka, associated with Mousterian artifacts. The exceptional state of preservation and minimal wear of the molar enabled a detailed description and comparative analysis of the inner and outer dental structure, including non-metric dental traits and morphometric features of the crown, roots, and dental tissues. The results of this study strongly support the identification of Pes-3 as Neanderthal. Non-metric traits of the occlusal surface of the crown, enamel-dentine junction, and roots are consistent with Neanderthal morphology. The crown shows morphometric features typical for Neanderthal M1, such as a buccolingually skewed crown shape, internally compressed cusps, and a relatively large hypocone. The specimen also shows Neanderthal-like dental tissue proportions, characterized by relatively thin enamel and large coronal dentine and coronal pulp volumes. The discovery of the Pes-3 molar therefore confirms the presence of Neanderthals in the territory of Serbia and the Central Balkans at the end of Marine Isotope Stage (MIS) 5c.
... The Balanica cave complex, which comprises the Velika and Mala Balanica caves, is situated 10 km east of Niš at the exit of the Sićevo gorge. A hominin mandible attributed to Homo erectus s.l. ( Roksandic et al. 2011;Roksandic 2016), recently dated to a period prior to 397-525 ka BP ( Rink et al. 2013), was discovered in the lowest excavated layer (3b) at Mala Balanica in 2006. In addition to the mandible, a Charentiantype industry was confirmed from layer 2a-2c in Mala Balanica and layers 3a-3b in Velika Balanica ( Fig. 9.5), while Typical Mousterian was recorded in the upper layers of Velika Balanica (2a-2c) ( Mihailović 2008bMihailović , 2009a). ...
Chapter
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Recent archaeological investigations have enabled preliminary insight into the Lower to Middle Paleolithic transition in the Central Balkans. Industries containing tools made from pebbles and flakes, within which Levallois artifacts were present to a lesser (Kosovska Kosa) or greater (Samaila) extent, have been encountered at the sites in the Zapadna Morava valley. The Charentian, likely dating to the Middle Pleistocene (possibly MIS 7) on the basis of microfaunal remains, has been reported in Velika and Mala Balanica in Sićevo. With regard to later (MIS 5–4) industries, assemblages of Typical Mousterian (Crvena Stijena, Hadži Prodanova cave), Charentian (Pešturina) and assemblages where Taubachian–Charentian component, Charentian elements, and backed bifaces are combined (Petrovaradin fortress) are encountered in the Central Balkans. After examining all available data, we propose the hypothesis that in addition to climatic, ecological, and behavioral factors, demographic factors also probably had considerable impact on the variability of lithic assemblages. Migrations and cultural transmission could have resulted in the appearance of Near Eastern elements in the Central Balkans as well as Balkan elements in the Near East. The homogeneity and/or variability of industries could be considerably influenced by the degree of isolation of human groups living in this region.
... 2013;Mihailović 2014). The redating of the hominin mandible from the cave of Mala Balanica (Serbia) by combined application of ESR/U-series and infrared/post-infrared luminescence dating provided a minimum age between 397 and 525 ka (Rink et al. 2013), which makes the BH-1 mandible (Roksandic et al. 2011;Roksandic 2016) the first human fossil in the Central Balkans recovered from controlled excavations and the easternmost hominin specimen securely dated to the Middle Pleistocene. ...
Chapter
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Lower Paleolithic evidence from the Mediterranean region holds a prominent position in discussions about the earliest peopling of Europe. Most studies examining patterns of human occupation focus on purported behavioral capacity, habitat preference, and environmental tolerance of different hominins. This chapter employs a geoarchaeological perspective through the examination of landscape dynamics as a complementary approach. In this context, Lower Paleolithic records of the Mediterranean and the Balkans are reviewed with an emphasis on the geomorphological settings of the best-studied sites. Since most of the oldest, well dated and primary-context material occurs in open-air sites situated in basins, the last part of the chapter explores how basin dynamics could have conditioned the preservation and accessibility of artifact-bearing strata. Spain, Italy, and Greece are used as case-studies and a conceptual model is proposed as a means to assess possible patterned relationships of site locations. A “basin model” offers a working hypothesis for evaluating site distributions and outlines first steps towards a geosciences-based methodology, which can be used to locate new sites.
... This unfortu- nate situation is likely due to the lack of a strong tradition in basic Paleolithic research in the region. Nowhere is this data gap more evident than in the human fossil record (similarly to the situation in the Central Balkans, Bulgaria, and Anatolia, see Aytek and Harvati 2016;Roksandic 2016;Strait et al. 2016). This chapter reviews the existing human fossil evidence from Greece in the framework of the research questions outlined above. ...
Chapter
Greece lies at the crossroads between Europe, Asia, and Africa, and represents a logical gateway through which early human populations might have repeatedly passed on the way to and from Europe. It also represents one of the three European Mediterranean peninsulas which acted as a refugium for fauna, flora and, very likely, human populations during glacial times. Evidence from this region is therefore essential in order to test hypotheses about the course of human evolution in Europe. Despite the importance of the region, paleoanthropological research has until recently been relatively neglected. In recent years, however, renewed research efforts have produced new human fossils from Greece, recovered from excavated contexts. This chapter reviews the Greek human fossil evidence in the context of broader questions in European paleoanthropology.
... The fact that the Lower Paleolithic finds from Croatia are rare probably does not reflect a lack of human habitation during the Early/Middle Pleistocene, but more likely ecological-geological-climatic fluctuation and changes in the sea level during this time, preservation of sediments from this period, and the relative lack of research in the past. Croatia is very rich in archaeological heritage, especially in the coastal region, where the abundance of monuments and sites from antiquity and the Middle Ages has resulted in most research focusing on these younger periods, similarly to other areas discussed in this book (see Harvati 2016;Roksandic 2016;Dinçer 2016). This has changed over the years, and it is to be expected that new research will result in a more detailed knowledge of the earliest phases of the Paleolithic habitation of Croatia in the future. ...
Chapter
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In this chapter, we discuss Croatian sites that have yielded human skeletal remains from the Pleistocene. These include the well-known Neandertal localities Hušnjakovo (at Krapina) and Vindija cave, as well as the Late Upper Paleolithic hominin fossil site Šandalja II cave in Istria. The Krapina site played an important role in the historical development of paleoanthropology and is still the Neandertal site with the largest known minimum number of skeletal individuals to date. Finds from Vindija cave belong to one of the latest Neandertal groups in Europe and provide data for the study of both their behavioral, as well as biological characteristics (including genomics studies). The Šandalja II cave in Istria is the only site in Croatia with direct association of human skeletal finds and the late Paleolithic, an Epigravettian industry, providing us with data on the anatomy and behavior of the Late Paleolithic inhabitants of this region.
... Earlier hominins, as well as Upper Paleolithic humans, are not known, with a few possible exceptions. Chapter 2 (Roksandic 2016) presents the fossil record from the Central Balkans, highlighting the recent fossil human find from Mala Balanica. Roksandic puts forth the possibility for an alternative course for human evolution in this part of Europe, different from the one proposed by the accretion hypothesis for the Western part of the continent. ...
... Bu bağlamda, Anadolu'daki insan fosilleri hem Homo genusunun ilk temsilcilerinin, hem de sonraki Homo türlerinin göçlerinin aydınlatılması bakımından büyük önem taşımaktadır. Bu önemli coğrafi pozisyonu ve komşu ülkeler ile benzerliklerine rağmen Anadolu'da gerçekleştirilmiş çalışma çok azdır ve bu yüzden fosil insanlar çok az buluntu ile temsil edilmektedir (Harvati vd., 2009;Harvati, 2016;Roksandic, 2016;Strait vd., 2016). Paleolitik çağa ait kamp alanları açısından Anadolu büyük öneme sahiptir. ...
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İnsanın Afrika’dan çıkıp dünyaya yayılımı paleoantropolojinin en önemli konularından birisidir. Bu göçlerin ne zaman, hangi rotalar kullanılarak ve hangi insan türleri tarafından gerçekleştirildiği önemli sorular olarak karşımıza çıkmakta, bulunan her yeni fosil bu sorulara ait yeni cevapları ortaya koymaktadır. Anadolu, üç kıtanın arasında yer alması sebebiyle bu kıtalar için önemli bir köprü görevi görür. Coğrafi pozisyonuna rağmen, paleoantropolojik çalışmalar kısıtlı olduğu için fosil insan kalıntıları çok azdır. Fosil insan kalıntılarının çoğu erken dönem çalışmalarında bulunmakla beraber, son yıllarda yapılan yeni çalışma ve keşifler Anadolu’nun bu göçlerdeki önemini daha çok açığa çıkarmaktadır. Bu çalışma Anadolu’da bulunmuş ve Paleolitik döneme tarihlendirilmiş fosil insan kalıntıları hakkında yapılmış çalışmaların derlemesi olarak hazırlanmıştır.
... Earlier hominins, as well as Upper Paleolithic humans, are not known, with a few possible exceptions. Chapter 2 (Roksandic 2016) presents the fossil record from the Central Balkans, highlighting the recent fossil human find from Mala Balanica. Roksandic puts forth the possibility for an alternative course for human evolution in this part of Europe, different from the one proposed by the accretion hypothesis for the Western part of the continent. ...
... At Pešturina several fragmentary skeletal and dental remains were recovered from two layers, representing modern humans and Neanderthals, respectively (Roksandic et al., 2017a). Other possible Pleistocene humans have been refuted or questioned based on direct dating, morphological analyses, and review of archival records (Roksandic, 2017;Roksandic et al., 2014). In the greater region of the Balkans peninsula, Neanderthal fossils from Vindija, Croatia, have been radiocarbon dated in a number of studies, with increasingly rigorous pretreatment methods (Green et al., 2010;Higham et al., 2006;Krings et al., 2000). ...
Article
The Central Balkans, in present-day Serbia, was a potentially dynamic zone during the Middle and Upper Paleolithic (MP-UP), as it is situated between hypothesized dispersal routes of modern humans and refuges of late Neanderthals. However, the population history of the region remains poorly understood because there are little chronometric data from Late Pleistocene sites in Serbia. Here, we review the existing paleoanthropological record for the MP-UP in the Central Balkans and surrounding areas. Then, we add to it by reporting radiocarbon dates from two Serbian cave sites, Pešturina and Hadži Prodanova, which contain Middle Paleolithic and Gravettian assemblages. The results provide reliable human occurrence-dates older than 39 ka calibrated radiocarbon years before present (cal BP) and between 34 and 28 ka cal BP. As shallow palimpsest deposits with low artifact yields, the sites are not ideal contexts for establishing chrono-cultural stratigraphy. However, it is proposed that the occupants before 39 ka cal BP were Neanderthals producing MP artifacts, while those after 34 ka cal BP were modern humans with Gravettian traditions.
... Along with the recent finds of fossil hominins (e.g. Roksandic et al., 2011;Roksandic, 2016;Radovi c et al., 2019), the papionin teeth described in this paper add to the growing record of fossil primates from this country. The discovery confirms that primates were present in this region during the latest part of the Neogene period (i.e. ...
... The past decade has seen an increased interest in the paleoanthropology of the Balkan Peninsula (Harvati and Roksandic, 2016, and chapters therein;Tourloukis and Harvati, 2018). Sitting at the "crossroads of Europe" (Roksandic, 2016), the region is marked by geographical migration corridors that facilitated human and animal movement between Europe and the Levant. It also served as one of three major glacial refugia (Hewitt, 1999;Griffiths et al., 2004;Denell et al., 2011;Roksandic et al., 2018), but in contrast to the Iberian and Apennine Peninsulas, the Balkan refugium was never geographically isolated from the larger hominin "source" population in Asia. ...
Article
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The Central Balkans represents a significant geographical gap in the human fossil record of Eurasia. Here we present two new human fossils from Pešturina Cave, Serbia: a partial atlas vertebra (C1) and a fragment of radial diaphysis. The atlas (Pes-1) derives from the lower portion of Layer 2 and conforms to modern human morphology. This layer is characterized by Gravettian industry despite uncertainties caused by bioturbation and difficulties in separating Layers 2 and 3. The radial fragment (Pes-2) was recovered from the contact zone between Layers 3 and 4, both of which represent Mousterian industries, and is tentatively assessed as Neanderthal based on morphology. With the recent publication of a Neanderthal molar (Pes-3) from the same site, Pešturina currently stands as the only Neanderthal fossil-bearing site in Serbia. These additional finds make Pešturina Cave the only site in the Central Balkans which preserves both modern human and Neanderthal remains with associated lithic industries and highlights the importance of Pešturina in the current discourse on hominin dispersals and migrations in the Balkans.
... Furthermore, the "Eurocentric point of view" in the discussion about the initial colonisation of Europe and the migratory routes, is being challenged (Dinçer 2016: 213). The Balkans are currently viewed as a core demographic area, sustaining populations even during the glacial periods and permitting continuous movement and communication via multiple routes between southeast Europe and southwest Asia throughout the Pleistocene (Roksandic 2016;Dobos and Iovita 2016;Ivanova 2016;Strait 2016), while the perception of Anatolia as a land bridge connecting Asia and Europe in an East to West axis is proving to be oversimplified and needs to be reconsidered. Climatic and topographic variability over this vast area and geographical/natural barriers must have posed serious challenges for early hominins during the Early and Middle Pleistocene (Kuhn 2010a;Dinçer 2016). ...
Thesis
This thesis explores possibilities for hominin movement and occupation over the exposed dry land landscapes of the Aegean region during the Early and Middle Pleistocene (focusing more on the Middle Pleistocene ca. 0.8- 0.2 Mya). The point of departure and inspiration is the recent palaeogeographical reconstructions from the study area. Geological evidence reveals the existence of extended terrestrial landscapes, with attractive environments, connecting western Anatolia to Europe via the Greek mainland, during the glacial lowstands of the Middle Pleistocene, and possibly during certain interglacials. These lands are now lost, lying underwater, but, in spatial terms, a completely new spectrum of possibilities opens up for hominins moving across or settling over this part of Eurasia, affecting the wider narrative regarding the early settlements out of Africa. Yet, the research potential of the submerged landscapes of the Aegean has not been fully integrated in the way(s) we study and interpret the Lower Palaeolithic evidence from this region. The discussion about the early colonisation of Europe has been long focused on the western part of the continent due to the abundance of available evidence. The wider Aegean region was excluded, until recently, as a ‘cul de sac’ that blocked movement and dispersal towards the west, representing a gap in the European Lower Palaeolithic archive, with very little to contribute in terms of material culture or hominin fossil evidence. Advances in palaeogeography and geoarchaeology and exciting new finds urging now for a reconsideration. Could the Aegean exposed lands provide land bridges for movement and favourable niches for occupation, offering perhaps an eastern gateway to Europe during the Early and Middle Pleistocene? In order to answer these questions I drew information from archaeology and palaeoanthropology, palaeozoology and palaeoenvironments, and geology and palaeogeography. These multiple lines of evidence have been synthesised within an affordance-based GIS framework, which centres on the relationship between the hominins and their ‘affording’ world. The new methodological scheme developed here led to new hypotheses and scenarios of movement and occupation, predicting areas in the Aegean, onshore and offshore, with increased research potential for the Lower Palaeolithic, based on the level of suitability for the hominin survival, subsistence and dispersal. The findings of my study suggest that despite the serious methodological challenges imposed by landscape dynamics, temporal limitations and extensive discontinuities in the archaeological record, a cross - and inter - disciplinary approach can help us gain valuable insights into the nature of the past landscapes and land use by hominins. In this respect, the complex topography concept and the concept of affordances constitute the backbone of my approach. The first, by setting out the background against which suitability was built, and the second, by attributing a lived and experienced element into the past landscape. The contribution of this study is twofold: (a) offers a framing heuristic, to the newly founded discipline of the continental shelf prehistoric research, for testing further ideas on hominin movement and occupation in dynamic environments; and (b) proposes trans-Aegean corridors of opportunity for dispersal and occupation areas, complementing the current Lower Palaeolithic narrative with a potential eastern gateway to Europe.
... Hominin fossil finds from Turkey can therefore help answer questions about hominin dispersals, both for the dispersal event of early Homo species, as well as for the later Out of Africa migration of modern humans (see also Dinçer 2016). Despite its critical geographic position-and similarly to other neighboring countries (see Harvati et al. 2009;Harvati 2016;Roksandic 2016;Strait et al. 2016)-paleoanthropological research in Turkey has been limited, and the known fossil human record from this region is small. Turkey has substantial potential for Paleolithic sites, and the poor fossil record is most likely a consequence of the scarcity of excavations and surveys. ...
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The timing and route of early human dispersals out of the African continent are among the most important issues currently discussed in paleoanthropology. Several questions arise concerning both early and later dispersals: When did migration events happen? From which populations did these dispersing hominins stem? Which routes did they use? One of the likely dispersal corridors passes through Turkey, which is situated between three continents and therefore can be seen as an important bridge between them. Despite its geographic position, paleoanthropological research in Turkey has been limited, and the known fossil human record from this region is small. Although most of the known fossil human remains were found during early investigations, in the last decade new finds have further highlighted the region’s potential for paleoanthropological research. This chapter reviews the human fossil record from Turkey, and presents the results of a preliminary geometric morphometric study of the Kocabaş hominin, the oldest and most important fossil human specimen known from the country.
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In recent years, an increasing number of discoveries have supported the idea that human occupation of Europe took place earlier than expected, during the Villafranchian and significantly predating 1 Ma. Two hypotheses of dispersal toward Europe seem possible: (1) A direct dispersal from Africa with the earliest possible time frame being ca. 2.0–1.95 Ma; (2) A more recent dispersal, possibly from secondary nuclei of speciation in Asia Minor-Caucasus. The earliest “well documented” wave of Homo dispersal is probably related to the late Villafranchian/Epivillafranchian boundary, at ca. 1.3–1.2 Ma. Two routes of dispersal were possible: via the Bosphorus, or by a circuitous route around the Black Sea basin along the northern peri-Pontic coast. The time of the earliest human appearance in Europe could be related to conditions of increasing aridification and to a domination of open/mosaic landscapes, which roughly correspond to the ecological conditions experienced by African early Homo. The early Homo populations in Europe were likely not adapted to harsh climates and may have occupied only the southern-most areas of the continent.
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In this chapter, we discuss Croatian sites that have yielded human skeletal remains from the Pleistocene. These include the well-known Neandertal localities Hušnjakovo (at Krapina) and Vindija cave, as well as the Late Upper Paleolithic hominin fossil site Šandalja II cave in Istria. The Krapina site played an important role in the historical development of paleoanthropology and is still the Neandertal site with the largest known minimum number of skeletal individuals to date. Finds from Vindija cave belong to one of the latest Neandertal groups in Europe and provide data for the study of both their behavioral, as well as biological characteristics (including genomics studies). The Šandalja II cave in Istria is the only site in Croatia with direct association of human skeletal finds and the late Paleolithic, an Epigravettian industry, providing us with data on the anatomy and behavior of the Late Paleolithic inhabitants of this region.
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Recent archaeological investigations have enabled preliminary insight into the Lower to Middle Paleolithic transition in the Central Balkans. Industries containing tools made from pebbles and flakes, within which Levallois artifacts were present to a lesser (Kosovska Kosa) or greater (Samaila) extent, have been encountered at the sites in the Zapadna Morava valley. The Charentian, likely dating to the Middle Pleistocene (possibly MIS 7) on the basis of microfaunal remains, has been reported in Velika and Mala Balanica in Sićevo. With regard to later (MIS 5–4) industries, assemblages of Typical Mousterian (Crvena Stijena, Hadži Prodanova cave), Charentian (Pešturina) and assemblages where Taubachian–Charentian component, Charentian elements, and backed bifaces are combined (Petrovaradin fortress) are encountered in the Central Balkans. After examining all available data, we propose the hypothesis that in addition to climatic, ecological, and behavioral factors, demographic factors also probably had considerable impact on the variability of lithic assemblages. Migrations and cultural transmission could have resulted in the appearance of Near Eastern elements in the Central Balkans as well as Balkan elements in the Near East. The homogeneity and/or variability of industries could be considerably influenced by the degree of isolation of human groups living in this region.
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The timing and route of early human dispersals out of the African continent are among the most important issues currently discussed in paleoanthropology. Several questions arise concerning both early and later dispersals: When did migration events happen? From which populations did these dispersing hominins stem? Which routes did they use? One of the likely dispersal corridors passes through Turkey, which is situated between three continents and therefore can be seen as an important bridge between them. Despite its geographic position, paleoanthropological research in Turkey has been limited, and the known fossil human record from this region is small. Although most of the known fossil human remains were found during early investigations, in the last decade new finds have further highlighted the region’s potential for paleoanthropological research. This chapter reviews the human fossil record from Turkey, and presents the results of a preliminary geometric morphometric study of the Kocabaş hominin, the oldest and most important fossil human specimen known from the country.
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Southeastern Europe represents a key area in investigating hominin dispersals during the Pleistocene. However, the understanding of these phenomena is hampered by the scarcity of data, especially for the Lower and Middle Pleistocene. The discoveries from Romania assigned to these periods (either credited as in situ or from disturbed contexts) are rather doubtful. After reviewing the state of the art, our paper presents the site of Dealul Guran, discovered in 2010 during a systematic survey carried out in the province of Dobrogea, southeastern Romania. The site is a collapsed rockshelter located on a limestone hill, very rich in flint nodules. Three archaeological layers were identified, and absolute ages indicate that the two oldest archaeological units correspond to an OIS 11 occupation of the site. The assemblages consist mostly of cortical flakes and there are many tested blocks from these units, likely reflecting flint quarrying activities.
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The location of Bulgaria on the Balkan Peninsula makes it potentially important for evaluating biogeographic hypotheses related to human evolution. The country lies at the crossroads of Europe and Asia Minor and constitutes a key portion of one of the possible dispersal pathways that hominin populations would have employed as they entered and left Europe during the Pleistocene. Unfortunately, the Pleistocene human fossil record of Bulgaria is sparse, and perhaps more importantly, the specific biogeographic hypotheses that human fossil discoveries might address could be more fully articulated. In this chapter, we review the fossil hominins currently known from Bulgaria and discuss the framing of biogeographic hypotheses.
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Dental anthropologists focus on the variation around a commonly shared pattern, a variation expressed by differences in tooth size and morphology. This book centers on the morphological characteristics of tooth crowns and roots that are either present or absent in any given individual and that vary in frequency among populations. These nonmetric dental traits are controlled largely by genetic factors and provide a direct link between extinct and extant populations. The book illustrates more than thirty tooth crown and root traits and reviews their biological and genetic underpinnings. From a database of more than 30,000 individuals, the geographic variation of twenty-two crown and root traits is graphically portrayed. A global analysis of tooth morphology shows both points of agreement and disagreement with comparable analyses of genetic and craniometric data. These findings are relevant to the hotly contested issue of timing and geographic context of modern human origins.
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Bones and teeth represent an exceptionally valuable and useful source of information for bioarchaeological research, since they may remain unaltered for a long period of time. The morphological characteristics of teeth are often used to explain certain historical problems which are the focus of interest in anthropological research. Purpose: the aim of this study was to compare morphological particularities of human molar crown traits in Croatian medieval and contemporary populations. Materials and methods: This research was conducted on 252 human molars, both of archeological (172) and of recent (80) origin. Upper molars were studied to determine the frequency of occurrence of: metaconus, hypoconus, metaconulus, parastyle and Carabelli's trait. Lower molars were studied to register the frequency of occurrence of: anterior fovea, mid trigonid crest, groove pattern, deflecting wrinkle, molar cusp number, protostylid, distal accessory ridge, cusp 5, cusp 6 and cusp 7. Each morphological feature's degree of expression was classified according to Arizona State University Dental Anthropological System. Statistically significant differences were established through analysis and comparison of these teeth of archeological and recent origin. Results: on the upper molars there were differences in the frequency of the occurrence of the hypoconus and metaconulus. On the lower molars there were differences in the frequency of the occurrence of the, anterior fovea and 5th cusp. Conclusion: these differences can't be simply explained by wear, since the teeth selected for research all displayed a low degree of abrasion. Additional explanation should be sought among available historical and archeological data, since the medieval period was marked by numerous wars and population migration.
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Palaeogeographic and climatic changes in the Eastern Mediterranean during the Neogene/Quatemary led to extended mammalian migrations and faunal exchanges between Eurasia and Africa. At the same time, the Beringian landbridge was activated several times, and American faunal elements entered Eurasia. It appears that the main factor affecting migration potential and faunal changes/exchanges during the Neogene was palaeogeography, while after the early Pliocene migrations were mainly controlled by climatic changes. Several mammalian migrations can be distinguished, but the most important was that of the middle Orleanian at about 17.0-18.0 Ma when Africa and Eurasia were connected after a long separation and a great number of African faunal elements entered Eurasia and vice versa. Some more important faunal changes also occurred: 1. at ∼5.5 Ma, marking the beginning of the Pliocene, 2. at ∼2.0-1.8 Ma, marking the beginning of the Pleistocene, and 3. at ∼1.0 Ma, defining the early/middle Pleistocene boundary and the establishment of modem mammal fauna. During the Pleistocene, oscillation of glacial and inter-glacial periods caused an alternation of cold-steppic faunas with temperate ones in the Eastern Mediterranean. Endemic late Pleistocene mammalian faunas developed in the Mediterranean islands after their isolation ; "dwarf" elephants, cervids and hippos occurred, as well as giant rodents.
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Two views prevail concerning the significance of H. heidelbergensis in Middle Pleistocene human evolution. H. heidelbergensis sensu stricto refers to a European chronospecies of H. neanderthalensis while H. heidelbergensis sensu lato is considered to be an Afro-European species ancestral to modern humans and Neandertals. Here, we test the phylogenetic validity of H. heidelbergensis using a cladistic analysis based on cranial morphological data of Pleistocene fossils. We perform a low-level analysis to ascertain the information content of the morphological features, a high-level analysis with reweighted characters resulting in a single most parsimonious cladogram and a bootstrap analysis to assess the robustness of this cladogram. Our results show that (i) the identification of a coherent H. heidelbergensis s.l. species is not well supported and is equivocal; (ii) the hypothetical last common ancestor of H. sapiens and H. neanderthalensis has more affinities with African specimens than European; (iii) two Middle Pleistocene European fossils (Atapuerca SH5 and Steinheim) should be classified as H. neanderthalensis.
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In 1925 Josef Szombathy (1853-1943) published a full, and for its time, relatively complete account of the excavations, geology, paleontology, archaeology, and anthropology of the Mladeč Caves1. It is unclear how Szombathy learned of Mladeč but in 1881 and 1882 he was commissioned by the Vienna Academy of Sciences to conduct exploratory research in the caves (Fig. 1.). The property was then owned by Prince Johann von and zu Liechtenstein, who as Szombathy commented, provided some "meager" financial support to run the excavations. The days Szombathy spent there were devoted to mapping the Main Cave and putting in test excavations, primarily in an area Szombathy called the "Dome of the Dead." As luck would have it, his excavations, though intended to be preliminary, produced major collections of human remains and prehistoric artifacts. Szombathy identified the locus of some discoveries, made a sketch of the vertical stratigraphy, and saved a great deal of the excavated material. After completing his work at the Main Cave in the late 19th century, all of the human remains and archaeological materials, and all of the faunal materials were brought to the Naturhistorisches Museum Wien, where they still reside. Szombathy returned to Mladeč twice more, once in 1904 to study some of the new discoveries by Knies and Smyčka in the Quarry Cave and again in 1925 to examine new specimens excavated from the Main Cave. On the last trip, it seems he was expecting to have this material transferred to him in Vienna, but he was only allowed to study them in Litovel. He wrote2 "it was impossible for me, however, to undertake intensive investigation [⋯] because I was equipped only with my traveling tool kit" (1925, 73).
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Systematic excavations of the Mala Balanica cave, Sicevo, (Serbia) yielded a left semi mandible: BH-1, the only specimen from the Balkan Peninsula securely dated to the Middle Pleistocene. The primitive morphology of the mandibular body and the lack of derived Neandertal traits place this specimen outside the variation of Middle Pleistocene European hominins. The specimen’s primitive morphology is more consistent with the new radiometric age estimate that places it into the earlier part of the Middle Pleistocene (Rink et al. 2013). Here I examine the significance of this specimen for the role that the Balkan Peninsula – the only refugium that never experienced isolation – could have played in maintaining gene flow and allowing primitive traits to remain present in the population for a longer period of time.
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The countries of the Balkans and the eastern Mediterranean, located at the junction of three continental regions, are located in a climatically diverse region that has had a profound effect on the development of the fauna and flora. Zolitschka et al (2000) highlight the importance of palaeoecological research in this region across three broad fronts; these being the potential of obtaining very long records of environmental change from basins that have not been over-ridden by extensive glaciations (unlike northern Europe); the fact that the region is a `frontier zone’ where the tropical (monsoonal) climatic system of northern Africa meets and interacts with the North Atlantic climatic system, and the long history of human occupation and civilisation in this region.
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By applying a fieldwork-based, geoarchaeological approach, Tourloukis examines in this study the evidence from Greece within the framework of the earliest occupation of Europe. Although the Greek Peninsula lies within a core area of early hominin movements between Africa and Europe but also within Eurasia itself, the Lower Palaeolithic record of Greece remains as yet extremely poor. Choosing the scanty Greek record as a case-study, Tourloukis elaborates on a hitherto largely overlooked subject in the Eurasian Early-Middle Pleistocene archaeology: the role of geomorphic processes in biasing archaeological distribution patterns of early human presence. This study identifies the current status of the record, explains this status from a geoarchaeological and geomorphological perspective, and prospects its future enrichment. The analysis is carried-out on a landscape-scale and it assesses preservation potential in conjunction with archaeological visibility. Finally a conceptual model emerges that can assist in interpreting and/or predicting early Palaeolithic site locations in tectonically active settings, such as those of the Mediterranean Basin.
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With the exact nature of the interactions between Neanderthals and anatomically modern humans, and the identity of the of the bearers of early Upper Paleolithic technology still open questions essential to expand the human fossil data of Southeast Europe. In our attempt to do so, we investigated a small collection of six previously unpublished human cranial fragments from Serbia, housed at the Natural History Museum in Belgrade and the National Museum in Kraljevo. Tenuous contextual evidence suggested a possible Pleistocene age for the specimens. We conducted a macro-morphological analysis and accelerator mass spectrometry radiocarbon dating in order to assess taxonomical positions and absolute dates for the specimens. Thorough prescreening and chemical characterization of bone samples were used to ensure high reliability of 14C dates. Although the results showed all specimens to be Holocene-aged anatomically modern humans, this should not discourage future research. On the contrary, if indeed we want to understand the early presence of modern humans in the Central Balkans, more research is needed. This includes further checking of old museum collections, but emphasis will need to be placed on new excavations of Pleistocene sites in the region.
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Seventeen Middle Pleistocene crania from the Sima de los Huesos site (Atapuerca, Spain) are analyzed, including seven new specimens. This sample makes it possible to thoroughly characterize a Middle Pleistocene hominin paleodeme and to address hypotheses about the origin and evolution of the Neandertals. Using a variety of techniques, the hominin-bearing layer could be reassigned to a period around 430,000 years ago. The sample shows a consistent morphological pattern with derived Neandertal features present in the face and anterior vault, many of which are related to the masticatory apparatus. This suggests that facial modification was the first step in the evolution of the Neandertal lineage, pointing to a mosaic pattern of evolution, with different anatomical and functional modules evolving at different rates.
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The application of microtomography (mCT) to dental morphological studies has unveiled a new source of palaeobiological information, particularly in the analysis of the internal structures of teeth. In this study, we assess the expression of talonid crests at the enamel and dentine surfaces in lower permanent and second deciduous molars (M2 and dm2) of H. sapiens, H. neanderthalensis and Atapuerca-Sima de los Huesos (SH) hominins. In modern humans, talonid crests are described exclusively in the deciduous teeth (Korenhof, 1982) and interpreted as a primitive mammalian remnant of the talonid attachment to the trigonid. Here we report for the first time the expression of talonid crests of deciduous and permanent molars in H. sapiens, H. neanderthalensis and Middle Pleistocene hominins. We discuss possible evolutionary interpretations and suggest the importance of recording this feature in future studies.
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Trigonid crest patterning in lower molars is distinctive among Late Pleistocene hominins such as Homo neanderthalensis, fossil Homo sapiens and modern humans. In this paper, we present an examination of trigonid crest patterning in the Middle Pleistocene permanent lower molar sample (n=62) of Homo heidelbergensis from Sima de los Huesos (SH). Crest expression was assessed from 3D models of the enamel and the dentine surfaces that were produced using micro-computed tomography (microCT). The aims of our analysis are to: 1) characterize the pattern of trigonid crest expression at the outer enamel and enamel-dentine junction surfaces (OES and EDJ) of the SH sample, 2) evaluate the concordance of expression between both surfaces, and 3) place trigonid crest variation in the SH sample into a phylogenetic context. Our results reveal a greater variability in the expression of trigonid crests at the EDJ (14 types) compared to the OES (4 types). Despite this variability, in almost all cases the expression of a continuous mid-trigonid or distal crest at the OES corresponds with the expression of a continuous mesial/mid-trigonid or distal trigonid crest, respectively, at the EDJ. Thus, it is possible to predict the type of trigonid crest pattern that would be at the OES in the case of partially worn teeth. Our study points to increased variability in trigonid crest expression in M3s compared to M1s and M2s. Moreover, our analysis reveals that the SH sample matches broadly the trigonid crest patterns displayed by H.neanderthalensis and differs from those exhibited by H.sapiens, particularly in the almost constant expression of a continuous middle trigonid crest at the EDJ. However, SH hominins also exhibit patterns that have not been reported in H.neanderthalensis and H.sapiens samples. Other aspects of the variability of the trigonid crest expression at the dentine are presented and discussed.
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The site of Dmanisi, Georgia, has yielded an impressive sample of hominid cranial and postcranial remains, documenting the presence of Homo outside Africa around 1.8 million years ago. Here we report on a new cranium from Dmanisi (D4500) that, together with its mandible (D2600), represents the world's first completely preserved adult hominid skull from the early Pleistocene. D4500/D2600 combines a small braincase (546 cubic centimeters) with a large prognathic face and exhibits close morphological affinities with the earliest known Homo fossils from Africa. The Dmanisi sample, which now comprises five crania, provides direct evidence for wide morphological variation within and among early Homo paleodemes. This implies the existence of a single evolving lineage of early Homo, with phylogeographic continuity across continents.
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Ce travail decrit les restes humains de la grotte ď Et-Tin (Israel). Ces ossements humains sont dates du Neolithique Levantin ancien ou de la fin de l'Epipaleolithique. Ces restes humains du Wadi Et-Tin compares a des pieces natoufiennes presentent en regle generale des traits et des dimensions semblables a ceux des fossiles natoufiens.
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The Visogliano shelter, in north-eastern Italy, is an important Middle Pleistocene occupation site where human remains were found together with an archaic lithic industry, including choppers, chopping tools and a few protobifaces. It is of utmost importance to try to document this period, when a second wave of settlement colonised Western Europe, carrying new flaking techniques and tools.Combined ESR/U-series analyses, integrated with biostratigraphical and environmental data, define a chronological frame for the layers from which the artefacts were unearthed. The lower levels, including human remains, can be dated to the 350–500kyr time span, in agreement with micromammal and stratigraphical studies.These data make Visogliano one of the oldest palaeoanthropological sites in Italy, where human remains are directly associated with protobifaces, choppers and chopping tools. In Western Europe, Visogliano is contemporaneous to the G soil of the Arago Cave, France, with which it shares several similarities in faunal assemblages and radiometric data, and which contains human remains also. These data make Visogliano as one of the oldest sites in Europe where the Acheulian culture is observed.
Book
Teeth are one of the best sources of evidence for both identification and studies of demography, biological relationships and health in ancient human communities. This text introduces the complex biology of teeth and provides a practical guide to the: • excavation, cleaning, storage and recording of dental remains • identification of human teeth including those in a worn or fragmentary state • methods for studying variation in tooth morphology • study of microscopic internal and external structure of dental tissues, and methods of age-determination • estimation of age-at-death from dental development, tooth wear and dental histology • recording of dental disease in archaeological and museum collections Dental Anthropology is the text for students and researchers in anthropology and archaeology, together with others interested in dental remains from archaeological sites, museum collections or forensic cases.
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Balkan Biodiversity is the first attempt to synthesise our current understanding of biodiversity in the great European hot spot. The conservation of biodiversity is one of today’s great ecological challenges but Balkan biodiversity is still poorly understood, in a region with complex physical geography and a long history of political conflict. The Balkans exhibit outstanding levels of endemism, particularly in caves and ancient lakes such as Ohrid; lying at the crossroads of Europe and Asia they are also renowned as a focus of Pleistocene glacial refugia. This volume unites a diverse group of international researchers for the first time. Its interdisciplinary approach gives a broad perspective on biodiversity at the level of the gene, species and ecosystem, including contributions on temporal change. Biological groups include plants, mammals, spiders and humans, cave-dwelling organisms, fish, aquatic invertebrates and algae. The book should be read by zoologists, botanists, speleobiologists, palaeoecologists, palaeolimnologists and environmental scientists.
Chapter
Extensive fieldwork and detailed studies during the last three decades have enriched our understanding of the Plio-Pleistocene large mammal record of Greece. While the unearthed material is abundant, it is not evenly distributed throughout the Plio-Pleistocene; therefore, there are time intervals in this period for which the known large mammal fauna is limited and our knowledge is poor. The Greek Plio-Pleistocene large mammal record reveals a paleoenvironmental transition from open woodlands in late Pliocene, to savannah-like landscape during the early Pleistocene, and to open grasslands during the late early Pleistocene. During this environmental shift, several taxa arrived in Greece in their westward expansion, whereas others made their last European appearance. The arrival of Homo in Europe is discussed in relation to the Greek faunal record. The available data cannot clearly distinguish between an African or an Asian origin, but the latter is supported by more evidence.
Chapter
Greece lies at the crossroads between Europe, Asia, and Africa, and represents a logical gateway through which early human populations might have repeatedly passed on the way to and from Europe. It also represents one of the three European Mediterranean peninsulas which acted as a refugium for fauna, flora and, very likely, human populations during glacial times. Evidence from this region is therefore essential in order to test hypotheses about the course of human evolution in Europe. Despite the importance of the region, paleoanthropological research has until recently been relatively neglected. In recent years, however, renewed research efforts have produced new human fossils from Greece, recovered from excavated contexts. This chapter reviews the Greek human fossil evidence in the context of broader questions in European paleoanthropology.
Chapter
Traces of settlement possibly dating to the Lower Paleolithic have recently been discovered in Bulgaria, including Kozarnika cave and surface sites from the Rhodope Mountains. Chopping tools, cores, flakes, and other stone tools are present in some flint assemblages. In rare cases, bifacial forms have been found. Based on their biostratigraphic position, the assemblages from Kozarnika are estimated to date from the period ranging between 1.6 Ma and 400 ka (Sirakov et al. 2010). This suggested dating is discussed here. The age of the surface sites in the Rhodope Mountains is estimated on the basis of the typo-technological characteristics and the stratigraphic location of the artifacts. The surface sites in the Western Rhodopes may date from the Middle Pleistocene, while the surface sites in the Eastern Rhodopes might be even older. The results of the research on Lower Paleolithic sites in Bulgaria are discussed in the framework of hypothesized repeated waves of dispersal towards Europe.
Article
Late Pleistocene Neandertals, so-called "Cro-Magnons," as well as earlier Homo erectus, had been identified as early as the late 19th century. Primarily, these fossils share some derived characteristics with later humans, in particular a large brain size. Recognizing the identity of the Neandertal lineage has been paramount in the re-interpretation of Middle Pleistocene hominins. Two models have been proposed for the emergence of the Neandertal phenotype. The first model proposes that, before MIS 8 or 7, a quite distinct and stable hominin phenotype assigned to a distinct species (Homo heidelbergensis) can be identified. A second model purports that the emergence of the Neandertal morphology results from an "accretion" of derived features throughout the whole second half of the Middle Pleistocene and possibly starting shortly after the separation of the populations ancestral to Neandertals from their African counterparts.
Article
This chapter discusses Daka hominid material, and presents a detailed ectocranial description of the Daka calvaria (BOU-VP-2/66) and other cranial and mandibular specimens from the Daka Member. It presents a detailed comparative description, tomographic analysis, and systematic interpretation of the cranium. The chapter also presents the postcranial elements in depth, discussing the functional and systematic information they provide. The most important conclusion that can be drawn from the comparison of Daka Homo erectus calvaria to other cranial specimens referred to Homo erectus is that ectocranial features were distributed in a complex mosaic across the Pleistocene within Africa and Eurasia.
Article
Human evolutionary studies exist in their own right due to our own anthropocentricity, but particularly through developments within the natural sciences. The emergence of an autonomous evolutionary biology has largely engulfed human evolutionary studies. Most perspectives of earlier workers are inexplicit in regard to phylogenetic inference, taxonomic resolution, process, pattern, tempo, and other aspects of hominin evolution, and thus are archaic, irrelevant, or both. Matters of epistemology have scarcely merited explicit, critical consideration; even inference as to the best explanation (abduction) has rarely been employed, or employed consistently. Human populations, their genic structure, variability, affinities and histories are now elucidated and directly quantified through molecular biology and population genetics. Past hominin populations are increasingly composed of samples necessary and sufficient to characterize paleo-demes (p-demes) and, ultimately, species clades representative of spatio-temporally bounded entities, the nature and affinities of which are informed through functional, cladistic, and morphometrical investigation. Diverse aspects of earlier hominin habitats, distributions, adaptations, and behavioral parameters are increasingly revealed through multifaceted approaches, all within the framework of paleoanthropology and focused on fuller recovery and elucidation of the Pleistocene archaeological record. Here, some central aspects of Pleistocene hominin evolution are broadly set out from such perspectives. Controversial issues exist, of course, but overall are secondary, in view of the prevalence of normal scientific practice.
Chapter
Every subdiscipline investigating the fossil record has its own holy grail, be they soft body tissues, or ancient DNA, or missing links in human ancestry, the level of its desirability determined not only by the importance in understanding particular aspects of the history of life, but also by the degree of its elusiveness. In the case of European Quaternary palaeobotany the search for cold stage refugia of temperate trees would probably qualify under this category. Ever since the first full-glacial pollen diagrams from southern Europe (e.g Wijmstra, 1969; Florschütz et al 1971) showed steppe-dominated landscapes, implying that forest biomes had not simply shifted en mass southwards as ice sheets expanded, the whereabouts of temperate elements of the flora has been a topic of continuous discussion. The prevailing hypothesis has been that remnant tree populations found refuge in the southern peninsulas of Europe where they survived in suitable microhabitats in mid-altitude zones and in locally moist sites in lowland and coastal areas (e.g Beug, 1968, 1975; Frenzel, 1968, 1979; Lang, 1970; van der Hammen et al 1971; Bennett et al 1991). The presumed small size of such populations and the relative lack of full-glacial evidence means that direct palaeobotanical detection has been difficult, and it is precisely this aspect of the problem that provides the element of fascination in the search for glacial refugia. Indeed, when reviewing the large number of publications on this issue, one is often struck by a tendency (if not a desire) to infer the presence of refugial populations on the basis of sometimes indirect and even tenuous evidence.
Article
While extensive Pleistocene loess deposits have been identified across Eurasia, Holocene age loess (typically nonglaciogenic) is rarely recognized. We explore possible loess deposits in the Mureş River Valley of western Romania, providing a regional signal of increased aridity during the mid‐late Holocene. This proposed aridity may be responsible for the abandonment of Middle Bronze Age tell settlements along the major drainages of the eastern Carpathian Basin (Pannonian plain). This hypothesis centers on a proposed aeolian deposit (the “Pecica deposit”), a ca. 50–80 cm thick, relatively homogeneous, gray layer blanketing the top of the Bronze Age tell of Pecica—Şanţul Mare. Comparing the morphological, geochemical, and physical characteristics of this specific tell deposit with two representative profiles near the site containing glaciogenic calcareous loess and potential Holocene loess deposits developed in Chernozems, we find significant similarities to support this hypothesis. We then review various forms of proxy data published from elsewhere in Central and Eastern Europe suggesting a warming trend during this period. The temporal placement of the Pecica deposit is bracketed using diagnostic artifacts, radiocarbon dates, and the degree of soil development, suggesting a period of increased aridity likely occurring soon after the 17th century B.C.
Article
Establishing a reliable chronology on the extensive hominin remains at Sima de los Huesos is critical for an improved understanding of the complex evolutionary histories and phylogenetic relationships of the European Middle Pleistocene hominin record. In this study, we use a combination of ‘extended-range’ luminescence dating techniques and palaeomagnetism to provide new age constraint on sedimentary infills that are unambiguously associated with the Sima fossil assemblage. Post-infrared-infrared stimulated luminescence (pIR-IR) dating of K-feldspars and thermally transferred optically stimulated luminescence (TT-OSL) dating of individual quartz grains provide weighted mean ages of 433 ± 15 ka (thousands of years) and 416 ± 19 ka, respectively, for allochthonous sedimentary horizons overlying the hominin-bearing clay breccia. The six replicate luminescence ages obtained for this deposit are reproducible and provide a combined minimum age estimate of 427 ± 12 ka for the underlying hominin fossils. Palaeomagnetic directions for the luminescence dated sediment horizon and underlying fossiliferous clays display exclusively normal polarities. These findings are consistent with the luminescence dating results and confirm that the hominin fossil horizon accumulated during the Brunhes Chron, i.e., within the last 780 ka. The new bracketing age constraint for the Sima hominins is in broad agreement with radiometrically dated Homo heidelbergensis fossil sites, such as Mauer and Arago, and suggests that the split of the H. neanderthalensis and H. sapiens lineages took place during the early Middle Pleistocene. More widespread numerical dating of key Early and Middle Pleistocene fossil sites across Europe is needed to test and refine competing models of hominin evolution. The new luminescence chronologies presented in this study demonstrate the versatility of TT-OSL and pIR-IR techniques and the potential role they could play in helping to refine evolutionary histories over Middle Pleistocene timescales.
Article
The debate surrounding the initial appearance of anatomically modern Homo sapiens and subsequent evolution into modern Homo sapiens sapiens had centered on two competing hypotheses. The “Multiregional Continuity Model” and the “Out of Africa 2 Model”. Evidence for these models has primarily come from analyses of pertinent fossilized remains and genetic data. White et al. (2003) suggests that their multivariate analyses of one of three fossil crania, from the Herto formation of the Bouri member in the Middle Awash, Ethiopia, places the fossil intermediate between Archaic Homo sapiens (i.e. Kabwe) and more modern Homo sapiens (i.e. Qafzeh and Skhul V). This, White et al. (2003) suggests, would be strong evidence for the “Out of Africa 2 Model”, albeit in a somewhat gradualistic sense. The purpose of the study undertaken here is to reanalyze the Herto specimen (i.e. BOU-VP-16-1) using c-score (i.e. shape) principal coordinates and Euclidean distances to determine if the Herto cranium is morphometrically intermediate between Archaic Homo sapiens and more modern Homo sapiens sapiens. Our results indicate that indeed these data can be reduced to show that the Herto cranium is relatively similar to our Upper Paleolithic Homo sapiens sample, Skhul V, and our modern Homo sapiens sapiens sample, in that order. Furthermore, our analyses suggest that Herto is not intermediate between Kabwe and more modern Homo sapiens.
Article
Morphometric relationships among late middle and early late Pleistocene fossil hominids from southwestern Asia are examined utilizing multivariate analyses of frontal bones. Particular emphasis is placed on assessing the interaction of size and shape factors in determining relationships among these specimens. Results of these analyses can be summarized in five points. First, techniques which maximize pattern recognition based primarily on shape factors provide the most reliable information pertinent to phylogenetic relationships among these hominids. Second, the Zuttiyeh specimen does not exhibit a greater similarity to the early “modern” Skhu¯l/Qafzeh hominids than to Levantine Neandertals. Third, the Shanidar Neandertals do not cluster closely with Levantine archaic humans. Fourth, the Mousterian-associated Skhu¯l/Qafzeh hominids exhibit distinct similarities to archaic humans, but the Skhu¯l hominids give no unequivocal indication of being “hybrids” between the Qafzeh people and Neandertals. Finally, clear patterns of change occur in frontal bone morphology from the early “modern” (Mousterian-associated) Skhu¯l/Qafzeh hominids to early Upper Paleolithic-associated humans from the Levant.