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Palaeontology and Zooarchaeology of Mezmaiskaya Cave (Northwestern Caucasus, Russia)
Abstract
Over 6000 large mammal and numerous small vertebrate remains have been recovered from preliminary excavations at Mezmaiskaya Cave, situated at 1300–1350 m above sea level in the northwestern foothills of the Caucasus Mountains. Most remains were recovered from layers containing Mousterian artefacts that date to the late Middle Pleniglacial (35,000bpand older). The faunal assemblage reflects a very low degree of weathering, and many bones exhibit traces of stone tool cuts and carnivore modification. Steppe bison (Bison priscus), Caucasian goat (Capra caucasica), and Asiatic mouflon (Ovis orientalis) are the most common large mammals. The remains of reindeer (Rangifer tarandus) were encountered for the first time in the Caucasus. Although most of the smaller vertebrate remains appear to have been accumulated by nonhuman biotic processes (e.g. owl predation), the majority of the ungulate remains probably represent animals hunted by the Mousterian occupants of the cave.
... 1998;Голованова, Дороничев, 2010Дороничева и др., 2013;Несмеянов, 1999;Понс де Леон и др., 2007;Поспелова и др. 2011;Golovanova, Doronichev, 2012;Golovanova et al. 1998;2010;Baryshnikov et al. 1996;Cleghorn, 2006;Doronicheva, Kulkova, Grégoire, 2012 и др.), а остановимся на анализе среднепалеолитического слоя 2В4. ...
... Они также могли приносить в пещеру части убитых ими животных (подробнее см. Baryshnikov, Hoffecker, 1994;Baryshnikov et al., 1996;Cleghorn, 2006). С помощью методики автоматизированного построения микропрофилей был проанализирован характер залегания каменных изделий внутри слоя 2В4 (рис. ...
... Интересно, что и в нижнем, и в верхнем уровнях практически не представлены такие категории костей, как позвонки, кости таза, черепа. Этот факт позволил исследователям палеонтологических материалов предположить, что разделка туш животных происходила на месте забоя (Baryshnikov et al. 1996;Cleghorn, 2006;Golovanova et al., 2005). В целом, как и в нижнем уровне, здесь видна тенденция увеличения количества костного материала вглубь пещеры. ...
... Occasional hunting for young cave bears is also possible, especially during the winter, as suggested in Europe on the basis of isotopic evidence for the diet of the cave lion (Bocherens et al. 2011). During battles with herbivorous adult cave bears, cave lions might be killed and their carcasses then disarticulated in caves by trampling by cave bear and other visitors (Diedrich 2011 (Vereshchagin 1971;Lioubine 1989;Baryshnikov et al. 1996;Hoffecker et al. 2003). ...
... In the Southern Caucasus leopard bone remains are known from many cave localities: Akhstyrskaya Cave in Russia, Bronzovaya Cave, Medvezhiya Cave and Verkhnyaya Cave in Georgia, Azykh Cave (layer 5) and Dashsalakhty in Azerbaijan (Lioubine 1989). In the cave deposits of the Northern Caucasus, leopard remains are rare, known for example in Mezmaiskaya Cave and in Treugolnaya Cave (Baryshnikov et al. 1996;Hoffecker et al. 2003). ...
... Remains of A. jubatus have been identified from Binagady near Baku in Azerbaijan (Vereshchagin 1951). Felids of the Late Pleistocene age are represented in the Northern Caucasus by Panthera pardus and P. spelaea (Vereshchagin 1959;Baryshnikov et al. 1996). The cave lion became extinct at the end of the Pleistocene, while other species survive in the modern Caucasian fauna. ...
The Kudaro Paleolithic site complex in Southern Ossetia includes five species of felids: Panthera onca gombaszoegensis, P. spelaea, P. pardus, Felis silvestris and possibly Lynx lynx. The fossil jaguar P. onca gombaszoegensis was identified from the lowest stratigraphic level of the Middle Pleistocene (Likhvian = Holsteinian Interglacial). Remains of P. pardus and Felis silvestris were recovered from all layers. Panthera spelaea and Lynx lynx represent northern migrants appeared there at the end of the Middle Pleistocene and Late Pleistocene respectively.
... Dans le cas des bisons, on les découpait sur le terrain de chasse, et on apportait dans le site des morceaux de viande. Les carcasses de petits capridés étaient apportées dans la grotte entièrement, ce qui indique la composition complète des parties de leurs squelettes (Baryshnikov et al., 1996). Les sites Ilskaya1 et Ilskaya 2 sont les seuls où on a trouvé des restes de mammouth. ...
... Cet arc traversait la zone fouillée presque en . Les sites sont présentés dans l'ordre chronologique (selon : Baryshnikov et Hoffecker, 1994 ;Baryshnikov et al., 1996). Ratio of bison versus other ungulates represented in MNI per each prey mammal class in the Eastern Micoquian sites in Northwestern Caucasus. ...
... Sites: 1 -Il'skaya 2, layers 5,6; 2 -Il'skaya 2, layers 4a, 4b, 3,2; 3 -Il'skaya 1; 4 -Mezmaiskaya, layers 2B-1-2B-4,3; 5 -Barakaevskaya; 6 -Monasheskaya; 7 -Gubs rockshelter 1; 8 -Mezmaiskaya, layers 2,2A. Statistic data after (Baryshnikov and Hoffecker, 1994;Baryshnikov et al., 1996). ...
A geographical position of the Caucasus in the border between Europe and Asia defines a complex character of the Middle Paleolithic in the region as a whole and in the Northern Caucasus in particular. Today, we can recognize three major cultural areas existed during the Middle Paleolithic in the Northern Caucasus: (1) a local North Caucasian variant of Eastern Micoquian, which is closely related to Eastern Micoquian of Central and Eastern Europe, in the Northwestern Caucasus; (2) a specific Caucasian industry type (called Khostinian Mousterian), which penetrated during later MIS 3 to the Northwestern Caucasus from the Northeastern Black sea cost in the Southern Caucasus; and (3) a similar to Zagros Mousterian industry, which is presented in Weasel Cave in the Northeastern Caucasus.
... The alternate use of cave sites by hominins and carnivores is well documented in a number of sites (Straus ibid, Baryshnikov et al. 1996;Stiner 1999 among others). Spatial modifications made by these agents often make the archaeological context difficult to study and understand . ...
... Occupation of Pleistocene sediments in Eurasian caves by predatory species other than hominins (e.g. wolves (Canis lupus), foxes (Vulpes spp.), spotted hyenas (Crocuta crocuta) and bears (Ursus spp.) is not uncommon (Baryshnikov 1993;Baryshnikov et al. 1996;Stiner 1999). Throughout the course of human evolution, there seems to be a consistent overlap both in the use of space and in foraging strategies employed by hominins and other predators. ...
Short-term settlement of Middle Paleolithic hunters leaves a specific toolkit on an archaeological site. In spite of this well-known fact, in some cases, concerning the duration of stay of groups of Neanderthals, mere techno-typological analysis of lithic assemblages seems insufficient. Analysis of raw material exploitation, combined with information about long use, or reworking of certain artifacts appears to be helpful. On most sites from the Middle Paleolithic, archaeological data concerning the raw material procurement shows that it generally had a local character. However, on a range of sites known from uplands of Hungary, Slovakia, and Poland, artifacts prepared of raw material transported from distant outcrops can be found. Such artifacts are usually reworked, showing traces of their long use and value for prehistoric people. It can be assumed that there exists a link between the settlement duration and the knowledge about the explored area, which can be observed in some lithic assemblages. Situation, where a large variability of regional raw material is represented on a site, can be treated as an indicator of stable settlement, while varied exotic raw materials’ presence documents an increase in group mobility.
... Notwithstanding this previous work, the so-called mountain ungulates, ibex (Capra ibex) and chamois (Rupicapra rupicapra), have been studied poorly in Mousterian sites, so that the hunting of these animals has traditionally been associated with the Upper Paleolithic (Freeman, 1973;Straus, 1987;Gamble, 1995). Nevertheless, evidence from a number of European sites suggests that Neanderthals hunted ibex, chamois and Caucasian tur (Capra caucasica) at certain times during the Middle Paleolithic (Baryshnikov et al., 1996;Valensi, 1996;Tillet, 1997;Moncel et al., 1998;Fiore et al., 2004;Bar-Oz and Adler, 2005;Díez et al., 2008;Adler and Bar-Oz, 2009), yet ibex, chamois and Caucasian tur are not frequent at European Mousterian sites. Furthermore, their presence has usually been attributed to carnivore consumption (Straus, 1982;Blasco, 1995;Geraads, 1997;Davis, 2004;Yravedra, 2011;Mallye et al., 2012). ...
... The remains found in several sites in the Iberian Peninsula such as Esquilleu, Valdegoba, Vanguard Cave, Fuente del Trucho, Covalejos, Ermitia and El Guardia confirm that both animals were hunted by Neanderthals. A similar situation is observed in several regions of Europe (Baryshnikov et al., 1996;Valensi, 1996;Tillet, 1997;Moncel et al., 1998;Fiore et al., 2004;Bar-Oz and Adler, 2005;Díez et al., 2008;Adler and Bar-Oz, 2009). Hence, these species were clearly hunted some time before the Upper Paleolithic. ...
... Материал из Ахштырской пещеры (без разделения на слои) кластируется с материалом из мустьерских слоев 3-4 пещеры Кударо 3 (подвид U. kudarensis kudarensis). Обе выборки хорошо отделены от зубов из ашельского слоя 5 в пещере Кударо 1, которые принадлежат более архаичному подвиду U. kudarensis praekudarensis Baryshnikov, 1996. Эволюционные изменения зубной морфологии спелеарктосов (подрод Spelearctos) связаны с переходом зверей от всеядной диеты, характерной для большинства представителей рода Ursus, к Рис. 6. Строение четвертого нижнего премоляра р4 у кударского пещерного медведя (Ursus kudarensis) из Ахштырской пещеры; лингвальная сторона. ...
... Современное распространение азиатского муфлона на Кавказе ограничено пустынными областями Армении. Вид отсутствует в фауне позднего плейстоцена Кударских пещер, но он был обычен в это время на Северном Кавказе (Мезмайская пещера, Монашеская пещера) наряду с другими видами открытых пространств (Baryshnikov et al. 1996). В холодные эпохи, особенно в стадию максимального похолодания позднего плейстоцена, когда площадь лесных массивов сильно сократилась, муфлоны могли широко расселяться по пологим склонам Северного Кавказа и достигать на юге района Ахштырской пещеры. ...
The Pleistocene fauna of vertebrates from the Akhshtyrskaya Paleolithic cave site comprises nine species of birds and 34 species of mammals, including 5 extinct taxa (Arvicola cf. chosaricus, A. cf. kalmalkensis, Ursus kudarensis, Panthera spelaea, Megaloceros giganteus). Most of fossil bone-fragments of large mammals (over 95%) belong to the Kudaro cave bear (Ursus kudarensis). Cave-bear remains predominantly represent males. The set of skeletal elements suggests the death of bears by natural causes, inside the cave. Traces of damage on bear bones produced by predators are scarce; no unambiguous cut marks from stone tools were revealed. The Akhshtyrskaya Cave presumably was used by cave bears for overwintering, serving from time to time as a shelter for ancient hominins.
... К сожалению, данные, которыми мы располагаем, очень ограничены или недоступны по публикациям. Специальных зооархеологических анализов фауны (Baryshnikov et al, 1996;Golovanova et al, 1999;Cleghorn, 2006) не проводилось на большинстве среднепалеолитических стоянок Кавказа. Работ современного уровня, в которых рассматриваются вопросы экологии и стратегий жизнедеятельности мустьерского неселения Кавказа, пока опубликовано очень мало (Baryshnikov, Hoffecker, 1994;Hoffecker, Baryshnikov, 1998;Hoffecker, Cleghorn, 2000;Adler, 2002). ...
... Отличался и способ доставки добычи на стоянку. Если бизонов разделывали на месте охоты, а на стоянку приносили наиболее мясные части туши, то более мелких козлов/баранов доставляли целиком в пещеру, на что указывает более полная представленность частей скелета последних (Baryshnikov et al, 1996). ...
... In a very real sense the southern Caucasus represents a terra incognita to most Palaeolithic researchers, especially those in the West. Moreover, because of this practical constraint the little that is known about the Middle Palaeolithic of the southern Caucasus is not directly comparable to the vast majority of material collected and studied in neighboring regions such as the northern Caucasus (e.g., Mezmaiskaya Cave: Baryshnikov et al. 1996;Golovanova et al. 1998;Golovanova et al. 1999;Hoffecker 1999;Matouzka Cave: Golovanova et al. 1990;Hoffecker 1999; Barakaevskaïa Cave: Liubin 1984Liubin , 1998Filipov and Liubin 1993;Hoffecker 1999;and Il'skaya I: Hoffecker et al. 1991;Hoffecker 1999); the Levant (e.g., Boker Tachtit: Marks and Friedel 1977;Marks 1983Marks , 1992; Amud Cave: Hovers 1998;Kebara Cave: Meignen and Bar-Yosef 1988;Bar-Yosef et al. 1992;Meignen 1995;andDouara Cave: Akazawa 1979, 1987); the Zagros-Taurus (e.g., Kunji Cave: Baumler and Speth 1993;Lindly 1997;Warwasi Cave: Dibble and Holdaway 1993;Lindly 1997; Shanidar Cave: R. S. Solecki and R. L. Solecki 1993;Lindly 1997;Bisitun Cave: Dibble 1984;Lindly 1997;and Karain Cave: Yalcinkaya et al. 1992;Ceylan 1994;Otte et al. 1998;Otte et al. 1995aOtte et al. , 1995b; or Crimea (e.g., Kabazi and Starosele: see papers in Marks andChabai 1998 andpapers in Chabai andMonigal 1999;Marks and Chabai 2001). Therefore, data from this region is not integrated easily or meaningfully into inter-regional syntheses such as that attempted by Cohen and Stepanchuk (1999). ...
... Tushabramishvili et al. 1999), dominates the faunal assemblage from Ortvale Klde. This species had a wide distribution during the middle and late Pleistocene (Vereshchagin and Baryshnikov 1980) but is generally poorly represented at Middle Palaeolithic sites in the Caucausus (Hoffecker et al. 1991;Baryshnikov and Hoffecker 1994;Baryshnikov et al. 1996). Other species present at the site are listed in table 5. ...
Occupying an intermediate position between Africa, Asia, and Europe the southern Caucasus has represented a northern geographic terminus for major expansions and migrations of human populations, both Archaic and Modern, throughout much of prehistory. During the Middle Palaeolithic, the high elevations and glaciated passes of the Caucasus served as a natural barrier to mobility in a northerly direction. Therefore the southern Caucasus provides an opportunity to examine Neanderthal behavioral patterns within an environmental and geographical cul de sac. Unfortunately, our current understanding of Middle Palaeolithic settlement and subsistence patterns within this region suffers from a dearth of well-excavated, dated, and documented sites. Previous excavations at the rockshelter Ortvale Klde, Djruchula Cave, and Bronze Cave, located in the western Georgian Republic, hint at a variable system of settlement and subsistence linked closely to prevailing environmental and topographical conditions. Although mountainous, warm, humid, and well forested, the numerous deep river valleys that drain the Caucasus form a patchwork of ecological niches populated by a wide array of floral and faunal species. The discontinuous nature of environmental communities and the natural impediments to mobility presented by deep valleys, fast rivers, and high elevations, likely influenced the settlement and subsistence behaviors of Neanderthals more than the cultural factors often cited. Likewise, we argue that climate change fed a cycle of regional abandonment and resettlement, which in turn fostered the technological diversity witnessed in the archaeological record. Traditional views of settlement and subsistence within the southern Caucasus are presented and evaluated in light of data retrieved during the recent re-excavation and dating of Ortvale Klde. Résumé. Occupant une position intermédiaire entre l'Afrique, l'Asie et l'Europe, le Caucase méridional a constitué une barrière géographique pour nombre d'expansions et de migrations de populations humaines, tant archaïques que modernes, au cours de la Préhistoire. Au Paléolithique moyen, l'altitude et l'enneigement des cols ont rendu la chaîne du Caucase infranchissable, de même que les mers, Noire et Caspienne, ont joué le rôle MIDDLE PALAEOLITHIC PATTERNS OF SETTLEMENT AND SUBSISTENCE IN THE SOUTHERN CAUCASUS 93 DANIEL S. ADLER, NICHOLOZ TUSHABRAMISHVILI 92 d'une barrière naturelle vers le Nord. Le Caucase méridional se prête de ce fait à l'étude du comportement néandertalien dans une situation environnementale, géographique et bio-culturelle de cul-de-sac. Une difficulté non négligeable pour notre compréhension des habitats et des modes de subsistance du Paléolithique moyen, au sein de cette région, est due au manque de sites bien fouillés, documentés et datés. La fouille récente de l'abri d'Ortvale Klde et des grottes Djruchula et Bronze, situées à l'Ouest de la République géorgienne, suggère des modes d'occupation et de subsistance variables, étroitement liés aux conditions topographiques et environnementales. Malgré un environnement montagneux, des conditions chaudes et humides et une couverture forestière dense, les nombreuses profondes vallées, qui drainent le Caucase, forment une mosaïque de niches écologiques avec un large spectre d'espèces animales et florales. Nos observations tendent à montrer que cette discontinuité environnementale, à côté des barrières naturelles que constituent les vallées profondes, les cours d'eau torrentiels et l'altitude des cols, davantage que les facteurs culturels souvent cités, ont influencé de façon déterminante le comportement des groupes néandertaliens. Par ailleurs, nous suggérons que l'existence de phases d'expansion et de contraction régionales, peut-être stimulées par des changements climatiques, ont entraîné la diversité technologique observée dans l'inventaire archéologique. Les visions traditionnelles d'occupation et de subsistance pour le Caucase méridional sont présentées et évaluées à la lumière des résultats et datations issus des fouilles récentes d'Ortvale Klde.
... Notwithstanding this previous work, the so-called mountain ungulates, ibex (Capra ibex) and chamois (Rupicapra rupicapra), have been studied poorly in Mousterian sites, so that the hunting of these animals has traditionally been associated with the Upper Paleolithic (Freeman, 1973;Straus, 1987;Gamble, 1995). Nevertheless, evidence from a number of European sites suggests that Neanderthals hunted ibex, chamois and Caucasian tur (Capra caucasica) at certain times during the Middle Paleolithic (Baryshnikov et al., 1996;Valensi, 1996;Tillet, 1997;Moncel et al., 1998;Fiore et al., 2004;Bar-Oz and Adler, 2005;Díez et al., 2008;Adler and Bar-Oz, 2009), yet ibex, chamois and Caucasian tur are not frequent at European Mousterian sites. Furthermore, their presence has usually been attributed to carnivore consumption (Straus, 1982;Blasco, 1995;Geraads, 1997;Davis, 2004;Yravedra, 2011;Mallye et al., 2012). ...
... The remains found in several sites in the Iberian Peninsula such as Esquilleu, Valdegoba, Vanguard Cave, Fuente del Trucho, Covalejos, Ermitia and El Guardia confirm that both animals were hunted by Neanderthals. A similar situation is observed in several regions of Europe (Baryshnikov et al., 1996;Valensi, 1996;Tillet, 1997;Moncel et al., 1998;Fiore et al., 2004;Bar-Oz and Adler, 2005;Díez et al., 2008;Adler and Bar-Oz, 2009). Hence, these species were clearly hunted some time before the Upper Paleolithic. ...
... The ibex and chamois hunting described at El Esquilleu can be recognised in the NISP and MNI results: they share common characteristics with other Upper Palaeolithic deposits that show specialised hunting (Straus 1987(Straus , 1992Delpech & Villa 1993;Gamble 1995;Tagliacozzo & Fiore 2000;Phoca-Cosmetatou 2009), such as Mezmayeskata (Baryshnikov et al. 1996). This hunting strategy is clearly not exclusive to the Upper Palaeolithic, but has earlier origins. ...
... In addition to these specialised strategies (Straus 1987(Straus , 1992Delpech & Villa 1993;Gamble 1995;Tagliacozzo & Fiore 2000;Phoca-Cosmetatou 2009), Neanderthals also exploited a wide spectrum of different taxa at other sites (Scott 1986;Madella et al. 2002;Blasco 2008;Stringer et al. 2008). Early hunting strategies are now also known to have focused on ibex (Baryshnikov et al. 1996), caribou (Gaudzinski & Roebroeks 2000;Rendu et al. 2012), deer (Yravedra 2013) and large bovids (Girad & David 1982;Jaubert & Brugal 1990). The next step is to understand the causes that motivated selection strategy. ...
Traditional views of Neanderthal hunting strategies envisage them preying on herd species such as bison and deer, rather than the sophisticated tracking of solitary animals. Analysis of faunal remains from El Esquilleu Cave in northern Spain, however, demonstrates that during certain periods of the Middle Palaeolithic occupation, Neanderthals focused on the hunting of ibex and chamois, small solitary species that inhabited the mountainous terrain around the site. These results indicate that Neanderthal hunting practices may have had more similarity to those of their Upper Palaeolithic relatives than is usually assumed.
... К сожалению, данные, которыми мы располагаем, очень ограничены или недоступны по публикациям. Специальных зооархеологических анализов фауны (Baryshnikov et al, 1996;Golovanova et al, 1999;Cleghorn, 2006) не проводилось на большинстве среднепалеолитических стоянок Кавказа. Работ современного уровня, в которых рассматриваются вопросы экологии и стратегий жизнедеятельности мустьерского неселения Кавказа, пока опубликовано очень мало (Baryshnikov, Hoffecker, 1994;Hoffecker, Baryshnikov, 1998;Hoffecker, Cleghorn, 2000;Adler, 2002). ...
... Отличался и способ доставки добычи на стоянку. Если бизонов разделывали на месте охоты, а на стоянку приносили наиболее мясные части туши, то более мелких козлов/баранов доставляли целиком в пещеру, на что указывает более полная представленность частей скелета последних (Baryshnikov et al, 1996). ...
... Additionally, an early MP industry with bifacial leaf points was found in Layer 2 at Sredniy Khadjokh (Hadjoh), which was dated preliminarily to MIS 5a-5b based on an IR-OSL date of 87.8±6.8 ka . According to multidisciplinary research at Mezmaiskaya cave (Baryshnikov et al. 1996;Golovanova andDoronichev 2003, 2017;Golovanova et al. 1998;1999;Nesmeyanov Gunz et al. 2012;Hajdinjak et al. 2018;Ponce de Leon et al. 2008;Weaver et al. 2016). Based on the results of radiometric dating, palynological, and geoarchaeological studies in Mezmaiskaya cave, we can identify now three major stages of the Eastern Micoquian Neanderthal occupation of the NWC, which are separated by pronounced cold stages that we identify as periods of environmental stress (see Table 1): • The first stage corresponds to the interval from late MIS 5 through the end of MIS 4, between about 90 to 60 ka. ...
Recent studies of the Middle and Upper Paleolithic in the northwestern Caucasus are focused on the research of relations between natural (climate and environment) and social (behavior and adaptations) factors that governed settlement dynamics of Neanderthal and anatomically modern human populations in the region. The majority of Middle Paleolithic sites in the region show temporal changes within a local variant of the Eastern Micoquian industry between approximately 90 and 40 thousand years (ka) ago. The final stage of the Eastern Micoquian occupation in the northwestern Caucasus is notable in that the number of Neanderthal sites increases, and these sites show a higher variety and spread towards the eastern boundary of the region. The research provides new data indicating that ecology and subsistence of late Neanderthals were affected by a large, catastrophic volcanogenic event, which likely caused the Neanderthal extinction, and that was followed by a subsequent reoccupation of the region by Upper Paleolithic modern humans. In addition, recent genetic analyses indicate that a population turnover is likely to have occurred, either in the Caucasus or throughout Europe, towards the end of Neanderthal history. In the northwestern Caucasus, Upper Paleolithic sites are found mostly in caves or rockshelters, and show two major periods of modern human occupation: (1) Upper Paleolithic, from ~39/38 ka to the onset of the Last Glacial Maximum; and, (2) Epipaleolithic, from the Last Glacial Maximum to ~11/10 ka. The Upper Paleolithic sites are rare, while the Epipaleolithic sites are quite numerous in the region. After the Last Glacial Maximum, milder conditions of the Late Glacial promoted an increase in the number of sites and mobility of the Epipaleo-lithic human groups. A high mobility is confirmed by the facts that similar Epipaleolithic industries are found in the Southern and Northern Caucasus and that the same obsidian sources were exploited in both regions. Results of recent studies indicate that the most crucial factors for hominin settlement during the entire Upper Pleistocene in the northwestern Caucasus were favorable climatic and environmental conditions. In comparison to other regions, including the Levant, the Caucasus' archaeological record shows distinct regional peculiarities and a specific pathway of Upper Paleolithic development, which we identify as the "Caucasus Upper Paleolithic". In support of this view, the results of two recent palaeogenomic analyses of two human individuals from the Southern Caucasus indicate that the first modern humans in the Caucasus shared ancestry with Upper Paleolithic humans of Western Asia, and that the first Upper Paleolithic modern humans in the Caucasus belonged to a distinct ancient clade, which split from the European Upper Paleolithic populations about 45 ka ago, shortly after the expansion of modern humans into Europe.
... The identification of cut marks on bones surface followed with its anatomical location, characteristics of depth and shape of marks, the color (Binford, 1981;Baryshnikov et al., 1996;Burova, 2002). The taphonomical damage on studied bone surface was described according Andrews (1990). ...
The paper is devoted to the role of hares in the composition of hunter's prey in the Upper Palaeolithic of the Russian (East European) Plain. Mammoths and hoofed mammals are traditionally considered as main hunting species for these sites. However, sometimes completely different species come to the fore, while the bones of the “main hunting animals” are presented only in small amounts. We have analyzed the faunistic materials of the sites confined to the basins of the main rivers of the Russian Plain (Dnieper, Desna, Seym, Oka, Don), the chronostratigraphic position of which is justified by radiocarbon dates. As a result of quantitative analysis, the sites were divided into 4 groups, where group 1 comprises the sites with the content of hare bones in relation to all definable bone residues of more than 50%, and group 4 - which does not contain them at all. A set of skeletal elements was also studied and their percentage in the collection was determined. Correlation of the data with the Late Pleistocene event scale showed the absence of a direct relationship between the number of hare bones in sites with climate changes. Throughout the entire Upper Palaeolithic, hares were part of the environment of the ancient population, spreading widely in forest landscapes and decreasing in the tundra-steppes. Hares served as universal game, especially when other types of hunting were limited (for example, during seasonal migrations of other species).
... See Tables 5-9 for separate bones. Calcaneus: GL (Flerov, 1976;Baryshnikov et al., 1996;Á lvarez-Lao and García-García, 2006;Vasiliev, 2008;Croitor, 2010a;Vercoutère and Guérin, 2010;Shpansky et al., 2016). The dimensions of the skulls and other specimens from Kaniv are characteristic for those of the Late Pleistocene steppe bison of Europe and Siberia. ...
Subfossil remains of the Quaternary megafauna from two Ukrainian natural history collections stored at Kaniv Nature Reserve and the National Museum of Natural History NAS of Ukraine were examined. Most of the bones were collected in 1965–1966, in the building pit and the gateway of the Kaniv hydroelectric power plant. The fauna is represented at least by ten taxa (Gulo gulo, Mammuthus trogonterii chosaricus, Mammuthus intermedius, Coelodonta antiquitatis, Megaloceros giganteus, Alces alces, Bison priscus, Cervus elaphus, Capra cf. ibex, and Equus ferus). Skeletal parts are represented mainly by skulls, long bones, horns, and tusks. Based on the species composition of proboscideans, at least part of the faunal assemblage is dated by the end of the Middle Pleistocene (Dnieper Stage = Saale, Warta, MIS 6), but majority of bones could be older or younger (Late Pleistocene and Holocene age) due to the alluvial origin of accumulation.
... ka BP) (Bar-Oz and Adler, 2005), Mezmaiskaya Cave (layers 2b3, 2b4 and 3, 70.6-46.5 ka BP) (Baryshnikov et al., 1996;Golovanova et al., 1998;Skinner et al., 2005;Soubrier et al., 2016), Kudaro 3 (layer 3, >41 600 BP, OxA-19611) (Baryshnikov, 2010). ...
The Middle Pleistocene – Late Pleistocene transition of European large mammal's fauna (Proboscidea, Artiodactyla, Perissodactyla, Carnivora, Hystrix and Castor) assemblages has been studied in 18 European regional faunal assemblages. This study is based on the data yielded from 423 palaeontological sites (758 localities) dated within interval of MIS 6–MIS 4. All the data was aggregated by 9 time intervals (time scale). For ten bioregions, we have been able to obtain descriptive models of evolution of their faunal assemblages. It allowed detecting common rules of changes in large mammals' fauna composition in Europe on the whole as well as changes in the distribution of individual species and their groups within the regions. We have studied the changes in biodiversity parameters (Shannon index, index of self-organization) and Mourelle–Ezcurra species turnover index within MIS 6–MIS 4 time interval. The evolution of European fauna was compared for MIS 6–MIS 5 transition and MIS 2–MIS 1 transition as well as influence of change in global temperature on these transitions was described. In addition, we have showed the correlation between modern species richness with the species richness in MIS 6, MIS 5 and MIS 4 and proposed the hypothesis of historical succession of European bioregions.
... carnivores in different units of Azokh 1 Cave has been indicated by various authors (Huseinov 1985;van der Made et al. 2016 (Baryshnikov 1993;Baryshnikov et al. 1996;Stiner 1999). Throughout the course of human evolution, there seems to be a consistent overlap both in the use of space and in foraging strategies employed by hominins and other predators. ...
The Caucasus is an important intercontinental passageway for fauna and hominin dispersal from Africa to Eurasia. Numerous Pleistocene sites emphasize the importance of this region for the study of human evolution and hominin “Out of Africa” dispersals. The Azokh 1 site in the Southern Caucasus provides a stratigraphic sequence, the renewed excavations of which have shown the presence of well-contextualised lithic and faunal assemblages dated between 300 – 100 ka associated with hominin remains (Homo heidelbergensis and Homo neanderthalensis) also found in the site.
Faunal assemblages are dominated by cave bear (Ursus spelaeus) remains resulting from their hibernation at the rear of the cave. Recent taphonomic studies indicate the in-situ exploitation of some of these remains. Other faunal remains, mainly herbivore, some showing signs of human activity, were most likely introduced into the cave by hominins. The study of lithic artefacts suggests an incomplete operative chain for all raw materials with a general absence of knapping debris, unknapped cobbles/pebbles, rare cores and refits. Techno-typologically these assemblages display characteristics that link them to Late Acheulean or Early Mousterian and Levallois Mousterian traditions.
The faunal and lithic assemblages originate from the rear of the cave. Research results, including some preliminary data on lithic use-wear, along with analyses of spatial distribution and post-depositional modification, indicate that occupation of the cave was short and seasonal in character. Cave bears were an important factor affecting the duration of hominin occupation of the cave. The characteristics of the lithic assemblages suggest mobile toolkits, with some isolated evidence of in-situ knapping and retouching activities.
... carnivores in different units of Azokh 1 Cave has been indicated by various authors (Huseinov 1985;van der Made et al. 2016 (Baryshnikov 1993;Baryshnikov et al. 1996;Stiner 1999). Throughout the course of human evolution, there seems to be a consistent overlap both in the use of space and in foraging strategies employed by hominins and other predators. ...
The Caucasus is an important intercontinental passageway for fauna and hominin dispersal from Africa to Eurasia. Numerous Pleistocene sites emphasise the importance of this region for the study of human evolution and hominin ‘Out of Africa’ dispersals. The Azokh 1 site in the Southern Caucasus provides a stratigraphic sequence, the renewed excavations of which have shown the presence of well-contextualised lithic and faunal assemblages dated between 300 and 100 ka associated with hominin remains (Homo heidelbergensis and Homo neanderthalensis) also found in the site.
... The study of several bone assemblages suggests that bovines were an important source of subsistence for Middle Palaeolithic hunters. Large accumulations of butchered carcasses have been recovered in western-central Europe, at La Quina (Chase, 1999;Rendu and Armand, 2009), Mauran (David et al., 1994;David and Fosse, 1999;Rendu et al., 2012), Coudoulous (Jaubert et al., 2005), La Borde , Biache-Saint-Vaast (Auguste, 1995), Hénin-sur-Cojeul (Marcy et al., 1993), Wallertheim (Gaudzinski, 1995(Gaudzinski, , 1996, Il'skaja (Hoffecker et al., 1991), and Mezmaiskaja Cave (Baryshnikov et al., 1996). A group of these sites provides evidence that Neanderthal groups used favourable landscape topography, such as cliffs, swamps or gorges to drive and trap the herds. ...
In northern Italy, Fumane Cave (Lessini Mountains -Verona), San Bernardino Cave, and De Nadale Cave (Berici Hills - Vicenza) provide data to interpret the exploitation dynamics of the Pleistocene's large bovids Bos primigenius and Bison priscus between 70 and 42 ky BP. Through the taphonomic study of bone assemblages, we have attempted to reconstruct the strategies, methods, and butchery practices in the exploitation of these game animals adopted by different cultural groups of Middle Palaeolithic hunters. Therefore, Neanderthal hunting behaviour has been examined using different proxies such as the choice of anatomical parts, selective transport of elements, prey selection, and age estimation. Results suggest bovines were an important subsistence resource in some cases, even if they were not the most exploited taxa in others, indicating differences and similarities depending on the context across the Italian peninsula. These results highlight additional aspects of Neanderthal landscape use and hunting strategies.
... The mouflon (Ovis orientalis) appears rarely in the faunal spectra of the Upper Pleistocene in the Caucasus. It is present in the Mousterian caves of Barakaevskaya and Mezmaiskaya, on the northwest flank of the Greater Caucasus (Baryshnikov & Hoffecker 1994;Baryshnikov et al. 1996), but does not seem to be in evidence in Georgia (Fig. 7.1). ...
Discovered in 2004 during the planning of a road along the river Barepat, the open air site of Kalavan 1
is located at an altitude of 1630m a.s.l. in the Aregunyats mountain range, which dominates the northern
shore of Lake Sevan in Armenia. In the present state of our knowledge, Kalavan 1 appears to present the
first evidence of human presence in the Lesser Caucasus after the Last Glacial Maximum.
The remains of the identified mammals in good stratigraphic context show a predominance of wild Caprinae
(Ovis sp. /Capra sp.), among which very few could be attributed to Ovis sp. The human groups of Kalavan
1 hunted wild Caprinae which had reached their maximum weight, thus furnishing a large quantity of meat,
as well as other products (hide, bone, tendons, blood, intestines, marrow, etc.).
It is assumed that the species hunted was Ovis orientalis gmelini (‘Armenian mouflon’), which migrates
seasonally between the valleys during the winter and the high meadows during the summer. Hypothetically,
the site of Kalavan 1 could have been occupied at the end of the summer, when the mouflon descended again
to the lower altitudes, and the hunters had already procured obsidian to fabricate their hunting arms from
the neighbouring deposits surrounding Lake Sevan.
... Not only do these assemblages resemble others in eastern Europe and the Levant (Golovanova and Doronichev 2003: 129-134), but the stratigraphic contexts at Mezmaiskaya and the other North Caucasian sites suggest that Neandertals and/or other archaic humans survived in this isolated area into the Upper Paleolithic or even later (Baryshnikov et al. 1996;Hoffecker 2002: 143;Soffer 2001: 236-244 Adler and Tushabramishvili 2004;Meshveliani, et al. 2004), which is several thousand years later than at established sites in Central and ...
This study investigates the genetic diversity and ethnohistory of Svaneti and its neighboring highland Georgian and breakaway regions in order to better understand the complex population history of the South Caucasus. The objectives of this project are to (1) document the biological diversity in contemporary settlements in the region of Svaneti; (2) compare patterns of gene diversity with Svaneti’s western and eastern neighbors, Abkhazia and South Ossetia, respectively; and (3) determine whether gene frequencies in Svaneti are evently distributed across geographic space by characterizing any village-level structuring. We will contextualize the findings within broader studies that address major regional population settlement events during the Upper Paleolithic, Neolithic, and Metal Ages, as well as the putative ‘Alan migration’ in the 4th century AD. To accomplish these goals, biological samples were collected from participants in Svaneti for genetic analysis, providing a more thorough coverage of village districts in Svaneti than has been achieved in previous studies. In addition, local-level ethnohistorical interviews were conducted in an effort to distinguish patterns of diversity resulting from long-term inhabitation versus those arising from recent immigration into the region. These DNA samples were characterized for mitochondrial DNA and Y-chromosome variation, and the resulting data analyzed with statistical and phylogenetic methods to define the biological affinities of highland Georgian populations, and reconstruct the migration and settlement history of the region. Data from published and unpublished sources on the genetic diversity of the greater Near East and Caucasus, specifically Abkhaz and Ossete populations, were used for phylogeographic and statistical comparison. The results revealed reduced Y-chromosome haplogroup diversity in Svans, with a predominance of G2a, although their paternal lineages occurred at frequencies comparable to those of neighboring highland populations. By contrast, mtDNA haplogroup diversity in Svans was both very high and reasonably similar in terms of frequency to other regional populations, with W6 and X2 occurring at unusually high frequencies. Interestingly, there was no geographic patterning of Y-chromosome or mitochondrial DNA diversity within Svaneti at the village level. Nevertheless, strong Y-chromosome affinities with eastern and western populations (i.e., Ossete and Abkhaz, respectively) living adjacent to the Svans indicated a common gene pool for these three ethno-linguistic groups in spite of linguistic differences at the language family level, and minimal contribution to the Ossete gene pool from Indo-European-speaking Alans.
... The recently analyzed fauna from Mezmaiskaya Cave (Cleghorn, 2006) indicates that while the frequency by NISP of Capra/Ovis is not as large as that for Bos/Bison, the very large number of size 2 bovid/cervids and ungulates likely represent sheep and goat. Thus, with only a few exceptions, Caucasian tur is generally poorly represented at most LMP and EUP sites in the Caucasus (e.g., Hoffecker et al., 1991;Baryshnikov and Hoffecker, 1994;Baryshnikov et al., 1996;Hoffecker and Cleghorn, 2000;Hoffecker, 2002). ...
The transition from hunting and gathering nomadic societies to true fanning communities took place during the Levantine Epipaleolithic sequence (20,000-10,000 BP) of Kebaran, Geometric-Kebaran, and Natufian cultures. Starting with the advent of sedentism during the Natufian, a change in demography and intensification of environmental exploitation occurred. We studied the faunal remains from Hefzibah, a large Geometric-Kebaran open-air site on the northern coastal plain of Israel. The Geometric-Kebaran economy and paleoecology are key to understanding the Natufian shift in culture and economy. The bone assemblage comprises 10 mammalian and 2 reptilian species. Bone fractures, burnt bones, and cut marks indicate consumption of most of these species. The major prey species at Hefzibah, as in other Epipaleolithic sites in the area, are mountain gazelle (Gazella gazella, 73%) and fallow deer (Dama mesopotamica, 20%). All body parts are represented at the site. Similar proportions of these species were found in the nearby sites of Nahal Hadera V (Kebaran) and Newe-David (Geometric-Kebaran). Other game species include auroch (Bos primigenius), hartebeest (Alcelaphus buselaphus), roe deer (Capreolus capreolus), wild boar (Sus scrofa), and wild horse (Equus sp.). Small game species include hare (Lepus capensis), tortoise (Testudo graeca), hedgehog (Erinaceus europeus), and fox (Vulpes vulpes).
... The recently analyzed fauna from Mezmaiskaya Cave (Cleghorn, 2006) indicates that while the frequency by NISP of Capra/Ovis is not as large as that for Bos/Bison, the very large number of size 2 bovid/cervids and ungulates likely represent sheep and goat. Thus, with only a few exceptions, Caucasian tur is generally poorly represented at most LMP and EUP sites in the Caucasus (e.g., Hoffecker et al., 1991;Baryshnikov and Hoffecker, 1994;Baryshnikov et al., 1996;Hoffecker and Cleghorn, 2000;Hoffecker, 2002). ...
... According to multidisciplinary research in Mezmaiskaya cave (Baryshnikov et al., 1996;Golovanova et al., 1998a, b;Nesmeyanov, 1999;Golovanova and Doronichev, 2003), hominid occupation has started soon after the cave was completely opened by erosion (in late MIS 5). According to a complex lithological, geomorphological, and paleontological research a relatively mild climate during the deposition of layers 3 and 2B4. ...
Modern data do not show a local evolutionary development of lithic industries throughout the Lower to Middle Paleolithic transition in the Northwestern Caucasus. Human occupation of these territories was not continuous during the Pleistocene. On the contrary, the lacunarity of archaeological record in some periods and a generally cooler climate characteristic for Eastern Europe and the Northwestern Caucasus in late Middle Pleistocene suggest that major climatic deteriorations had a great impact on human settlement of the regions. The human colonization of the Northwestern Caucasus at the interface of the Lower and Middle Paleolithic demonstrates a peculiar dynamics associated with climate cycles of the late Middle Pleistocene – the first half of Upper Pleistocene.
... In the specimens from the early Middle Pleistocene localities of Europe, the value of this index is considerably higher, varying from 53.5% to 56.5% and constituting in average 55.5% (n=7) (Schütt & Hemmer, 1978). Similar index has been obtained by us for the isolated lower carnassial tooth m1 (NHM F66, length 30.5 mm, width 17.Sukachev et al., 1966;Vereshchagin, 1971;Soffer, 1985;Kalinovski & Kuzmina, 1993;Baryshnikov et al., 1996;Averianov et al., 1999;Enloe et al., 2000;Chubur, 2001;Burova, 2002;Sotnikova & Nikolskiy, 2006; and original data). ...
... Few archaeologists nowadays hold to the once fashionable notion that Neanderthals were scavengers, at least of larger ungulates (Auguste 1988(Auguste , 1992Baryshnikov et al 1996;Bocherens et al 2001;Boyle 2000;Burke 2000;Conard & Prindiville 2000;Farizy et al 1994;Gaudzinski 1996Gaudzinski , 2000Gaudzinski & Roebroeks 2000;Grayson & Delpech 1994;Hoffecker & Cleghorn 2000;Jaubert et al 1990;Marean & Kim 1998;Patou 1989;Patou-Mathis 2000;Stiner 1990Stiner , 1991aStiner , 1991bStiner , 1994Stiner , 2004Stiner , 2005 Milo 1997Milo , 1998). ...
We examine the larger mammals from late Middle Palaeolithic Kebara Cave (Israel), and offer eight principal conclusions concerning Neanderthal hunting activities at the site. (1) Regardless of prey size, most procurement was by hunting, not scavenging. The major prey were gazelle and fallow deer, but also aurochs, red deer, and boar. (2) Hunting was seasonal, with most hunts in winter and/or spring. (3) Hunters took male and female deer in similar numbers, but a preponderance of female gazelle. These sex ratios probably reflect local availability and encounter rates. (4) More juvenile deer than juvenile gazelle were taken. The frequency of juveniles has not been severely impacted by taphonomic processes. Because of their small size and limited body fat, juveniles were probably low-ranked resources by comparison to their adult counterparts and may often have been excluded from the hunters' optimal diet. If so, fluctuations in the numbers of juveniles do not track changes in hunting season, but instead indi...
... Introduction e history of bovines in relation to the evolution of Man is, in one sense, a summary of our economic and alimentary history across thousands of years, beginning with the Paleolithic hunters and down to the modern breeder. For the Middle and Upper Paleolithic hunters, bovines were a great source of food, so important in many contexts as it has been evidenced by a large number of archaeozoological assemblages (Baryshnikov et al., 1993;Baryshnikov, 1999, Boscato & Crezzini, 2006, Brugal & Valente, 2007, Cardoso, 1996, Chase, 1999, David et al., 1994, David & Fosse, 1999, Delpech, 1999, Fiore et al., 2004, Gaudzinski, 1995Gaudzinski, 1996, Julien, 2011, Martinez-Moreno, 1999, Minieri et al., 1995, Sala, 1990, Tagliacozzo et al., 2013, Turner, 1999, Turq et al., 1999. e probable techniques of hunting are only a matter of hypothezises, based on faunal data of some important European sites (es. ...
In northern Italy, Grotta di Fumane (VR) Grotta di San Bernardino
Cave (VI) give us important data for the interpretation of exploitation dynamics
Pleistocene’s large bovines, Bos primigenius and Bison priscus in a chronological
period between 50-25 Ky BP. +e archaeozoological study was carried out on the
bovines remains found in the archaeological units: A8, A9, A5, A5+A6, A6 (Musterian
Levallois); A3 e A4 (Uluzzian); A2, A2R, A1, D6, D3+D6 e D3 (Aurignazian);
D1c, D1d, D1e e D1f (Gravettian and Hyenas den) of Fumane Cave and IV and II
(Musterian) of San Bernardino Cave. +rough the study of bone surfaces we have attempted
to reconstruct strategies, methods and practices of butchery adopted by the
di5erent groups of Palaeolithic hunters. +e taphonomic analysis indicates human
being as the main responsible for the accumulation of remains of the large bovines,
and put it in comparison.
... Liubin 1998). By and large, though, the Caucasian goat is poorly represented at Middle Palaeolithic sites in the Caucasus (Hoffecker et al. 1991;Baryshnikov and Hoffecker 1994;Baryshnikov et al. 1996). ...
... Traditionally, the entire contents of sites containing stone tool industries have been attributed exclusively to the actions of humans as the predominant hunters responsible for killing and transporting prey to such sites (e.g. Hoffecker et al., 1991; Baryshnikov & Hoffecker, 1994; Baryshnikov et al., 1996). Over the last couple of decades, a growing appreciation of taphonorny, and of the wide diversity of agents contributing to site formation, has led to reconsideration of interpretations of Mousterian hunting patterns (e.g. ...
Prolom II, a stratified archaeological cave deposit in the eastern Crimean Peninsula, Ukraine, dates back to approximately 135 000–60 000 years ago. Stone tool industries from four human occupation levels are characteristic of the Middle Palaeolithic, typically associated with Neanderthals. In addition to the stone tool artifactual material, there is abundant faunal material, including saiga antelope, horse, bison, hyena and bear. This zooarchaeological investigation examined approximately 3500 specimens of animal bones to discern between natural and cultural modifications, and to elucidate human patterns of exploitation of faunal resources in prehistoric subsistence.
... pointing to the addition of some food items with less negative d 13 C values than the potential prey analyzed in this study. In Late Pleistocene sites from western Europe, such a food item is represented by reindeer Rangifer tarandus (Bocherens et al., 2011a) but in the Late Pleistocene of the Caucasus, this species is almost completely missing, with only a handful of bones from this species found in Mezmaiskaya Cave in Northern Caucasus (Baryshnikov et al., 1996). ...
Bone fragments of large anadromous salmon in the Middle Palaeolithic archaeological layers of Kudaro 3 cave (Caucasus) suggested fish consumption by archaic Hominins, such as Neandertals. However, large carnivores such as Asiatic cave bears (Ursus kudarensis) and cave lions (Panthera spelaea) were also found in the cave and could have been responsible for such an accumulation. The diet of these carnivores was evaluated using carbon, nitrogen and sulphur isotopes in faunal bone collagen. The results suggest that anadromous fish were neither part of the diet of either cave bear (vegetarian) or cave lion (predators of herbivores from arid areas) and therefore provide indirect support to the idea that Middle Palaeolithic Hominins, probably Neandertals, were able to consume fish when it was available.
... This economic focus on Caucasian tur differs from the pattern we documented at the neighboring and contemporaneous EUP site of Dzudzuana Cave, where steppe bison represented 57% of the assemblage and Caucasian tur 40% (Bar-Oz et al. 2004). The abundance of Caucasian tur at Ortvale Klde is remarkable and, as far as can be determined from published data, cannot be matched at other Palaeolithic sites in the Caucasus (see, e.g., Hoffecker, Baryshnikov, and Potapova 1991;Baryshnikov and Hoffecker 1994;Baryshnikov, Hoffecker, and Burgess 1996;Hoffecker 2002). Barakaevskaïa Cave, located roughly 350 km northwest of Ortvale Klde in the northern Caucasus, contains a faunal assemblage with one of the highest frequencies of Caucasian tur (28.2% [Liubin 1998]). ...
... It includes, in addition, remains of several species of small carnivores, mainly fox. In contrast, other Middle and Upper Palaeolithic faunal assemblages from the Caucasus are typically dominated by one or two prey taxa often including the wild goat and/or steppe bison (Bison priscus) or aurochs (Bos primigenius) Bar-Oz et al., 2007;Baryshnikov et al., 1996;Cleghorn, 2006;Hoffecker and Cleghorn, 2000). Studies of Palaeolithic assemblages from the sites of Table 6 Frequencies of fracture angle, fracture outline, fracture edge and shaft circumference of ungulates and cave bear bones from Hovk-1. ...
Information on mammal remains from 665 sites/localities was used to estimate the range of size diversity of European species in the Eem interglacial (MIS 5e) and the GS 2.1 stadial (MIS 2). Review of faunal composition and comparison of the rank distributions of the occurrence of species remains across sites allowed determination of the composition of groups of typical and indicator species for each of the time intervals. Entropy, dominance index and self-organization index were calculated from the rank distributions of the occurrence of large, small and medium-sized herbivorous, carnivorous and representatives of the order Eulipotyphla. The parameters of the non-linear function describing the increase in the average number of species that can be found in 1, 2, 3, etc. randomly selected sites were calculated. The relationship between the parameters of these functions and between them and species richness is shown. The paper discusses hypotheses about the relationship between parameter variations and changes in range size ratios in different mammal groups during the Eem interglacial and at the end of the Last Glacial Maximum.
The genetic and morphological variability of the southern form of the European chromosomal race of the small wood mouse Apodemus (Sylvaemus) uralensis (Pallas, 1811) in the North Caucasus was studied. The results of the sequence analysis of the cytochrome b gene fragment from 9 geographical samples from the western, central and eastern parts of the North Caucasus indicate a low degree of their genetic isolation with a relatively high diversity of haplotypes. Analysis of morphological variability revealed high variety: three morphological groups were identified. Representatives of all three or two groups were found in the same geographical samples, but their distribution did not conform to the hypothesis of random occurrence. Based on indirect evidence, hypotheses about possible causes of the observed features of regional small forest mouse biodiversity are formulated, which require verification in the course of future studies. In particular, we hypothesised that in the Late Pleistocene (130–11.7 thousand cal yr BP) there were several isolated populations in the region that could have been sources of species diversity in the Holocene.
Climatic and environmental fluctuations as well as anthropogenic pressure have led to the extinction of much of Europe’s megafauna. Here we show that the emblematic European bison has experienced several waves of population expansion, contraction and extinction during the last 50,000 years in Europe, culminating in a major reduction of genetic diversity during the Holocene. Fifty-seven complete and partial ancient mitogenomes from throughout Europe, the Caucausus and Siberia reveal that three populations of wisent ( Bison bonasus ) and steppe bison ( B. priscus ) alternated in Western Europe correlating with climate-induced environmental changes. The Late Pleistocene European steppe bison originated from northern Eurasia whereas the modern wisent population emerged from a refuge in the southern Caucasus after the last glacial maximum. A population overlap in a transition period is reflected in ca. 36,000 year-old paintings in the French Chauvet cave. Bayesian analyses of these complete ancient mitogenomes yielded new dates of the various branching events during the evolution of Bison and its radiation with Bos that lead us to propose that the genetic affiliation between the wisent and cattle mitogenomes result from incomplete lineage sorting rather than post-speciation gene flow.
Significance
Climatic fluctuations during the Pleistocene had a major impact on the environment and led to multiple megafaunal extinctions. Through ancient DNA analyses we decipher these processes for one of the largest megafauna of Eurasia, the bison. We show that Western Europe was successively populated during the Late Pleistocene by three different bison clades or species originating from the Caucasus and North-Eastern Europe that can be correlated to major climatic fluctuations and environmental changes. Aurignacian cave artists were witnesses to the first replacement of bison species ~35,000 years ago. All of these populations went extinct except for one that survived into the Holocene where it experienced severe reductions of its genetic diversity due to anthropogenic pressure.
Background: Climatic and environmental fluctuations as well as anthropogenic pressure have led to the
extinction of much of Europe’s megafauna. The European bison or wisent (Bison bonasus), one of the last wild
European large mammals, narrowly escaped extinction at the onset of the 20th century owing to hunting and
habitat fragmentation. Little is known, however, about its origin, evolutionary history and population dynamics
during the Pleistocene.
Results: Through ancient DNA analysis we show that the emblematic European bison has experienced several
waves of population expansion, contraction, and extinction during the last 50,000 years in Europe, culminating
in a major reduction of genetic diversity during the Holocene. Fifty-seven complete and partial ancient mitogenomes
from throughout Europe, the Caucasus, and Siberia reveal that three populations of wisent (Bison bonasus) and
steppe bison (B. priscus) alternately occupied Western Europe, correlating with climate-induced environmental
changes. The Late Pleistocene European steppe bison originated from northern Eurasia, whereas the modern
wisent population emerged from a refuge in the southern Caucasus after the last glacial maximum. A population
overlap during a transition period is reflected in ca. 36,000-year-old paintings in the French Chauvet cave. Bayesian
analyses of these complete ancient mitogenomes yielded new dates of the various branching events during the
evolution of Bison and its radiation with Bos, which lead us to propose that the genetic affiliation between
the wisent and cattle mitogenomes result from incomplete lineage sorting rather than post-speciation gene flow.
Conclusion: The paleogenetic analysis of bison remains from the last 50,000 years reveals the influence of
climate changes on the dynamics of the various bison populations in Europe, only one of which survived into
the Holocene, where it experienced severe reductions in its genetic diversity. The time depth and geographical
scope of this study enables us to propose temperate Western Europe as a suitable biotope for the wisent
compatible with its reintroduction.
Keywords: Ancient DNA, Bison, Population
Background
Climatic and environmental fluctuations as well as anthropogenic pressure have led to the extinction of much of Europe’s megafauna. The European bison or wisent (Bison bonasus), one of the last wild European large mammals, narrowly escaped extinction at the onset of the 20th century owing to hunting and habitat fragmentation. Little is known, however, about its origin, evolutionary history and population dynamics during the Pleistocene.
Results
Through ancient DNA analysis we show that the emblematic European bison has experienced several waves of population expansion, contraction, and extinction during the last 50,000 years in Europe, culminating in a major reduction of genetic diversity during the Holocene. Fifty-seven complete and partial ancient mitogenomes from throughout Europe, the Caucasus, and Siberia reveal that three populations of wisent (Bison bonasus) and steppe bison (B. priscus) alternately occupied Western Europe, correlating with climate-induced environmental changes. The Late Pleistocene European steppe bison originated from northern Eurasia, whereas the modern wisent population emerged from a refuge in the southern Caucasus after the last glacial maximum. A population overlap during a transition period is reflected in ca. 36,000-year-old paintings in the French Chauvet cave. Bayesian analyses of these complete ancient mitogenomes yielded new dates of the various branching events during the evolution of Bison and its radiation with Bos, which lead us to propose that the genetic affiliation between the wisent and cattle mitogenomes result from incomplete lineage sorting rather than post-speciation gene flow.
Conclusion
The paleogenetic analysis of bison remains from the last 50,000 years reveals the influence of climate changes on the dynamics of the various bison populations in Europe, only one of which survived into the Holocene, where it experienced severe reductions in its genetic diversity. The time depth and geographical scope of this study enables us to propose temperate Western Europe as a suitable biotope for the wisent compatible with its reintroduction.
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Between 2002 and 2009, renewed investigations of Units II, III, IV and V at Azokh 1 cave were undertaken following rigorous systematic methods of excavation and recording. New dates suggest an age of 184–100 ka for Unit II and ~300 ka for Unit Vm. The excavations produced a range of fossil faunas dominated by cave bears, and 387 lithic artifacts: 68 from Unit Vm, 4 from Unit III, and 315 from Unit II. Although a range of rock types was exploited for tool production, most artifacts are on siliceous rocks, with a few made from non-local obsidian. There is little evidence for the early stages of production on-site. It is possible that initial working may have occurred elsewhere, and cores, blanks and tools transported to the cave. However, given the restricted area of excavations at the rear of the cave, we cannot discount the possibility of knapping activities having occurred in other areas of the cave, the sediments of which were removed in earlier excavations. The stone artifacts from Unit II, with their Levallois component, are clearly Middle Paleolithic, and may be among the earliest evidence for Middle Paleolithic presence in the Southern Caucasus. The material from Unit Vm could be late Acheulean on the basis of dating, lack of Levallois technology, the general larger sizes of the pieces (although no bifaces have been found), and its stratigraphic position below Units II, III, and IV.
The craniometric differences between two closely related species, Sorex minutus and S. volnuchini ("minitus" species group) and fossil shrews of this group from the Northern Caucasus were studied by the discriminant analysis. The significant morphological differences between the current forms were revealed. The data obtained confirm the species rank of S. volnuchini. The Pleistocene Caucasian shrews of the "minutus" group belong to the species S. volnuchini. Sorex minutus and S. volnuchini existed as individual species from the early Pleistocene period. Problems of subspecies taxonomy of S. volnuchini are discussed.
The late Pleistocene remains of insectivorous mammals belonging to the order Insectivora from four localities of the Northern Caucasus (the caves of Matuzka, Mezmaiskaya, Monasheskaya, and Myshtulagty) were studied. Most of the samples is referred to the Chasoval'skii faunistic complex. Generally, the Insectivora fauna dealt with is somehow intermediate between the Middle Pleisocene and recent ones. For instance, Sorex minutissimus, S. cf. runtonensis, and Neomys hintoni, which inhabited the Caucasus in the Middle Pleistocene, and recent S. cf. satunini, S. raddei, Neomys teres, and Erinaceus concolor were discovered within the same deposits together with Talpa sp. However, S. doronochevi, Drepanosorex rupestris, and N. newtoni, which are khown for the Middle Pleistocene, occurred to be completely eliminated. The distinguishing feature of this period is the predominance of the Caucasian shrew S. satinini. Presently, it is the dominant species of subalpine meadows in the Caucasus. The presence of S. raddei and Erinaceus concolor among the findings indicates the existence of forest habitats over those times.
The culmination of more than a decade of fieldwork and related study, this unique book uses analyses of perimortem taphonomy in Ice Age Siberia to propose a new hypothesis for the peopling of the New World. The authors present evidence based on examinations of more than 9000 pieces of human and carnivore bone from 30 late Pleistocene archaeological and palaeontological sites, including cave and open locations, which span more than 2000 miles from the Ob River in the West to the Sea of Japan in the East. The observed bone damage signatures suggest that the conventional prehistory of Siberia needs revision and, in particular, that cave hyenas had a significant influence on the lives of Ice Age Siberians. The findings are supported by more than 250 photographs, which illustrate the bone damage described and provide a valuable insight into the context and landscape of the fieldwork for those unfamiliar with Siberia. © Christy G. Turner II, Nicolai D. Ovodov and Olga V. Pavlova 2013.
A revision of the Caprini from the Caucasus provides new information on the dispersal of the species and on the biochronology of the Mousterian sites of Kudaro I and III. In both Capra caucasica is recognized. Remains from Kudaro I show an archaiq morphology in comparison with Kudaro III. The discovery of Hemitragus bonali and Ovis ammon antiqua is in accordance with a Middle Pleistocene age of the base of Kudaro I site. In Kudaro III, a well preserved male ibex skull is closer to the recent sub-species living in the western margins of Caucasus Mountains than the Elbrouz ibex which is distributed in its Center.
This is the first book to provide a systematic overview of social zooarchaeology, which takes a holistic view of human–animal relations in the past. Until recently, archaeological analysis of faunal evidence has primarily focused on the role of animals in the human diet and subsistence economy. This book, however, argues that animals have always played many more roles in human societies: as wealth, companions, spirit helpers, sacrificial victims, totems, centerpieces of feasts, objects of taboos, and more. These social factors are as significant as taphonomic processes in shaping animal bone assemblages. Nerissa Russell uses evidence derived from not only zooarchaeology, but also ethnography, history, and classical studies to suggest the range of human–animal relationships and to examine their importance in human society. Through exploring the significance of animals to ancient humans, this book provides a richer picture of past societies.
For us, the experience of reading Steve Churchill's book Thin on the Ground: Neandertal Biology, Archeology, and Ecology was like that of reading The Origin of Species for the first time. In both Churchill's and Darwin's books, the reader is led carefully and meticulously through a beautifully organized presentation of all the evidence bearing on a vexed and long-standing problem, arriving at a novel answer that resolves many issues all at once. Like Darwin, Churchill makes his case with such a wide-ranging, comprehensive, and judicious presentation that when the overall conclusion is fully laid out in the last chapter, its force is inescapable.
This paper presents a short review of the data and new approaches to Middle Paleolithic chronology and variability in the Caucasus. A climatostratigraphic scheme of the Caucasian Middle Paleolithic is proposed on the basis of the oxygen isotope stages. The authors attempt to identify tool types characteristic of the Caucasian Middle Paleolithic industries, as well as to reveal its development during more than 100,000 years. A new approach to treating Middle Paleolithic variability in the Caucasus recognizes three general cultural areas: (1) North Caucasian Micoquian in the Northwestern Caucasus; (2) Khostinian and several cultural entities possibly rooted in Levantine D (Djruchulian), Levantine B (Tskhinvali group), and Karain (?) Mousterian (Tskhaltsitela–Tsutskhvati group) in the South-central Great Caucasus; (3) Zagros Mousterian in the southernmost part of the Caucasus.
Intraspecific variability of the Pleistocene speciesSorex runtonensis Hinton, 1911 from different Polish and Russian (Caucasus Mts.) localities and its relationships to Recent red-toothed shrews
of similar body size and mandibular morphology,S. caecutiens Laxmann, 1788 andS. tundrensis Merriam, 1990, are explained on the grounds of multivariate statistics. The remains ofS. runtonensis from different localities form a single group and differ fromS. caecutiens. They resembleS. tundrensis by the first canonical root related to a mandibular size and proportions. This may indicate thatS. runtonensis andS. tundrensis are closely related but separated components of the species complextundrensis.
Pleistocene remains from Chionomys from Palaeolithic cave sites of Northern Caucasus are described. Ch. gud and Ch. roberti are present in the region at least from the beginning of Middle Pleistocene while Ch. nivalis seems not to appear before Early/Middle Weichselian boundary. -Authors
Some ungulate species at Upper Pleistocene and Holocene archeological sites in South Africa exhibit catastrophic mortality profiles, while others exhibit attritional ones. The awareness by Stone Age people that some species are especially amenable to driving or snaring probably accounts for the catastrophic profiles. Natural catastrophic death immediately followed by human scavenging is a much less likely explanation because the species samples comprise material lumped from deposits that accumulated more or less continuously over hundreds or even thousands of years during which period there is no reason to suppose the repeated occurrence of natural catastrophes nearby.
The inability of Stone Age people to obtain prime-age adults in species that are not particularly amenable to driving or snaring presumably accounts for the attritional mortality profiles. Although the species that display attritional profiles conceivably were scavenged, the high proportion of very young individuals in the profiles suggests active hunting. Very young individuals are much less abundant in attritional profiles from local non-archeological sites, probably because their carcasses were removed from the record before burial, primarily by carnivore or scavenger feeding. Scavenging would account for the abundance of very young individuals in the archeological sites only in the unlikely event that people could regularly locate carcasses before other predators did.
In general, geomorphic/sedimentologic context is probably the best criterion for determining whether a species characterized by a catastrophic profile in an archeological site was hunted or scavenged. At the majority of known sites, active hunting is suggested. In the case of a species characterized by an attritional profile in an archeological site, the proportion of very young individuals in the sample probably provides the best criterion for distinguishing hunting from scavenging. A relatively high proportion of very young individuals suggests active hunting.
In growing numbers, archeologists are specializing in the analysis of excavated animal bones as clues to the environment and behavior of ancient peoples. This pathbreaking work provides a detailed discussion of the outstanding issues and methods of bone studies that will interest zooarcheologists as well as paleontologists who focus on reconstructing ecologies from bones. Because large samples of bones from archeological sites require tedious and time-consuming analysis, the authors also offer a set of computer programs that will greatly simplify the bone specialist's job. After setting forth the interpretive framework that governs their use of numbers in faunal analysis, Richard G. Klein and Kathryn Cruz-Uribe survey various measures of taxonomic abundance, review methods for estimating the sex and age composition of a fossil species sample, and then give examples to show how these measures and sex/age profiles can provide useful information about the past. In the second part of their book, the authors present the computer programs used to calculate and analyze each numerical measure or count discussed in the earlier chapters. These elegant and original programs, written in BASIC, can easily be used by anyone with a microcomputer or with access to large mainframe computers.
Primate fossils from the Acheulian handaxe site at Olorgesailie, Kenya, are analyzed in an attempt to evaluate Isaac's suggestion that Locality DE/89 B preserves the remains of the hunting and butchering of giant gelada baboons. The age and sex structure of the 90 individuals suggests that attritional mortality occurred. This evidence is consistent with predation on small numbers of individuals at a time by either hominids or carnivores. A comparative method of analyzing breakage patterns is used. The type and frequency of breaks on each skeletal element are compared statistically with those on the same skeletal elements of primates from broadly contemporaneous sites where hunting and butchering are not suggested to have occured but where carnivore activity and other sources of damage have occurred. Some aspects of the breakage pattern on the Olorgesailie baboon material are significantly different from those at the nonhominid sites. The breaks in question could have occurred during disarticulation of the primates. The most probable interpretation of the evidence is that the hominids at Olorgesailie systematically and repeatedly hunted and butchered giant geladas. This is the first evidence of such a behavior in the fossil record.
Recent investigations have produced a group of important new Mousterian sites in the NW Caucasus (Russia). These sites, which range in elevation from 100 to 1350 m above sea level, include Il'skaya II (100 m)) Matuzka Cave (750 m), Barakaevskaya Cave (900 m), and Mezmaiskaya Cave (1350 m). Each contains a large faunal assemblage that, in conjunction with faunal remains excavated earlier from Il'skaya I (100 m) and Dakhovskaya Cave (500 m), constitute a rich source of information on Mousterian paleoecology in this part of Europe. Sites at lower elevations yield remains of mammoths and giant deer; at higher elevations, these taxa are absent but goats and sheep are common. Steppe bison is the most common species in a majority of assemblages, and bone modification and demographic data indicate that it was probably hunted (as opposed to scavenged) by Mousterian groups. Cave bear remains exhibit an inverse relationship to artifacts, and most probably reflect natural mortality. The economic role of mammoths is problematic; many bones and tusks may have been scavenged or collected.
A new analysis of vertebrate remains from the Mousterian open-air site of Il'skaya I was undertaken. The site is located near Krasnodar in the northern Caucasus. The faunal samples available to us were excavated in 1926, 1928, and 1936–1937, and comprise c. 1500 identifiable bones and teeth; mixture of remains from different stratigraphic levels significantly affects the results of the analysis. Steppe bison (Bison priscus) dominates the mammalian assemblage (92%), which also includes giant deer, horse, red deer, wolf, hyaena, mammoth, wild ass, corsac, bear, saiga, and dhole. Bird remains (rare for Mousterian open-air sites) include black vulture, magpie, and mallard. Bison is represented by virtually all skeletal parts, although isolated teeth are especially abundant; variations in skeletal-part frequencies primarily reflect weathering effects. Age profiels for bison, obtained from molar crown-height measurements, indicate the predominance of prime-age adults; both males and females appear to be represented, but estimation of sex ratio is problematic. The site was probably occupied during several seasons. The results are discussed within the context of general issues concerning Mousterian subsistence strategy.
Mortality patterns in archaeofaunas can be informative of prehistoric human foraging habits, land use, and, ultimately, evolutionary changes in hominid sociality and ecological niche. The analytical value of mortality patterns is only as great, however, as archaeologists' understanding of the full range of possible causes for patterns in these data. Here, the relationships between mortality patterns in death assemblages and their documented causes are examined. Interspecific comparisons reveal that, while mortality patterns alone cannot diagnose cause, these data are potentially powerful tools for studies of hominid subsistence if supported by taphonomic analyses of bone assemblage formation. Mortality analyses are particularly effective if age frequency data are divided according to life history characteristics of prey species. Comparisons of known modern cases to ungulate assemblages created by Holocene and Pleistocene hominids of westcentral Italy present new information on humans as predators and evolutionary changes therein. These data indicate significantly greater strategic variation in the Middle Paleolithic cases than for all subsequent cultural periods combined. The variation certainly corresponds to two or more distinct foraging/land use strategies, scavenging and ambush hunting—the latter of which became more specialized with time.
Bones of recent mammals in the Amboseli Basin, southern Kenya, exhibit distinctive weathering characteristics that can be related to the time since death and to the local conditions of temperature, humidity and soil chemistry. A categorization of weathering characteristics into six stages, recognizable on descriptive criteria, provides a basis for investigation of weathering rates and processes. The time necessary to achieve each successive weathering stage has been calibrated using known-age carcasses. Most bones decompose beyond recognition in 10 to 15 yr. Bones of animals under 100 kg and juveniles appear to weather more rapidly than bones of large animals or adults. Small-scale rather than widespread environmental factors seem to have greatest influence on weathering characteristics and rates. Bone weathering is potentially valuable as evidence for the period of time represented in recent or fossil bone assemblages, including those on archeological sites, and may also be an important tool in censusing populations of animals in modern ecosystems.
Criteria for distinguishing between bone assemblages accumulated by hominids versus those accumulated by hyenas are discussed. These criteria are based on observations from southern African sites, particularly the Upper Pleistocene brown hyena accumulations from Swartklip and Equus Cave. When contextual evidence is lacking or ambiguous, these criteria can be used to determine if a bone accumulation is primarily the result of hyena or hominid activities. The criteria are as follows: 1) the tendency for carnivore remains to be relatively common in carnivore accumulations, 2) the presence of distinctive hyena damage on bone surfaces (depending on bone surface preservation, such damage may not always be common in hyena assemblages); 3) the tendency for bones from hyena accumulations to have relatively complete shafts but lack epiphyses (i.e., being bone “cylinders” while those from hominid accumulations have broken shafts and intact epiphyses; 4) the tendency for the cranial-postcranial ratio to decrease with ungulate size in hyena accumulations; 5) the tendency for small, hard, bones to be uncommon in hyena accumulations, regardless of state of preservation; and 6) the tendency for age profiles to be attritional in hyena accumulations.
Recent excavation and subsequent analysis have been undertaken for the express purpose of deriving quantitative techniques for the analysis of faunal remains from archaeological sites. Experiments were conducted which indicate that significant amounts of bones are lost through the various meshes of screen used by archaeologists. Factors were then calculated to alleviate this problem and to yield results amenable to rigorous quantitative treatment. Results obtained in this manner can add accuracy to most salvage operations and credibility to the more ideal excavation conditions. The proposed method was applied to data from Smoky Creek Cave, a Medithermal site approximately 40 mi. north of Gerlach, Washoe County, Nevada; results indicate a pattern of cottontail rabbit and mountain sheep hunting similar to the model which Flannery has proposed for the Tehuacan Valley, Mexico.
Mary Stiner uses ecological niche theory to analyze and interpret several Middle Paleolithic archaeological and paleontological sites in southern Europe. Her concern is with how the hunting, scavenging, and foraging behavior of Neandertals compared and contrasted with the subsistence behavior of other large predators living in the region at the time - lions, hyenas, and wolves, for example - and with how Neandertal subsistence behavior related to the behavior of the anatomically modern humans who subsequently came to dominate the area in the Upper Paleolithic. Her conclusion, very broadly stated, is that Neandertals entered the Middle Paleolithic in direct and successful competition with lions, hyenas, and wolves, but ended the period in direct and ultimately unsuccessful competition for the ecological niche that we came to occupy with our slightly more advanced technology and slightly more sophisticated ambush hunting strategies and techniques
Morphological distinctions between the mandibular teeth of young sheep (Ovis) and goats (Capra) are described, and their value in the study of bone samples from archaeological sites is briefly discussed.
"Amongst scientists involved [in taphonomy], C. K. Brain stands out as the pioneer; this impressive book is a statement of his investigations. . . . The Hunters or the Hunted? is a very important book for paleoanthropology. It presents the first thorough analysis of the Sterkfontein Valley assemblages, contributes significantly to the resolution of lingering controversies and, by placing the old information in a fresh perspective, enables new and more sophisticated questions to be asked not only of the South African material but of similar assemblages elsewhere. Another contribution is that it reinforces the recent change in feelings as to what constitutes data, for the value of looking at fossil and contemporary bones as closely as this is clear. Brain urges the necessity of recovering fossils with a high regard for subtle detail. I hope excavators of any vertebrate fossil site will be persuaded to follow his advice and pay more attention to these features of bone accumulations that have been previously neglected; for taphonomy can be a powerful tool in elucidating the problems of fossil assemblages, especially when handled with the care and caution that Brain brings to the subject."âAndrew Hill, Nature
In growing numbers, archeologists are specializing in the analysis of excavated animal bones as clues to the environment and behavior of ancient peoples. This pathbreaking work provides a detailed discussion of the outstanding issues and methods of bone studies that will interest zooarcheologists as well as paleontologists who focus on reconstructing ecologies from bones. Because large samples of bones from archeological sites require tedious and time-consuming analysis, the authors also offer a set of computer programs that will greatly simplify the bone specialist's job. After setting forth the interpretive framework that governs their use of numbers in faunal analysis, Richard G. Klein and Kathryn Cruz-Uribe survey various measures of taxonomic abundance, review methods for estimating the sex and age composition of a fossil species sample, and then give examples to show how these measures and sex/age profiles can provide useful information about the past. In the second part of their book, the authors present the computer programs used to calculate and analyze each numerical measure or count discussed in the earlier chapters. These elegant and original programs, written in BASIC, can easily be used by anyone with a microcomputer or with access to large mainframe computers.
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