Abstract
The Crimean Peninsula has a rich archaeological record characterised by numerous multi-layered sites from the Middle Palaeolithic and by a number of important sites representing the transition from the Middle to the Upper Palaeolithic. Absolute dates place the region as one of the last Neanderthal refuges in Europe. Combining dental meso- and microwear analyses of worn molars found at faunal assemblages from different sites across Crimea, we can characterise the feeding habits of ungulates at different scales in their lifetime. The proxy data resulting from these analyses provide evidence not only on diet but also on (1) the habitat where fauna were hunted and (2) the duration of the events of human occupation in each archaeological level. For this palaeoecological reconstruction, we analysed three sites: Kabazi II and Chokurcha I from the Middle Palaeolithic and Siuren I from the Early Upper Palaeolithic. We focused particularly on the repeated Neanderthal subsistence pattern observed at Kabazi II throughout the sequence (around 80 to 48 ky). Analyses revealed that Equus hydruntinus (wild ass) was feeding on abrasive grasses, suggesting it was occupying open habitats, while Saiga tatarica (saiga antelope) was mixed feeding and probably consuming significant amounts of browse and lichens in relatively open habitats. Furthermore, variability in microwear signalled short seasonal occupations for all assemblages. During the Middle Palaeolithic and the early Upper Palaeolithic, these sites were occupied during the warm season.
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References
Ackermans NL, Martin LF, Codron D, Hummel J, Kircher PR, Richter H, Kaiser TM, Clauss M, Hatt JM (2020) Mesowear represents a lifetime signal in sheep (Ovis aries) within a long- term feeding experiment. Palaeogeogr Palaeoclimatol Palaeoecol 553:109793. https://doi.org/10.1016/j.palaeo.2020.109793
Allsworth-Jones P (1990) The Szeletian and the stratigraphic succession in central Europe and adjacent areas: main trends, recent results, and problems for resolution. In: Mellars P (ed) The emergence of modern humans: an archaeoogical perspective. Cornell University Press, New York, pp 160–242
Bahloul K, Pereladova OB, Soldatova N, Fisenko G, Sidorenko E, Sempéré AJ (2001) Social organization and dispersion of introduced kulans (Equus hemionus kulan) and Przewalski horses (Equus przewalski) in the Bukhara reserve, Uzbekistan. J Arid Environ 47:309–323. https://doi.org/10.1006/jare.2000.0714
Banks WE (2020) Puzzling out the Middle-to-Upper Palaeolithic transition. Nat Ecol Evol 4:775–776. https://doi.org/10.1038/s41559-020-1162-1
Barron E, van Andel TH, Pollard D (2003) Glacial environment II: reconstructing the climate of Europe in the last glaciation. In: van Andel TH, Davis W (eds) Neanderthals and modern humans in the European landscape during the last glaciation: archaeological results of the stage 3 project. McDonald Institute for Archaeological Research, Cambridge, pp 57–78
Baryshnikov G (2003) Mammuthus primigenius from the Crimea and the Caucasus. Deinsea 9:41–56
Baskin L, Danell K (2003) Ecology of ungulates: a handbook of species in Eastern Europe and Northern and Central Asia. Springer-Verlag, Berlin Heidelberg
Bataille G (2010) Recurrent occupations of the Late Middle Palaeolithic Station Kabazi II, Unit II, Level 8 (Crimea, Ukraine)–seasonal adaptation, procurement and processing of resources. Quartär 57:43–77
Bataille, G (2012) Stones and bones. The reconstruction of occupational palimpsests in the Late Middle Palaeolithic of Crimea (Ukraine). In Cascalheira J, Gonçalves C (eds) Actas das IV Jornadas de Jovens em Investigação Arqueológica–JIA 2011(2) Promontoria Monográfica 16, Universidade do Algarve, Faro, pp. 201–209
Bataille G (2016) Neanderthals of Crimea–creative generalists of the late Middle Paleolithic. Contextualizing the leaf point industry Buran-Kaya III, Level C. Quat Int 435:211–236. https://doi.org/10.1016/j.quaint.2015.12.071
Bekenov AB, Grachevand IA, Milner-Gulland EJ (1998) The ecology and management of the Saiga antelope in Kazakhstan. Mamm Rev 28:1–52. https://doi.org/10.1046/j.1365-2907.1998.281024.x
Bennett EA, Champlot S, Peters J, Arbuckle BS, Guimaraes S, Pruvost M, Bar-David S, Davis SJM, Gautier M, Kaczensky P, Kuehn R, Mashkour M, Morales-Muñiz A, Pucher E, Tournepiche JF, Uerpmann HP, Bălăşescu A, Germonpré M, Gündem CY, Hemami MR, Moullé PE, Ötzan A, Uerpmann M, Walzer C, Grange T, Geigl EM (2017) Taming the late Quaternary phylogeography of the Eurasiatic wild ass through ancient and modern DNA. PLoS One 12:1–28. https://doi.org/10.1371/journal.pone.0174216
Boulbes N, van Asperen EN (2019) Biostratigraphy and Palaeoecology of European Equus. Front Ecol Evol 7. https://doi.org/10.3389/fevo.2019.00301
Burke A (2000) The view from Starosele: faunal exploitation at a Middle Palaeolithic site in Western Crimea. Int J Osteoarchaeol 10:325–335. https://doi.org/10.1002/1099-1212(200009/10)10:5<325: AID-OA556>3.0.CO;2-T
Burke A (2006) Neanderthal settlement patterns in Crimea: a landscape approach. J Anthropol Archaeol 25:510–523. https://doi.org/10.1016/j.jaa.2006.03.003
Burke A, Ebert D, Cardille J, Dauth D (2008) Paleoethology as a tool for the development of archaeological models of land-use: the Crimean Middle Palaeolithic. J Archaeol Sci 35:894–904. https://doi.org/10.1016/j.jas.2007.06.014
Burke A, Eisenmann V, Ambler GK (2003) The systematic position of Equus hydruntinus, an extinct species of Pleistocene equid. Quat Res 59:459–469. https://doi.org/10.1016/S0033-5894(03)00059-0
Burke A, Markova AK, Mikhailesku C, Patou-Mathis M (1999) The animal environment of Western Crimea. In: Chabai VP, Monigal K (eds) The Middle Paleolithic of Western Crimea. Vol. 2. ERAUL 87, Liège, pp 143–151
Carrión JS, Ochando J, Fernández S, Blasco R, Rosell J, Munuera M, Amorós G, Martín-Lerma I, Finlayson S, Giles F, Jennings R, Finlayson G, Giles-Pacheco F, Rodríguez-Vidal J, Finlayson C (2018) Last Neanderthals in the warmest refugium of Europe: Palynological data from vanguard cave. Rev Palaeobot Palynol 259:63–80. https://doi.org/10.1016/j.revpalbo.2018.09.007
Catalano G, Modi A, Mangano G, Sineo L, Lari M, Bonfiglio L (2020) A mitogenome sequence of an Equus hydruntinus specimen from Late Quaternary site of San Teodoro Cave (Sicily, Italy). Quat Sci Rev 236:106280. https://doi.org/10.1016/j.quascirev.2020.106280
Chabai VP (1998a) Kabazi II: introduction. In: Marks AE, Chabai VP. (eds) The Middle Paleolithic of Western Crimea.Vol.1. ERAUL 84, Liège, pp 167–200
Chabai VP (1998b) The history of Crimean middle Paleolithic studies. In: Marks A, Chabai VP (eds) The Middle Paleolithic of Western Crimea. Vol.1. ERAUL 84, Liège, pp 1–16
Chabai VP (2004a) Chokurcha I: introduction. In: Chabai VP, Monigal K, Marks AE (eds) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea. Vol. 3. ERAUL 104, Liège, pp 343–353
Chabai VP (2004b) The Middle Paleolithic of Crimea. Simferopol: Shlyakh
Chabai VP (2005) Kabazi II: stratigraphy and archaeological sequence. In: Chabai VP, Richter J, Uthmeier Th (eds) Kabazi II: last interglacial occupation, Environment & Subsistence. Simferopol - Cologne, pp. 1–24
Chabai VP (2007) Kabazi V: the lithological and archaeological sequences. In: Chabai VP, Richter J, Uthmeier, T (eds), Kabazi V: interstratification of Micoquian and Levallois-Mousterian Camp Sites. Paleolithic sites of Crimea 3, part 1. Simferopol – Cologne: Shlyakh, pp. 1–26
Chabai VP, Marks AE, Monigal K (2004) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea. ERAUL 104, Liège
Chabai VP, Monigal K (eds) (1999) The Middle Paleolithic of Western Crimea, volume 2. ERAUL 87, Liège
Chabai VP, Patou-Mathis M (2009) Kabazi V, late Middle Palaeolithic camps: raw material and fauna exploitation. Archaeol Alm 20:59–76
Chabai VP, Richter J, Uthmeier Th (eds) (2005) Kabazi II: last interglacial occupation, environment and subsistence. Palaeolithic Sites of Crimea. Vol.1. Simferopol – Cologne: Shlyakh
Chabai VP, Richter J, Uthmeier Th (eds) (2006) Kabazi II: the 70 000 years since the last interglacial. Palaeolithic Sites of Crimea. Vol.2. Simferopol – Cologne: Shlyakh
Chabai VP, Richter J, Uthmeier Th (eds) (2007) Kabazi V: interstratification of Micoquian & Levallois-Mousterian Camp Sites. Part 1. Palaeolithic Sites of Crimea. Vol. 3. Simferopol – Cologne: Shlyakh
Chabai VP, Richter J, Uthmeier Th (eds) (2008) Kabazi V: interstratification of Micoquian & Levallois-Mousterian Camp Sites. Part 2. Palaeolithic Sites of Crimea. Vol.3. Simferopol – Cologne: Shlyakh
Chabai VP, Uthmeier Th (2006) Settlement Systems in the Crimean Middle Palaeolithic. In: Chabai VP, Richter J, Uthmeier Th (eds) Kabazi II: 70.000 years since the last interglacial. Palaeolithic Sites of Crimea. Vol.2. Simferopol – Cologne: Shlyakh, pp 297–359
Chabai VP, Uthmeier Th (2017) New excavations at the Middle Paleolithic site Zaskalnaya V, Crimea: the 2012-2013 field seasons. A preliminary Report. Quartär 64: 27–61
Chen L, Wolf AB, Fu W et al (2020) Identifying and interpreting apparent Neanderthal ancestry in African individuals. Cell 180:677–687.e16. https://doi.org/10.1016/j.cell.2020.01.012
Cordova CE, Gerasimenko NP, Lehman PH, Kliukin AA (2011) Late Pleistocene and Holocene paleoenvironments of Crimea: pollen, soils, geomorphology, and geoarchaeology. Geol Soc Am 473:133–164. https://doi.org/10.1130/2011.2473(09)
Demidenko YE (2004) Buran-Kaya III Layer B: the lithic assemblage. In Chabai VP, Monigal K, Marks AE (ed) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea, Vol. 3. ERAUL 104, Liège, pp 113-149
Demidenko YE (2008) The Early and Mid-Upper Palaeolithic of the North Black Sea region: an overview. Quartär 55:99–114
Demidenko YE (2012) Interpretation of the Middle Paleolithic component in the Early Aurignacian Units H and G and the 1 920s Lower Layer. In Demidenko YE, Otte M, Noiret P (eds) Siuren I Rock-shelter. From Late Middle Paleolithic and Early Upper Paleolithic to Epi-Paleolithic in Crimea. ERAUL 129, Liège, pp 321-332
Demidenko YE (2014) Siuren I Rockshelter: From the Late Middle Paleolithic and Early Upper Paleolithic to the Epipaleolithic in Crimea. Encycl Glob Archaeol 3:6711–6721
Demidenko YE, Otte M (2001) Siuren-I (Crimea) in the context of a European Aurignacian. Préhistoire Eur 16–17:133–146
Demidenko YE, Otte M, Noiret P (eds) (2012) Siuren I Rock-shelter. From Late Middle Paleolithic and Early Upper Paleolithic to Epi-Paleolithic in Crimea. ERAUL 129, Liège
Demidenko YE, Noiret P (2012) Radiocarbon dates for the Siuren I Sequence. In: Demidenko YE, Otte M, Noiret P (eds) Siuren I rock-shelter. From Late Middle Paleolithic and Early Upper Paleolithic to Epi-Paleolithic in Crimea. ERAUL 129, Liège, pp 49–53
Demidenko YE, Uthmeier Th (2013) Kiik-Koba Grotto, Crimea (Ukraine). Re-analysis of a key site of the Crimean Micoquian. Kölner Studien zur Prähistorischen Archäologie. Vol. 3. Maria Leidorf Verlag, Rahden/Westf
Feh C, Munkhtuya B, Enkhbold S, Sukhbaatar T (2001) Ecology and social structure of the Gobi khulan Equus hemionus subsp. in the Gobi B National Park, Mongolia. Biol Conserv 101:51–61. https://doi.org/10.1016/S0006-3207(01)00051-9
Fewlass H, Talamo S, Wacker L, Kromer B, Tuna T, Fagault Y, Bard E, McPherron SP, Aldeias V, Maria R, Martisius NL, Paskulin L, Rezek Z, Sinet-Mathiot V, Sirakova S, Smith GM, Spasov R, Welker F, Sirakov N, Tsanova T, Hublin JJ (2020) A 14 C chronology for the Middle to Upper Palaeolithic transition at Bacho Kiro Cave, Bulgaria. Nat Ecol Evol 62:794–801. https://doi.org/10.1038/s41559-020-1136-3
Fortelius M, Solounias N (2000) Functional characterization of ungulate molars using the abrasion-attrition wear gradient: a new method for reconstructing paleodiets. Am Museum Novit 3301:1–36. https://doi.org/10.1206/0003-0082(2000)301<0001: FCOUMU>2.0.CO;2
Gerasimenko N (1999) Late Pleistocene vegetational history of Kabazi II. In: Chabai VP, Monigal K (eds) The Middle Palaeolithic of Western Crimea. Vol.2. ERAUL 87, Liège, pp 115–142
Gerasimenko N (2004) Vegetational history of Buran-Kaya III. In: Chabai VP, Monigal K, Marks AE (eds) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea.Vol.3. ERAUL 104, Liège, pp 19–34
Gerasimenko N (2005) Vegetation evolution of the Kabazi II Site. In: Chabai VP, Richter J, Uthmeier Th (eds) Kabazi II: last interglacial occupation, environment & subsistence. Palaeolithic Sites of Crimea.Vol.1. Simferopol – Cologne: Shlyakh, pp 25–50
Gerasimenko N (2007) Environmental changes in the Crimean mountains during the last interglacial-middle pleniglacial as recorded by pollen and lithopedology. Quat Int 164–165:207–220. https://doi.org/10.1016/j.quaint.2006.12.018
Green RE, Krause J, Briggs AW, et al (2010) A draft sequence of the Neandertal genome. Science 328:710–722. https://doi.org/10.1126/science.1188021
Grine FE (1986) Dental evidence for dietary differences in Australopithecus and Paranthropus: a quantitative analysis of permanent molar microwear. J Hum Evol 15:783–822. https://doi.org/10.1016/S0047-2484(86)80010-0
Gubonina ZP (1985) Environmental conditions of the Paleolithic in the Crimean foothills. In: Palynology of the Quaternary (in Russian). Nauka, Moscow, pp 95–103
Higham T, Douka K, Wood R, Ramsey CB, Brock F, Basell L, Camps M, Arrizabalaga A, Baena J, Barroso-Ruíz C, Bergman C, Boitard C, Boscato P, Caparrós M, Conard NJ, Draily C, Froment A, Galván B, Gambassini P, Garcia-Moreno A, Grimaldi S, Haesaerts P, Holt B, Iriarte-Chiapusso MJ, Jelinek A, Jordá Pardo JF, Maíllo-Fernández JM, Marom A, Maroto J, Menéndez M, Metz L, Morin E, Moroni A, Negrino F, Panagopoulou E, Peresani M, Pirson S, de la Rasilla M, Riel-Salvatore J, Ronchitelli A, Santamaria D, Semal P, Slimak L, Soler J, Soler N, Villaluenga A, Pinhasi R, Jacobi R (2014) The timing and spatiotemporal patterning of Neanderthal disappearance. Nature 512:306–309. https://doi.org/10.1038/nature13621
Hublin J-J, Sirakov N, Aldeias V, Bailey S, Bard E, Delvigne V, Endarova E, Fagault Y, Fewlass H, Hajdinjak M, Kromer B, Krumov I, Marreiros J, Martisius NL, Paskulin L, Sinet-Mathiot V, Meyer M, Pääbo S, Popov V, Rezek Z, Sirakova S, Skinner MM, Smith GM, Spasov R, Talamo S, Tuna T, Wacker L, Welker F, Wilcke A, Zahariev N, McPherron SP, Tsanova T (2020) Initial Upper Palaeolithic Homo sapiens from Bacho Kiro Cave, Bulgaria. Nature 581:299–302. https://doi.org/10.1038/s41586-020-2259-z
Jürgensen J, Drucker DG, Stuart AJ, Schneider M, Buuveibaatar B, Bocherens H (2017) Diet and habitat of the saiga antelope during the late Quaternary using stable carbon and nitrogen isotope ratios. Quat Sci Rev 160:150–161. https://doi.org/10.1016/j.quascirev.2017.01.022
Kaiser TM, Solounias N (2003) Extending the tooth mesowear method to extinct and extant equids. Geodiversitas 25:321–345
King T, Andrews P, Basak Boz (1999) Effect of taphonomic processes on dental microwear. Am J Phys Anthropol 108:359–373. https://doi.org/10.1002/(SICI)1096-8644(199903)108:3<359: AID-AJPA10>3.0.CO;2–9
Kuhlwilm M, Gronau I, Hubisz MJ, de Filippo C, Prado-Martinez J, Kircher M, Fu Q, Burbano HA, Lalueza-Fox C, de la Rasilla M, Rosas A, Rudan P, Brajkovic D, Kucan Ž, Gušic I, Marques-Bonet T, Andrés AM, Viola B, Pääbo S, Meyer M, Siepel A, Castellano S (2016) Ancient gene flow from early modern humans into Eastern Neanderthals. Nature 530:429–433. https://doi.org/10.1038/nature16544
Majkic A, Evans S, Stepanchuk V et al (2017) A decorated raven bone from the Zaskalnaya VI (Kolosovskaya) Neanderthal site, Crimea. PLoS One 12:1–33. https://doi.org/10.1371/journal.pone.0173435
Marín J, Saladié P, Rodríguez-Hidalgo A, Carbonell E (2017) Neanderthal hunting strategies inferred from mortality profiles within the Abric Romaní sequence. PLoS One 12:1–42. https://doi.org/10.1371/journal.pone.0186970
Markova AK (1999) Small mammal fauna from Kabazi II, Kabazi V, and Starosele: paleoenvironments and evolution. In: Chabai VP, Monigal K (eds) The Middle Paleolithic of Western Crimea. Vol.2. ERAUL 87, Liège, pp 75–98
Markova AK (2004a) Small mammal fauna from Buran-Kaya III. In: Chabai VP, Monigal K, Marks AE (eds) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea. Vol.3. ERAUL 104, Liège, pp 35–48
Markova AK (2004b) Rodent fauna from the Middle Paleolithic site of Karabi Tamchin. In: Chabai VP, Monigal K, Marks AE (eds) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea. Vol. 3. ERAUL 104, Liège, pp 289–293
Markova AK (2004c) Rodent (Rodentia) fauna from Chokurcha I Unit IV. In: Chabai VP, Monigal K, Marks AE (eds) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea. Vol. 3. ERAUL 104, Liège, pp 371–376
Markova AK (2005) Small mammals from the Palaeolithic site Kabazi II, Western Crimea. In: Chabai VP, Richter J, Uthmeier Th (eds) Kabazi II: last interglacial occupation, environment & subsistence. Palaeolithic sites of Crimea.Vol.1. Simferopol-Cologne: Shlyakh, pp. 51–65
Markova AK (2012) Small mammals from Paleolithic site Siuren I. In: Demidenko YE, Otte M, Noiret P (eds) Siuren I rock-shelter. From Late Middle Paleolithic and Early Upper Paleolithic to Epi-Paleolithic in Crimea. ERAUL 129, Liège, pp 65–71
Marks AE, Monigal K, Usik VI, Ferring CR, Burke A, Rink J, McKinney C (1997) Starosele and the Starosele child: new excavations, new results. Curr Anthropol 38:112–123. https://doi.org/10.1086/204591
Marks AE, Demidenko YE, Monigal K, Usik VI (1998) Starosele: the 1993-1995 excavations. In Marks AE, Chabai VP (eds) The Middle Paleolithic of Western Crimea, Vol. 1. ERAUL 84, Liège, pp 67-100
Massé J, Patou-Mathis M (2009) Siuren I: new zooarchaeological results. In: Chabai VP (ed) Archaeological Almanac. Donetsk, pp 149–158
Massé J, Patou-Mathis M (2012) Zooarchaeological analysis of the faunal assemblage from Siuren I, Crimea (Ukraine). In: Demidenko YE, Otte M, Noiret P (eds) Siuren I rock-shelter. From Late Middle Paleolithic and Early Upper Paleolithic to Epi-Paleolithic in Crimea. ERAUL, Liège, pp. 55–64
Meignen L, Brugal JP (2001) Territorial exploitation, technical traditions and environment in a mid-altitude context: the Canalettes rockshelter (Grands Causses, France). In: Conard N. (ed) Settlement dynamics of the Middle Paleolithic and Middle Stone Age. Kerns Verlag, Tübingen, pp. 463–483
Mihlbachler MC, Rivals F, Solounias N, Semprebon GM (2011) Dietary change and evolution of horses in North America. Science 331:1178–1181. https://doi.org/10.1126/science.1196166
Mikhailesku C (1999) Malacology and paleoenvironments of Western Crimea. In: Chabai VP, Monigal K (eds) The Middle Paleolithic of Western Crimea. Vol.2. ERAUL 87, Liège, pp 99–114
Mikhailesku C (2004) Snails from Karabi Tamchin, Buran-Kaya III and Chokurcha I. In: Chabai VP, Monigal K, Marks AE (eds) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea.Vol.3. ERAUL 104, Liège, pp 299–305
Mikhailesku C (2005) Snail fauna from Kabazi II. In: Chabai VP, Richter J, Uthmeier T (eds) Kabazi II: last interglacial occupation, environment & subsistence. Palaeolithic Sites of Crimea.Vol.1. Simferopol – Cologne: Shlyakh, pp 67–76
Monigal K (2004) Introduction to the site of Buran-Kaya III. In Chabai VP, Monigal C, Marks AE (eds) The Middle Paleolithic and the Early Upper Paleolithic of Eastern Crimea, Vol. 3. ERAUL 104, Liège, pp. 3-18
Musil R (1985) Paleobiography of terrestrial communities in Europe during the Last Glacial. In: Acta Musei Nationalis Pragae. p 83
Nowell A (2010) Defining behavioral modernity in the context of Neandertal and anatomically modern human populations. Annu Rev Anthropol 39:437–452. https://doi.org/10.1146/annurev.anthro.012809.105113
Orlando L, Mashkour M, Burke A et al (2006) Geographic distribution of an extinct equid (Equus hydruntinus: Mammalia, Equidae) revealed by morphological and genetical analyses of fossils. Mol Ecol 15:2083–2093. https://doi.org/10.1111/j.1365-294X.2006.02922.x
Orlando L, Metcalf JL, Alberdi MT, Telles-Antunes M, Bonjean D, Otte M, Martin F, Eisenmann V, Mashkour M, Morello F, Prado JL, Salas-Gismondi R, Shockey BJ, Wrinn PJ, Vasil’ev SK, Ovodov ND, Cherry MI, Hopwood B, Male D, Austin JJ, Hanni C, Cooper A (2009) Revising the recent evolutionary history of equids using ancient DNA. Proc Natl Acad Sci U S A 106:21754–21759. https://doi.org/10.1073/pnas.0903672106
Patenaude B (2010) Faunal exploitation at the Middle Paleolithic site Kabazi II (Western Crimea). PhD dissertation, Université de Montréal
Patou-Mathis M (1999) Archeozoological analysis of the Middle Paleolithic Fauna from selected levels of Kabazi II. In: Chabai VP, Monigal K (eds) The Middle Palaeolithic of Western Crimea. Vol.2. ERAUL 87, Liège, pp 41–74
Patou-Mathis M (2000) Neanderthal subsistence behaviours in Europe. Int J Osteoarchaeol 10:379–395. https://doi.org/10.1002/1099-1212(200009/10)10:5<379: aid-oa558>3.3.co;2-w
Patou-Mathis M (2004) Archeozoological analysis of large mammals of Chokurcha I Unit IV. In: Chabai VP, Monigal K, Marks (eds) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea. Vol. 3. ERAUL 104, Simferopol – Cologne: Shlyakh, pp 355–370
Patou-Mathis M (2005) Analyses Archéologiques des Unités V et VI de Kabazi II. In: Chabai VP, Richter J, Uthmeier Th (eds) Kabazi II: last interglacial occupation, environment & subsistence. Simferopol – Cologne: Shlyakh, pp. 73–98
Patou-Mathis M (2006) Analyse Archéozoologique de l’Unite III. In: Chabai VP, Richter J, Uthmeier Th (eds) Kabazi II: the 70 000 years since the Last Interglacial. Simferopol – Cologne: Shlyakh, pp. 209–240
Patou-Mathis M, Chabai VP (2003) Kabazi II (Crimée, Ukraine): Un site d’abattage et de boucherie du Paléolithique moyen. L’ Anthropol 107:223–253
Péan S, Puaud S, Crépin L et al (2013) The Middle to Upper Paleolithic sequence of Buran-Kaya III (Crimea, Ukraine): new stratigraphic, paleoenvironmental, and chronological results. Radiocarbon 55:1454–1469. https://doi.org/10.2458/azu_js_rc.55.16379
Prat S, Péan SC, Crépin L, Drucker DG, Puaud SJ, Valladas H, Lázničková-Galetová M, van der Plicht J, Yanevich A (2011) The oldest anatomically modern humans from far southeast Europe: direct dating, culture and behavior. PLoS One 6:e20834. https://doi.org/10.1371/journal.pone.0020834
Richter J (2005) Hasty foragers: the Crimea Island and Europe during the Last Interglacial. In: Chabai VP, Richter J, Uthmeier Th (eds) Kabazi II: last interglacial occupation, environment and subsistence. Palaeolithic Sites of Crimea. Vol.1. Simferopol – Cologne: Shlyakh, pp 275–286
Rivals F, Drucker DG, Weber MJ, Audouze F, Enloe JG (2020) Dietary traits and habitats of the reindeer (Rangifer tarandus) during the Late Glacial of Northern Europe. Archaeol Anthropol Sci 12. https://doi.org/10.1007/s12520-020-01052-y
Rivals F, Kitagawa K, Julien MA, Patou-Mathis M, Bessudnov AA, Bessudnov AN (2018) Straight from the horse’s mouth: high-resolution proxies for the study of horse diet and its relation to the seasonal occupation patterns at Divnogor’ye 9 (Middle Don, Central Russia). Quat Int 474:146–155. https://doi.org/10.1016/j.quaint.2018.01.008
Rivals F, Mihlbachler MC, Solounias N (2007) Effect of ontogenetic-age distribution in fossil and modern samples on the interpretation of ungulate paleodiets using the mesowear method. J Vertebr Paleontol 27:763–767. https://doi.org/10.1671/0272-4634(2007)27
Rivals F, Prignano L, Semprebon GM, Lozano S (2015) A tool for determining duration of mortality events in archaeological assemblages using extant ungulate microwear. Sci Rep 5:1–12. https://doi.org/10.1038/srep17330
Rivals F, Schulz E, Kaiser TM (2008) Climate-related dietary diversity of the ungulate faunas from the middle Pleistocene succession (OIS 14-12) at the Caune de l’Arago (France). Paleobiology 34:117–127. https://doi.org/10.1666/07023.1
Rivals F, Schulz E, Kaiser TM (2009) Late and middle Pleistocene ungulates dietary diversity in Western Europe indicate variations of Neanderthal paleoenvironments through time and space. Quat Sci Rev 28:3388–3400. https://doi.org/10.1016/j.quascirev.2009.09.004
Rivals F, Semprebon GM (2017) Latitude matters: an examination of behavioural plasticity in dietary traits amongst extant and Pleistocene Rangifer tarandus. Boreas 46:254–263. https://doi.org/10.1111/bor.12205
Rodríguez-Hidalgo A, Morales JI, Cebrià A, Courtenay LA, Fernández-Marchena JL, García-Argudo G, Marín J, Saladié P, Soto M, Tejero JM, Fullola JM (2019) The Châtelperronian Neanderthals of Cova Foradada (Calafell, Spain) used imperial eagle phalanges for symbolic purposes. Sci Adv 5:1–12. https://doi.org/10.1126/sciadv.aax1984
Sánchez-Hernández C, Gourichon L, Pubert E, Rendu W, Montes R, Rivals F (2019) Combined dental wear and cementum analyses in ungulates reveal the seasonality of Neanderthal occupations in Covalejos Cave (Northern Iberia). Sci Rep 9:1–18. https://doi.org/10.1038/s41598-019-50719-7
Semprebon GM, Godfrey LR, Solounias N, Sutherland MR, Jungers WL (2004) Can low-magnification stereomicroscopy reveal diet? J Hum Evol 47:115–144. https://doi.org/10.1016/j.jhevol.2004.06.004
Semprebon GM, Rivals F (2007) Was grass more prevalent in the pronghorn past? An assessment of the dietary adaptations of Miocene to Recent Antilocapridae (Mammalia: Artiodactyla). Palaeogeogr Palaeoclimatol Palaeoecol 253:332–347. https://doi.org/10.1016/j.palaeo.2007.06.006
Semprebon GM, Rivals F (2010) Trends in the paleodietary habits of fossil camels from the Tertiary and Quaternary of North America. Palaeogeogr Palaeoclimatol Palaeoecol 295:131–145. https://doi.org/10.1016/j.palaeo.2010.05.033
Solounias N, Semprebon GM (2002) Advances in the reconstruction of ungulate ecomorphology with application to early fossil equids. Am Museum Novit 3366:1–49. https://doi.org/10.1206/0003-0082(2002)366<0001: AITROU>2.0.CO;2
Spindler L, Comeskey D, Chabai VP, et al (submitted) Dating the last Middle Palaeolithic of the Crimean Peninsula: new hydroxyproline AMS dates from the site of Kabazi II. J Hum Evol
Stepanchuk VN, Vasilyev SV, Khaldeeva NI, Kharlamova NV, Borutskaya SB (2015) The last Neanderthals of Eastern Europe: Micoquian layers IIIa and III of the site of Zaskalnaya VI (Kolosovskaya), anthropological records and context. Quat Int 428:132–150. https://doi.org/10.1016/j.quaint.2015.11.042
Stiner MC (2013) An unshakable Middle Paleolithic? Trends versus conservatism in the predatory niche and their social ramifications. Curr Anthropol 54:S288–S304. https://doi.org/10.1086/673285
Stringer C (2002) Modern human origins: Progress and prospects. Philos Trans R Soc B Biol Sci 357:563–579. https://doi.org/10.1098/rstb.2001.1057
Uerpmann H-P (2005) Betrachtungen zum Verhältnis zwischen Wildpferd (Equus ferus) und Hydruntinus (Equus hydruntinus) im Jungpleistozän und Holozän auf der Iberischen Halbinsel. Munibe Antropol - Arkeol 351–358
Uthmeier Th (2004) Planning depth and Saiga hunting: on-site and off-site activities of late Neandertals in level B1 of Buran-Kaya III. In: Chabai VP, Monigal K, Marks AE (eds) The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea. Vol.3. ERAUL 104, Liège, pp 175–191
Uthmeier Th (2012) The transition from Middle- to Upper Palaeolithic at Buran Kaya III, Crimea (Ukraine): a case of conceptual continuity of lithic artefact manufacture? In: Pastoors A, Peresani M (eds) Flakes not blades: the role of flake production at the onset of the Upper Palaeolithic. Wissenschaftliche Schriften des Neanderthal Museums 5, Mettmann, pp 239–260
Uthmeier Th (2013) From transformation analysis to land use pattern: the original assemblage of layer IV from Kiik-Koba grotto. In: Demidenko YE, Uthmeier Th (eds) Kiik-Koba Grotto, Crimea (Ukraine). Re-analysis of a key site of the Crimean Micoquian. Kölner Studien zur Prähistorischen Archäologie. Vol. 3. Leidorf, Rahden, pp. 129–160
Uthmeier Th, Chabai VP (2010) Neanderthal subsistence tactics in the Crimean Micoquian. In: Conard N., Delagnes A (eds) Settlement dynamics of the Middle Paleolithic and Middle Stone Age. Vol. III. Kerns Verlag, Tübingen, pp. 195–234
Uthmeier Th, Chabai VP (2018) Formation processes at sites with high-resolution sequences in the Crimean Middle Paleolithic: the Kabazi V rock shelter and the open-air site of Kabazi II compared. Quat Int 485:44–67. https://doi.org/10.1016/j.quaint.2018.01.017
Uthmeier Th, Chabai VP, Veselsky AP (2020) From the apron into the pit: the deposition of the complete debris from the manufacture of a bifacial preform at the middle Paleolithic site of Kabazi V, level III/4-2. Archaeol Anthropol Sci 12. https://doi.org/10.1007/s12520-019-00968-4
van Asperen EN (2010) Ecomorphological adaptations to climate and substrate in late Middle Pleistocene caballoid horses. Palaeogeogr Palaeoclimatol Palaeoecol 297:584–596. https://doi.org/10.1016/j.palaeo.2010.09.007
van Asperen EN, Stefaniak K, Proskurnyak I, Ridush B (2012) Equids from Emine-Bair-Khosar Cave (Crimea, Ukraine): co-occurrence of the stenonid Equus hydruntinus and the caballoid E. ferus latipes based on skull and postcranial remains. Palaeontol Electron 15: https://doi.org/10.26879/280
Vereshchagin N, Baryshnikov G (1980) Mammals of the northern Crimea in the Paleolithic. Tr Zool Instituta Akad Nauk SSSR 93:26–49
Wynn T, Coolidge FL (2004) The expert Neandertal mind. J Hum Evol 46:467–487. https://doi.org/10.1016/j.jhevol.2004.01.005
Xafis A, Nagel D, Bastl K (2017) Which tooth to sample? A methodological study of the utility of premolar/non-carnassial teeth in the microwear analysis of mammals. Palaeogeogr Palaeoclimatol Palaeoecol 487:229–240. https://doi.org/10.1016/j.palaeo.2017.09.003
Yevtushenko AI, Burke A, Ferring CR (2004) The site of Karabi Tamchin: introduction. In Chabai V, Monigal K, Marks AE (eds) The Paleolithic of Crimea. The Middle Paleolithic and Early Upper Paleolithic of Eastern Crimea, Vol. 3. ERAUL 104, Liège, pp. 277-282
Yevtushenko A, Chabai VP (2012) Karabai I, the Paleolithic site in Eastern Crimea. Donetsk, Donbass
Zilhão J (2006) Neandertals and moderns mixed, and it matters. Evol Anthropol 15:183–195. https://doi.org/10.1002/evan.20110
Funding
The research of FR and IRP is supported by projects 2017 SGR 836 from the Generalitat de Catalunya, project HAR2016-76760-C3-1-P and PID2019-103987GB-C31 from the Ministerio de Ciencia e Innovación and the CERCA Program/Generalitat de Catalunya. The sampling of teeth from the Crimean sites of Kabazi II, Siuren I and Chokurcha I at Institut de Paléontologie Humaine of the Muséum National d’Histoire Naturelle (MNHN) in Paris was funded by a grant of the Deutsche Forschungsgemeinschaft (DFG) ‘The dispersal of modern humans into an Eastern European refugial area of late Neanderthals: interdisciplinary studies of contemporaneous industries from the Middle to Upper Palaeolithic transition in the Crimea (Ukraine)’ (UT 41/2-1) given to Thorsten Uthmeier. The sampling of extant saiga specimens at the Naturhistorisches Museum in Vienna was supported by the European Commission’s Research Infrastructure Action via the SYNTHESYS Project AT-TAF-4385 to FR.
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Ramírez-Pedraza, I., Rivals, F., Uthmeier, T. et al. Palaeoenvironmental and seasonal context of the Late Middle and Early Upper Palaeolithic occupations in Crimea: an approach using dental wear patterns in ungulates. Archaeol Anthropol Sci 12, 268 (2020). https://doi.org/10.1007/s12520-020-01217-9
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DOI: https://doi.org/10.1007/s12520-020-01217-9