Supplementary Information for

Archaeological evidence for two separate dispersals of into southern Siberia

Kseniya A. Kolobova, Richard G. Roberts, Victor P. Chabai, Zenobia Jacobs, Maciej T. Krajcarz, Alena V. Shalagina, Andrey I. Krivoshapkin, Bo Li, Thorsten Uthmeier, Sergey V. Markin, Mike W. Morley, Kieran O’Gorman, Natalia A. Rudaya, Sahra Talamo, Bence Viola and Anatoly P. Derevianko

Kseniya A. Kolobova, Richard G. Roberts Email: [email protected], [email protected]

This PDF file includes:

Supplementary text Figures S1 to S26 Tables S1 to S33 SI References

www.pnas.org/cgi/doi/10.1073/pnas.1918047117 1

SUPPLEMENTARY INFORMATION

Supplementary text Section S1  Description of the and stratigraphic sequence ……………………………… 3 Section S2  Sediment micromorphology analyses ……………………………………………. 6 Section S3  Palynology, palaeontology and Pleistocene environments ………………………. 9 Section S4  Radiocarbon and optical dating …………………………………………………. 10 Section S5  Palaeoanthropological data ………………………………………………………. 15 Section S6  Lithic assemblage from subunits/sublayers 6a–6c/2 …………………………….. 19 Section S7  Comparison with other Altai Middle Palaeolithic assemblages …………………. 21 Section S8  Comparison with European Micoquian assemblages ……………………………. 24 Section S9  Timing and routes of migrations …………………………………… 29 Figures S1 to S26 ………………………………………………………………………………… 31 Tables S1 to S33 ………………………………………………………………………………….. 57 References ………………………………………………………………………………………… 94

2

Section S1. Description of the cave and stratigraphic sequence (M.T.K.)

Chagyrskaya Cave (51° 26′ 34.6′′ N; 83° 09′ 18.0′′ E) is situated on the left bank of the Charysh River in the Tigirek Range in the foothills of the (Fig. S1). The cave faces north and is situated at an elevation of 353 m above sea level, about 19 m above the river level. It consists of two chambers with a total area of ~130 m2. The stratigraphic sequence (up to 3.5 m thick) includes both Holocene (lithoseries III) and Pleistocene sediments (Derevianko et al., 2013a). The Pleistocene deposits can be subdivided into a lower and upper part (lithoseries I and II), reflecting different sedimentation processes. The upper part (layer 5 and subunits 6a–d) is composed mostly of sub-aerial deposits, including loess-like sediments. The lower part (layers 7 and 8) comprises dense loamy sediments with quartz grains.

Lithoseries I – clays and gravels Layer 8 – red clay occurring locally in depressions in the bedrock. This sediment is preserved as small remnants that survived erosional events in pocket-like structures. The red clay is a typical weathered sediment (terra rossa type) that has accumulated as a residual material during karst dissolution of the limestone, most probably during warm pre-Pleistocene climatic phases. Layer 7 – red-brown (dry 7.5YR 6/7 – reddish yellow, moist 7.5YR 4/4 – brown/dark brown) clay or clayey loam, with quartz grains and fine strongly chemically weathered limestone clasts and riverine pebbles. Intercalations of greenish silt occur locally. This sediment is lying on the bedrock. The presence of pebbles and red clay indicates a complex origin of the layer. The pebbles probably originate from old alluvial terraces located above the cave, and were transported into the cave via chimneys, by colluvial processes. The red clay is a typical weathered sediment (terra rossa type), accumulated as a residual material during karst dissolution of the limestone. We assume that the alluvial sediments and the weathered clays are not contemporaneous, and were secondarily deposited together by colluvial activity. The complex inner structure of layer 7 is reflected in its division into subunits 7a, 7b and 7c (based on differences in colour and proportion of pebbles) during the excavations led by S.V.M. The maximum thickness of this layer is 100 cm.

Lithoseries II – silty sediments Layer 6 is a complex series composed of subunits 6d, 6c, 6b and 6a. Subunit 6d (Russian 6г) is a reddish-brown (dry 7.5YR 6/7 – reddish yellow, moist 7.5YR 4/4 – brown/dark brown) loam with fine weathered limestone clasts, sparse bones and fine riverine pebbles. Maximum thickness is 10 cm. This subunit contains clasts and packets of layer 7 mixed with sediment similar to subunit 6c, as a result of frost action. Some vertical rearrangement of sediments is confirmed by the presence of a diapir, which locally lifted the sediments of layer 7 and subunit 6d by ~30 cm, up to the elevation of subunit 6a (Fig. S2C). The spatial distribution of the diapir deforming layers 7 and 6 testifies to the localised plastic deformation of sediments after deposition of subunit 6a (i.e., long after Palaeolithic occupation). Subunit 6c (Russian 6в) is a gray carbonate silty loam with sparse fine riverine rounded pebbles, numerous bone fragments, lithic artefacts and sparse limestone clasts. Locally, the subunit has a complex structure and may be subdivided into two sublayers: 6c/1 (more brownish colour: dry 10YR 6/3 – pale brown, moist 10YR 4/3.5 – brown/dark brown/dark yellowish brown) and 6c/2 (more grayish to greenish colour: dry 10YR 6/4 – light yellowish brown, moist 10YR 3.5/2.5 – dark grayish brown/very dark grayish brown). Sublayer 6c/2 is a loess-like sediment, while sublayer 6c/1 resembles a palaeosol developed on the loess; both sublayers contain Middle Palaeolithic artefacts and fossil bones. Subunit 6c is plastically deformed by cryoturbation, similar to layer 7 and subunits 6d and 6a. The total primary thickness of subunit 6c remains unknown as it is cut by the erosional feature at the bottom of subunit 6a or, locally, layer 5. The maximum thickness of subunit 6c is 60 cm, but this includes convolutions produced by cryoturbation. The lower boundary of subunit 6c and the inner boundary between sublayers 6c/1 and 6c/2 are clear and marked by a colour change. The morphology of the diapir of layer 7 and subunits 6d and 6c is repeated in the form of plastic

3 deformation of the overlying sediments, which we interpret as evidence that cryoturbation occurred after deposition of subunit 6a (i.e., long after Middle Palaeolithic occupation). We did not, however, observe mixing of material between strata, except for subunit 6d, which incorporates material from layer 7 and sublayer 6c/2. This means that the sediments of subunit 6c have not been mixed with those of the overlying layers. Subunit 6b (Russian 6б) – this layer (dry 10YR 6/3 – pale brown, moist 10YR 4/3 – brown/dark brown) is known from previous excavations led by S.V.M. Based on the preserved sections from the earlier excavations, however, subunits 6b and 6a appear to form a complex colluvial series, built of more than two interbedded sedimentary units: 6b is more silty and brown, similar to sublayer 6c/1, while 6a is more clayey and orange, similar to layer 7. Subunits 6b and 6a are similar, but the sediments in subunit 6b are slightly denser, less porous and have a darker colour. Both subunits contain Middle Palaeolithic artefacts and fossil bones, and the entire subunit 6a/6b complex is more than 100 cm thick. Subunit 6a (Russian 6а) is a light brown (dry 10YR 7/3 – very pale brown, moist 10YR 4/4 – dark yellowish brown) carbonate silt with sparse angular limestone clasts, bone fragments, lithic artefacts and riverine pebbles. Subunit 6a is up to 60 cm thick, with a lower boundary that is erosional, undulating and inclined towards the cave entrance. Colluvial sediments of this subunit were transported northward by cohesive flow, from the inner part of the cave towards the cave mouth. Subunit 6a contains Middle Palaeolithic artefacts similar to those in subunit 6c, which suggests that subunit 6c served as a source of material for colluvial flow. Other sediments were also incorporated into this flow, as indicated by the lithological dissimilarity between subunits 6c and 6a. The latter subunit has survived as localised erosional remnants, with most of it eroded prior to the erosional event preceding the deposition of layer 5. Layer 5 is a yellowish (dry 10YR 7/3 – very pale brown, moist 10YR 4/4 to 5/6 – dark yellowish brown to yellowish brown) carbonate silt with limestone debris. From a sedimentological point of view, this layer is a complex of strata, composed of two types of sediment that may be regarded as separate subunits, here designated as subunit 5a (a silt with sparse, rounded pebbles and sparse, angular limestone clasts) and subunit 5b (limestone debris comprising angular clasts up to 0.5 m in size, with a silty matrix, but commonly without any fine material in intergranular spaces, indicating very rapid accumulation). Subunit 5a represents cohesive colluvial fill of loess-like sediments redeposited in erosional channels formed by flowing water. These erosional features are several decimeters in depth and have rounded basal profiles cut either into older sediments (usually subunit 6a) or into subunit 5b. Subunit 5b consists of rock fall, most probably triggered by seismic events, but preceded by intensive mechanical weathering (frost action). Subunits 5a and 5b can be clearly distinguished wherever they occur in juxtaposition. The entire layer 5 complex is up to 110 cm thick, with sharp and erosional boundaries at the base of the layer and between the subunits. Artefacts and fossil bones are less numerous in layer 5 than in layer 6. Layer 4 is a local variety of subunit 5a and has a more grayish colour.

Lithoseries III – sandy loams with riverine pebbles Layer 3 – grayish-brown (dry 2.5Y 5/1 – gray/grayish brown, moist 10YR 4/4 – very dark brown) loamy sand with abundant riverine rounded pebbles of variable lithology (sedimentary, igneous and metamorphic rocks). The pebbles and sand most probably derive from old river terraces situated on the slope above the cave, and were transported into the cave by colluvial processes via the karstic chimneys in the ceiling of the rear chamber. Fluvial activity can be excluded as a direct depositional agent, due to high elevation of the cave above the river bed and the poor sorting of sediments by grain size. The numerous archaeological finds in this layer and its grayish colour testify to the cultural character of the sediment. Layer 3 is up to 35 cm thick and has a clear lower boundary. Layer 2 – yellowish brown (dry 10YR 7/3 – very pale brown, moist 10YR 2/2 – dark yellowish brown) loamy sand. This layer is similar to layer 3, except that it is of a more yellowish colour. The imbrication of the pebbles is clearly visible in the longitudinal profiles, indicating the

4 transport direction northward from the cave interior towards the entrance. Solifluction (sediment creep under cold conditions) is the depositional process. Layer 2 is up to 60 cm thick and has a blurred lower boundary marked by a colour change. Layer 1 – gray to dark gray non-carbonate loamy sand, slightly compacted, with many small fluvial pebbles. This layer is up to 5 cm thick and has a clear lower boundary. Layer 1 represents the uppermost part of layer 2, altered by the input of organic matter and the effects of human trampling.

Stratigraphic position of the Middle Palaeolithic level Subunit 6c (and its sublayers 6c/1 and 6c/2) can be regarded as the primary depositional context of the Middle Palaeolithic assemblage at Chagyrskaya Cave. The occurrence of bones and lithic artefacts in subunit 6d is the result of post-depositional displacement due to frost action. Although signs of cryoturbation are also evident in subunit 6c, this process involved small-scale freezing and thawing of the sediments. Large-scale disturbances are limited to plastic deformation in the form of a diapir, which did not result in the mixing of sediments, and was easy to identify during excavation due to lithological differences between strata. The presence of Middle Palaeolithic artefacts in the overlying deposits (subunits 6b and 6a and layer 5) is the result of erosion of subunit 6c followed by redeposition of the sediments, bones and artefacts via colluvial processes.

5

Section S2. Sediment micromorphology analyses (M.W.M., M.T.K.)

Sample collection Three sediment blocks for micromorphological analyses (MM2, MM3 and MM4) were collected by M.W.M. in 2014 from two sections towards the rear of the cave, where Layers 5–7 were exposed during excavations led by S.V.M. (Figs S2A and S3E–G). These sections was chosen to maximise the potential to record an environmental signal, as hominin activities appear to have been concentrated near the mouth of the cave and taper off inside (Vasiliev, 2013). The blocks were extracted from key parts of the stratigraphic sequence, targeting interfaces between adjacent layers:  MM2: bottom of layer 5 and top of subunit 6a, east face of square M12.  MM3: bottom of layer 6a, all of subunit 6b and top of sublayer 6c/1, east face of square M12.  MM4: bottom of sublayer 6c/1 and top of layer 7, south face of square H12. In 2017, six further sediment blocks were collected by M.T.K. from newly exposed profiles nearer to the front of the cave, and one block was taken from the profile exposed previously towards the rear of the cave (Figs S2A and S3A–D). These sampling locations were selected for their proximity to the sediment samples collected for optical dating during the same field season. These blocks targeted the following parts of the sequence:  2969: middle part of subunit 5a, east face of square И8.  2970: bottom part of subunit 5a, east face of square И7.  2987: middle part of sublayer 6c/1, south face of square К8.  2985: middle part of sublayer 6c/2, mid part of square К7.  2988: middle part of subunit 6d, south face of square К8.  2984: upper part of layer 7, mid part of square К7.  2989: bottom of greenish intercalation within layer 7, south face of square Н11.

Sample preparation The sediment blocks were impregnated with resin, with two thin sections made from each of the 2014 blocks (distinguished by suffixes A and B) and one thin section from each of the 2017 blocks, following procedures described elsewhere (e.g., Morley et al., 2017, 2019). Diagnostic features observed using a polarizing microscope were recorded for each layer using standard protocols (Stoops, 2003).

Observations and interpretations Micromorphological (microstratigraphic) analysis of archaeological sequences can help elucidate the processes responsible for site formation, the depositional and post-depositional environments, and the context of archaeological objects and features and their interpretation (Goldberg and Berna, 2010; Mallol and Mentzer, 2017). Our results indicate often subtle changes in the depositional environment related to the use of the cave by animals (including hominins) and local changes in temperature and humidity. Detailed sediment descriptions and earlier micromorphological analyses were published by Derevianko et al. (2013a), who also reported a range of useful palaeoenvironmental indicators. The analyses reported here broadly concur with their findings. Summary descriptions of the thin section observations and interpretations are given in Table S1, and a selection of photomicrographs showing the key features is provided in Fig. S4. Layer 7 marks the base of the analysed part of the stratigraphic sequence. The top of this layer is a compact silty clay, with variable quantities of quartz sand and occasional rock fragments (mainly weathered metamorphic and volcanic lithologies, such as basalt and schist) (Fig. S4A). A notable characteristic is the common occurrence of angular, sub-rounded and rounded sand-sized clay aggregates, reworked fragments of clay coatings, and sand grains with clay pendant coatings

6

(Fig. S4B). These features indicate a reworking of older sediments into the sediment matrix of this layer, possibly with variable residence times in the karstic system. Different lithologies are represented in these aggregates, with variable colours and internal compositions, suggesting multiple sources for these features. Rock fragments with clay cappings and coatings indicate the vertical movement of fine sediments, but the partial preservation of some of these coatings indicates that the clay particles were deposited prior to erosion and redeposition. We attribute these features to changes in climatic conditions affecting site hydrology. These observations are compatible with humid conditions at the time of accumulation, followed by an erosional event (or events) caused by sheetwash or changes in karstic hydrology. Locally, the sediment is intercalated with greenish dense clay with rounded sand-sized grains of varying lithologies, probably connected with episodes of increased chemical weathering during warmer and wetter climatic phases. Subunit 6d exhibits a mixture of the characteristics of both layer 7 and sublayer 6c/2. The sediment is porous, composed of compact clay-silt aggregates, and is brownish in plane polarised light (ppl). These features are shared with sublayer 6c/2, but subunit 6d also contains fragments of orange clay aggregates or clay coatings, which are more typical of layer 7 (Fig. S4C,D). The chaotic orientation of elongated grains and aggregate microstructure indicates frost action as a process responsible for the mixing of the sediments. Sublayer 6c/2 is a variably compacted clay silt with a weakly developed aggregate microstructure. Numerous coprolite and bone fragments are present, consisting of coprolites and bones, which contribute around 10% of the total sediment volume. Bone fragments are usually rounded, most probably due to corrosion in carnivore digestive tracts. The large mammal data show that the dominant carnivores in the cave were wolf and (Vasiliev, 2013). Some bone fragments are fractured in situ forming fine angular pieces retaining close association (Fig. S4E). Such features are produced by post-depositional mechanical frost weathering (Krajcarz and Krajcarz, 2019). Together with the vertical orientation of some bone fragments and lithic artefacts (Fig. S4F), this suggests intensive frost action and cryoturbation. Coprolites are rounded, usually yellow with a brown center or brown ‘rinds’, or brown. Given their morphology, they are most likely of hyena origin (Horwitz and Goldberg, 1989; Larkin et al., 2000; Carrión et al., 2007). Inclusions are rare, but many have vesicular voids inside, formed by gas bubbles. The coprolites also contain corroded bones, thereby linking them to bone eaters, such as hyenas. No traces of clay aggregates, as recorded for the underlying strata (layer 7 and subunit 6d), were detected in sublayer 6c/2. This indicates that the sediments of this sublayer were derived from an external source, and that material from layer 7 and subunit 6d was not incorporated into this sublayer by post- depositional processes. The existence of hyena coprolites and bones, as well as bones digested by hyenas, clearly indicates that these carnivores were present in the cave at times and might have caused local disturbance in places. Although the sublayer 6c/2 assemblage could, therefore, have been bioturbated, no older assemblages are known from the cave and there is no micromorphological evidence for mixing of these sediments with those of the overlying layers. In addition, the optical ages indicate that sublayer 6c/2 accumulated over a short time interval (a few centuries or millennia at most; see Section S4), which further reduces the probability that the assemblage in sublayer 6c/2 is associated with multiple Neanderthal occupations. The micromorphological and chronological data thus support the stratigraphic integrity of this sublayer and its associated assemblage. Sublayer 6c/1 is variably compacted, becoming increasingly porous towards the upper contact with subunit 6b, possibly in association with bioturbation of its upper surface (Fig. S4G,H). In parts of this sublayer, the fine sediment matrix has a ‘granular’ microstructure (Fig. S4H), with distinctive birefringent clay alignments around the outer rim of these round and ovoid arrangements (Fig. S4I). These features are a sign of stress caused by the hydraulic force of ice expansion within the sediment (Van Vliet-Vanöe, 2010). Climatic deterioration and a shift towards cooler temperatures is most likely associated with frequent cycles of freezing and thawing, commonly recorded in the upper parts of soils (Cremaschi and Van Vliet-Lanöe, 1990; Van Vliet-Lanöe, 2010). These observations support inferences of a cold and dry climate and steppe landscape drawn

7 from the pollen and large mammal assemblages in layer 6 (Rudaya, 2013; Vasiliev, 2013; Rudaya et al., 2017). An important feature is the occasional occurrence of rounded clay aggregates with a dense structure, clay-silt lithology and brownish colour typical of sublayer 6c/2. This indicates that sublayer 6c/2 was eroded and redeposited, probably by sheetwash, serving as a source of material for sublayer 6c/1. This sublayer has low numbers of rounded coprolite fragments, indicating sporadic use of the cave by non-hominin animals (most likely carnivores, based on the morphological characteristics of these droppings). The coprolite fragments are generally very fine- grained, yellow in colour in ppl, and exhibit a darker brown ‘rind’; these features are compatible with published descriptions of hyena coprolites. The frequency of inclusions is generally low, but occasional vesicles (gas bubbles) and other organic inclusions (e.g., hair/fur) are present in some of the coprolites. The rounded shape and fragmentary state of the coprolites supports the idea that this sublayer is composed of materials originally deposited in sublayer 6c/2, so we consider that both sublayers contain the same artefact and faunal assemblages. Subunit 6b has a well-developed granular microstructure (Fig. S4J,M), also recorded in sublayer 6c/1, causing the fine fabric to separate into ovoid or round aggregates (Van Vliet-Lanöe, 2010). Freezing of the sediments is also apparent in the mechanically cracked and fractured clay aggregates and occasional coprolite fragments, which were also noted in the microscopic fracture patterns of quartz grains observed under a scanning electron microscope (Derevianko et al., 2013a). In some localised areas, finely laminated, limpid clay fragments might relate to the cracking, reorganisation and incorporation of clay crusts and caps by expansion and contraction (e.g., FitzPatrick, 1993). Large fragments of bone that show signs of breakage in situ might relate to trampling of the sediments by large animals (Fig. S4K), possibly hominins. Coprolites are present in low numbers (Fig. S4L), but with localised regions where frequencies are much higher, suggesting continued sporadic use of the cave by carnivores. Subunit 6a, towards the top of the studied sequence, shows a marked increase in the frequency of coprolites (Fig. S4N–P). The high intensity of use of this part of the cave by carnivores (most likely hyenas) is indicated by the increase in small bones displaying acid etching, and the absence of human-modified bones (Vasiliev, 2013; Rudaya et al., 2017). Freezing of the sediments is evident in the granular microstructure and occasional b-fabric associated with mineral and composite aggregate grains (Fig. S4Q). These features are not as well-developed as in subunit 6b and sublayer 6c/1, however, which might indicate climatic amelioration during this aggradational phase. Layer 5 marks the top of the studied sequence. The base of this layer bears similarities to subunit 6a, but with an increase in coarse mineral grains and rock fragments. An increase in rounded clay aggregates (‘rip up clasts’) and the more poorly-sorted composition of these sediments could signify a slight increase in water availability at this time. Minor signs of diagenesis in the chemically-modified speleothem and limestone fragments lend support to this interpretation, as does the presence of clay aggregates that may be related to water erosion. Microstructural features indicative of freezing conditions are infrequent and weakly expressed (suggesting a warmer environment) in the parts of the sequence sampled in 2014, whereas the sediments sampled in 2017 exhibit distinctive frost-related microstructures with ovoid aggregates (Van Vliet-Lanöe, 2010). Coprolite fragments are very common, often with brownish clay-silt coats, indicating redeposition from older sediments (Fig. S4R).

8

Section S3. Palynology, palaeontology and Pleistocene environments (N.A.R.)

The environmental reconstruction for the period of Neanderthal occupation of the Charysh River valley is based on pollen and palaeontological data from Chagyrskaya Cave (Derevianko et al., 2013a; Rudaya, 2013; Vasiliev, 2013; Rudaya et al., 2017; this study). These palaeoclimate proxies support the assignment of layers 6 and 5 to the end of Marine Isotope Stage (MIS) 4 and the onset of MIS 3. The absence of tundra components in the pollen spectra, and the low frequency of tundra species among the small and large mammals, reflect a relatively warm climate in the Charysh valley in comparison to the West Siberian Plain during the MIS 4 stadial (Volkova and Kulkova, 1984). Ecological-niche modelling (Glantz et al., 2018) also suggests that the Altai foothills likely acted as a hominin refugium during the , given the milder climate compared to the adjacent plain. Subunit 6a yielded only two molars of the Ob lemming (Lemmus sibiricus). The scarcity of these rodents, which are restricted today to subarctic regions (Derevianko et al., 2013a), reflects this warmer climate, but the remains of reindeer (Rangifer tarandus) in layer 5 and subunits/sublayers 6a–6c/2 (Vasiliev, 2013; Rudaya et al., 2017) indicate that cold conditions also prevailed at times. The presence of arid steppe landscapes during Neanderthal settlement of the Charysh valley is also confirmed by the finding of solitary remains of the yellow steppe lemming, Eolagurus luteus, a species uncommon in the modern Altai fauna. Bones of Eolagurus luteus were recorded in all layers, with higher frequencies in layer 5 and subunits 6a and 6b. At the present day, this animal inhabits desert steppes in eastern Kazakhstan (Lake Zaysan region), , and . Layer 5 and subunits 6a and 6b also contained the remains of the desert-steppe species Allactaga major (great jerboa), which is also unusual in the modern Altai fauna (Rudaya et al., 2017). The bird fauna of layer 6 is typical of the late Pleistocene bird composition of the northwestern Altai and includes, for example, Lagopus lagopus and Corvus corax (Martynovich et al., 2016). The occurrence in layer 6 of Syrrhaptes paradoxus, a species found in the modern dry steppes and semi deserts of Central Asia, Kazakhstan, Mongolia and the Volga region, is additional evidence for arid conditions in the Charysh valley in the final stages of MIS 4. Fossil plant taxa and the faunal composition suggest that steppe or semi-desert steppe had spread under a dry continental climate in the Charysh valley at the end of MIS 4. The palaeontological and palynological data from layer 5 reveal a complex environment at the start of MIS 3: steppe and forest-steppe developed in a relatively warm and humid climate, supplemented by dark coniferous and mixed birch coniferous forest in the river valleys (Rudaya et al., 2017). Palaeolithic occupation of Chagyrskaya Cave is associated with a high density of bones. A total of 186,688 specimens of Pleistocene fauna were recovered during the 2007–2013 excavation seasons from the deposits extending from layer 5 to sublayer 6c/2 (Rudaya et al., 2017). The taphocoenosis of layer 5 and subunit 6a was formed as a result of the feeding activities of large carnivores, and the taphocoenosis of subunits/sublayers 6b–6c/2 resulted from human activity. A detailed description of the bone assemblage from the various stratigraphic layers at Chagyrskaya Cave is given in Vasiliev (2013). Neanderthal hunting activity was focused on bison (Bison priscus), with abundant remains recovered from subunits 6b and 6c (up to 49.75% in the latter). Bison hunting may have been seasonal and connected to the annual migration of Bison priscus herds between the Altai piedmont lowlands and the mountainous interior. Juveniles and females were preferred as prey (Vasiliev, 2013), as they would have been easier game than adult males. Other prey hunted included the Ovodov horse (Equus (Sussemionus) ovodovi), Siberian mountain goat (Capra sibirica), (Ovis ammon) and reindeer, albeit to a much lesser degree than bison. Palaeontological remains from layer 5 and subunit 6a mostly demonstrate hunting activity by wolves and cave hyenas (Vasiliev, 2013).

9

Section S4. Radiocarbon and optical dating (R.G.R., Z.J., B.L., K.O’G., S.T.)

Radiocarbon dating Radiocarbon (14C) ages have been obtained for 20 Bison sp. remains recovered from layers 5 and 6 (Table S2). Ten, possibly 13, of the bones have cut or impact marks made by humans using stone tools (Derevianko et al., 2013a; Rudaya, 2013; Rudaya et al., 2017). These humanly modified bones were recovered from subunit 6b and sublayer 6c/1, which also yielded almost all of the Neanderthal remains. Ages were obtained on collagen extracted from these samples (laboratory code S-EVA) at the Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology (MPI-EVA) in Leipzig, , using the pretreatment procedures described by Talamo and Richards (2011). The pretreated samples were ultrafiltered to isolate the >30 kDa fraction and remove contaminants with lower molecular weights. This results in more accurate ages, particularly for samples older than ~30 ka (Brown et al., 1988; Talamo and Richards, 2011; Brock et al., 2013; Wood, 2015). Stable isotope ratios, C:N atomic ratios, and collagen yields were measured to determine the extent of collagen preservation. The stable isotope analyses were made using a Thermo Scientific Flash Elemental Analyzer, coupled to a Delta V isotope ratio mass spectrometer. Bones with >1% weight collagen and C:N ratios in the range 2.9–3.5 are commonly considered to have passed the evaluation criteria for collagen to proceed to accelerator mass spectrometry (AMS) analysis (van Klinken, 1999). Samples were graphitised and the 14C contents measured by AMS at the Mannheim facility (laboratory code MAMS; Kromer et al., 2013). The measured (conventional) ages listed in Table S2 have been calibrated, and their 68.2% and 95.4% confidence intervals estimated, using the IntCal13 data set (Reimer et al., 2013) and OxCal v4.3 (Bronk Ramsey, 2009). Sample MAMS-14962 failed both of the quality-assurance criteria (C:N ratio of 3.6 and a collagen yield of <1%), so we consider the resulting age (17,630 ± 50 yr BP, ~21 ka cal. BP) to be unreliable. We note that this age estimate is also discrepant with the other 12 ages obtained for sublayer 6c/1 (Table S2). The only other sample with an obviously anomalous age for its stratigraphic position, and that also yielded <1% collagen, is MAMS-14956 (4,497 ± 26 yr BP, ~5 ka cal. BP), which we attribute to the incorporation of this bison phalanx into layer 5 as a result of post-depositional disturbance. Of the remaining 18 samples, only four produced finite ages, eight have ages of >49 ka BP and a further six have ages of >52 ka BP. The youngest of the four finite ages is 33,760 ± 170 yr BP (~38 ka cal. BP) for MAMS-14954, which was collected from the uppermost horizon of layer 5. This sample may reflect the true age of this horizon, or it may have been incorporated subsequently into this horizon from ~38 ka cal. BP deposits that have since been eroded. The other three finite ages lie at the limits of reliable 14C dating and have calibrated ages of >47 ka cal. BP. These bones were collected from horizons 1 (MAMS-13033), 2 (MAMS-13034) and 3 (MAMS-13035) of sublayer 6c/1. We view these ages as minimum estimates of the true age, given that infinite ages were obtained for several other samples from horizons 1 and 3 in sublayer 6c/1 and from the stratigraphically overlying deposits (subunits 6b and 6a and layer 5). The incomplete removal of all sources of younger carbon during sample pretreatment can readily account for these apparent finite ages, as contamination of a 60 ka BP sample with just 0.5% modern carbon will result in a measured age of ~42 ka BP (Wood, 2015). The fourteen infinite ages of >49 and >52 ka BP obtained from layers 5 (horizon 2) and 6 indicate that these bison remains date to early MIS 3 or a preceding period, but alternative dating methods are needed to obtain finite ages for these deposits.

Optical dating Depositional ages for layers 5, 6 and 7 have been obtained for 27 sediment samples using optical dating procedures for sand-sized grains of potassium-rich feldspar (K-feldspar). Optical dating yields estimates of the time elapsed since the grains were last exposed to sunlight (Huntley et al., 1985; Hütt et al., 1988; Roberts et al., 2015). Ages are calculated in calendar years and are

10 estimated by dividing the sample equivalent dose (De, a measure of the radiation energy absorbed by grains during their period of burial) by the environmental dose rate (the rate of supply of ionizing radiation to the grains over the same period). The De is determined from laboratory measurements of the infrared stimulated luminescence (IRSL) emitted by the K-feldspar grains, and the dose rate is estimated from field and laboratory measurements of environmental radioactivity from the 238U, 235U and 232Th decay series and 40K, the internal 40K and 87Rb content of the grains, and the small contribution from cosmic rays. The sample De values and optical ages are listed in Table S3, along with the supporting dose rate data. Nine samples were collected in 2012 (denoted as CHAG12-), eleven in 2014 (CHAG14-) and seven in 2017 (CHAG17-) from the stratigraphic units listed in Table S3. Roberts et al. (2018) reported optical ages and supporting data for the 2012 samples, together with details of the sample collection, preparation, measurement and data analysis procedures. All but two of the 2012 and 2014 samples were collected from near the rear of the cave: CHAG12-1 to -9 from the south face of squares M11 and H11, and CHAG14-2 to -11 from the south face of square M12. CHAG12-10 and CHAG14-12 were collected from closer to the cave entrance (both from the east face of square Л7), and the 2017 samples were collected from this same general area (shaded yellow in Fig. S2A): the south faces of squares И6 (CHAG17-3 and -4), И8 (CHAG17-5 and -6) and К8 (CHAG17-7 to -9). The sampling locations are shown in Fig. S2A. Samples were collected at night (using dim red light for illumination), sealed in plastic bags and wrapped in black plastic to prevent light exposure during transport to the University of Wollongong. These sediment samples were used for IRSL measurements, to estimate the field water contents and to make laboratory measurements of the beta dose rate. Measurements of the in situ gamma dose rate were made at each sample location using a portable gamma-ray detector. In the laboratory, sand-sized grains of K-feldspar were extracted from the samples under dim red illumination using standard procedures (Aitken, 1998). Each sample was sieved to isolate grains of 180–212 µm diameter (or 125–150 µm for CHAG17-4 and 90–125 µm for some of the CHAG14- 10 analyses), which were treated with solutions of 10% hydrochloric acid and 30% hydrogen peroxide to remove carbonates and organic matter, respectively. CHAG12-8 yielded too few grains to proceed further. For the other 27 samples, K-feldspar grains were separated from quartz and heavy-mineral grains using solutions of sodium polytungstate and then etched in 10% hydrofluoric acid for 40 min (to clean the grain surfaces and remove, or greatly reduce in volume, the alpha- irradiated rinds), rinsed in hydrochloric acid (to remove any precipitated fluorides) and, finally, dried and sieved again. Measurements of the beta dose rate were made on dried, homogenised and powdered portions of each sample using a low-level beta counting system (Bøtter-Jensen and Mejdahl, 1988) and the data-analysis procedures described in Jacobs and Roberts (2015). We used a 1-inch diameter NaI(Tl) detector and the ‘threshold’ technique to estimate the gamma dose rates from the U and Th decay series and 40K (Mercier and Falguères, 2007), with the detector calibrated using the doped concrete blocks at Oxford (Rhodes and Schwenninger, 2007). Cosmic-ray dose rates were estimated following Prescott and Hutton (1994), taking into account the latitude, longitude and altitude of the site, the thickness and density of sediment overburden and bedrock shielding, and the zenith angle dependence of cosmic rays. The beta, gamma and cosmic-ray dose rates were calculated assuming a long-term water content of 20 ± 5% (at 1σ), which accommodates at 2σ the range of measured (field) water contents (Table S3) and any likely variations in the mean water content over the period of sample burial. The total dose rates also include an internal dose rate due to the decay of 40K and 87Rb inside the grains. K concentrations for individual grains have been shown to be variable due to the presence of perthitic textures; that is, Na-rich lamellae existing in a K-rich matrix (Smedley et al., 2012; Jacobs et al., 2019). We therefore estimated the K concentrations of 146 individual grains from samples CHAG12-1 and -6 using the FEI Quanta QEMSCAN 650F at the Centre for Advanced Microscopy, Australian National University. Quantitative evaluation of minerals energy- dispersive spectroscopy (QEM-EDS) measurements were made at an accelerating voltage of 15 kV

11 and beam current of 10 nA at 5 μm intervals, to capture spatial variation within individual grains. Calibration of this system makes it suitable for estimating K concentrations for perthitic grains with a standard error of ~0.5 wt%. Individual grains yielded K concentrations of 0–14 wt%, of which 88% of the grains (N = 128) gave values of 10–14 wt% (Fig. S5A). Grains with Tn intensities greater than 1000 counts/s (N = 75), the threshold value used to avoid De underestimation from inherently dim grains (see below), had a median K concentration of 12.2 wt% and arithmetic and geometric means of 11.9 and 11.8 wt%, respectively, with a standard deviation of 1.2 wt%. For all samples, we used a K concentration of 12 ± 1 wt%, together with an assumed Rb concentration of 400 ± 100 μg/g (Huntley and Hancock, 2001), resulting in effective internal dose rates of 0.80 ± 0.10, 0.49 ± 0.05 and 0.42 ± 0.05 Gy/ka for grains of 180–212, 125–150 and 90–125 μm diameter, respectively. De values for all 23 samples from layers 5 and 6 were estimated from post-infrared IRSL (pIRIR) measurements of 400–1600 individual K-feldspar grains using the procedure described by Blegen et al. (2015). Single-grain analysis allows for grains with aberrant luminescence properties to be identified and rejected before age determination, and to address any issues of incomplete bleaching before deposition or stratigraphic disturbance after deposition (Jacobs and Roberts, 2007; Duller, 2008; Roberts et al., 2015; Roberts and Jacobs, 2018). A multiple-aliquot pIRIR procedure (Li et al., 2017b) was used to obtain De values for the four samples from layer 7, and two of these samples (CHAG12-9 and CHAG14-10) were also analysed using the single-grain approach. We used the functions implemented in two R packages, Luminescence (Kreutzer et al., 2012) and numOSL (Peng et al., 2013; Peng and Li, 2017), for the data analyses, including curve fitting, De and error estimation, and graphical display and age-model analysis of the De distributions. The De estimation procedures described below have also been applied to K-feldspar grains from by Jacobs et al. (2019), where further details of these single-grain and multiple-aliquot methods can be found. All samples were measured using automated Risø TL-DA-20 instruments equipped with infrared (870 nm) light emitting diodes for stimulation of multi-grain aliquots, and focused infrared lasers (830 nm) for stimulation of individual grains loaded on to custom-made discs (Bøtter-Jensen et al., 2003). The violet/blue IRSL emissions were detected by Electron Tubes Ltd 9235QA photomultiplier tubes fitted with Schott BG-39 and Corning 7-59 filters, and beta doses were administered using calibrated 90Sr/90Y sources. For the single-grain measurements, we used a two-step, regenerative-dose pIRIR procedure in which an initial infrared bleach at 200°C is followed by infrared stimulation of the dating signal at 275°C (Blegen et al., 2015). De values were estimated using three methods: A. 2012 samples and two of the 2017 samples: projection of the sensitivity-corrected natural signal (Ln/Tn) on to the full dose-response curve regenerated for each grain. B. All samples: projection of the re-normalised Ln/Tn ratio for each grain on to the global standardised growth curve (SGC) developed for single grains of K-feldspar (Li et al., 2018) using the least-squares normalisation procedure (Li et al., 2016). Four of the CHAG12 samples (-1, -6, -9 and -10) were included in the data set used to construct the single-grain SGC of Li et al. (2018). C. All samples: projection of the weighted mean re-normalised Ln/Tn ratio for all grains used for De determination on to the SGC for K-feldspar grains. This ‘LnTn’ approach (Li et al., 2017a) does not suffer from the same saturation limitations as methods A and B.

We estimated the final De values for all single-grain samples from the Ln/Tn distributions obtained using method C (Fig. S5B), but all three methods yield statistically consistent De values (as these samples are not in, or close to, saturation). Fig. S6A compares the De values estimated for each of the 23 samples from layers 5 and 6 using methods B and C; the corresponding mean ratio is 1.000 ± 0.008 (standard error of the mean). Standard quality-assurance criteria (Jacobs et al., 2006) were applied to grains analysed using method A, including tests for dose recovery, recuperation, recycling, anomalous fading and the size

12 of residual dose remaining after bleaching. For method B, we omitted criteria associated with construction of the full dose-response curve and, for method C, criteria associated with signal saturation were also excluded. All three methods included a ‘threshold’ criterion, based on the pIRIR intensity of each grain measured after a test dose, Tn, following Roberts et al. (2018). They found that, for the 2012 samples, intrinsically dim grains yielded dose underestimates in dose recovery tests, with the same trend observed in the natural samples; some of the K-feldspar samples analysed from Denisova Cave also show this pattern (Jacobs et al., 2019). For the samples studied by Roberts et al. (2018), only grains with Tn intensities greater than 500 counts/s yielded dose recovery ratios consistent with unity. In the present study, we used a Tn threshold intensity of 1000 counts/s for all samples to avoid De underestimation from inherently dim grains. To determine appropriate De values for age determination, we examined each of the single- grain Ln/Tn distributions (Fig. S5B) for any patterns in the data, and calculated the amount of overdispersion for each distribution (i.e., the spread in values remaining after making allowance for measurement uncertainties) using the central age model (Galbraith et al., 1999; Galbraith and Roberts, 2012). The Ln/Tn distributions are overdispersed by between 28 ± 2% and 57 ± 4%, due mainly to the presence of of grains (11–46%) with values corresponding to a mean age of ~310 ka (Fig. S6B). Layer 7 was deposited at around this time (Table S3), so we interpret these grains as representing reworked older material, consistent with the micromorphological observations (see Sections S1 and S2). To separate the latter population of grains from the majority of grains in each distribution, we fitted the Ln/Tn distributions with a two- or three-component mixture using the finite mixture model (Roberts et al., 2000; Galbraith and Roberts, 2012). The overdispersion value was varied to determine the optimal model fits using maximum log likelihood and the Bayes Information Criterion (Schwartz, 1978); optimal fits were obtained using overdispersion values of 18–30% (Table S3). Two samples (CHAG12-9 and CHAG12-10) have single-grain Ln/Tn distributions optimally fitted by a single component, so we used the central age model to estimate the weighted mean De values, after rejecting statistical outliers based on the normalised median absolute deviation (Rousseeuw and Croux, 1993; Powell et al., 2002; Rousseeuw et al., 2006). For each of the distributions fitted by the finite mixture model, the weighted mean De value of the major Ln/Tn component was used to estimate the optical age. Fig. S6C shows the ages of the major and minor components in each Ln/Tn distribution (filled circles and open triangles, respectively; the latter are the same data as those plotted in Fig. S6B), compared with the six ages of the four samples from layer 7 (two single-grain and four multiple-aliquot ages; see below). As mentioned above, the age of the minor Ln/Tn component in samples from layers 5 and 6 is consistent with that estimated for layer 7. Individual K-feldspar grains of two samples from layer 7 (CHAG12-9 and CHAG14-10) were measured using method C, and all four samples from this layer were measured using the multiple- aliquot regenerative-dose (MAR) procedure of Li et al. (2017b). With the MAR procedure, six aliquots of each sample were stimulated successively at 50, 100, 150, 200 and 275°C, and the re- normalised Ln/Tn ratios were projected on to the multiple-aliquot SGC to estimate the corresponding De values. The highest stimulation temperature (275°C) is the same as that used for the single-grain measurements, thereby enabling a direct comparison of De values. CHAG12-9 and CHAG14-10 were analysed using both approaches and the weighted mean De estimates (calculated using the central age model) are statistically consistent for each sample. The 23 optical ages for layers 5 and 6 range from 63.2 ± 4.4 ka (CHAG17-6, subunit 6d) to 47.7 ± 3.0 ka (CHAG14-6, subunit 6a), but there is no relationship of age to either stratigraphic stratum or burial depth. Subunit 6d could be slightly older than the overlying sediments, but any such difference in age is smaller than the associated uncertainties of a few millennia at 1σ. The weighted mean of these ages is 54.0 ka and the arithmetic mean is 54.3 ka, both with random and total 1σ uncertainties of 0.8 and 2.5 ka, respectively. The four samples from layer 7 yield weighted mean and arithmetic mean ages of 329 and 327 ka, respectively, both with random and total 1σ uncertainties of 8 and 16 ka.

13

We note that the ages for the CHAG12 samples in Table S3 differ slightly from those reported by Slon et al. (2017a) and Roberts et al. (2018). Those studies obtained single-grain De values using method A and De values for CHAG12-9 using a single-aliquot ‘pre-dose’ procedure (Li et al., 2014). Marginally different values were also used for some of the dose rate components (e.g., internal K concentration and long-term water content). However, each pair of ages is statistically indistinguishable and the average age difference for all nine sample pairs is consistent with zero (1.1 ± 3.3%).

Chronological summary The lowest layer in the stratigraphic sequence at Chagyrskaya Cave (layer 7) does not contain any Neanderthal skeletal or cultural remains, and the fossil fauna and pollen records are also scant (Derevianko et al., 2013a; Rudaya, 2013; Rudaya et al., 2017). The optical age of ~329 ka (95.4% confidence interval: 361–297 ka) places deposition of this layer most likely within interglacial MIS 9 (337–300 ka), but the uncertainty at 2σ extends into the second half of the preceding glacial (MIS 10) and the start of the subsequent glacial (MIS 8). Layers 5 and 6 were deposited much more recently, ~54 ka (95.4% confidence interval: 59– 49 ka) based on the optical ages. This chronology is consistent with the 14C ages of >48 ka BP obtained for the bison remains, and indicate that layers 5 and 6 accumulated during the final phase of MIS 4 and/or near the start of MIS 3 (~57 ka). This matches the timing inferred from previous pollen, faunal and sedimentological analyses (Derevianko et al., 2013a; Rudaya et al., 2017), as well as the micromorphology data reported in Section S2. As subunit 6c is in primary depositional context (Section S1), we consider the optical ages to be reliable indicators of the time of deposition of the associated Micoquian-like artefacts and Neanderthal fossils. These may have accumulated over a few millennia or less, but we cannot resolve the duration of deposition more precisely because the optical ages each have a total relative systematic error of ~4.3% at 1σ, which cannot be reduced by averaging. In contrast to the optical ages, a DNA-based age estimate of ~80 ka (late MIS 5) has been proposed for the ‘Chagyrskaya Neanderthal’, Chagyrskaya 8 (Mafessoni et al., 2018; Bokelmann et al., 2019). This fossil was not found in primary context, but was recovered from the sieved sediments of subunit 6b (Table S4). The sediments, bones and artefacts in this subunit were redeposited via colluvial processes following erosion of subunit 6c (Section S1), so could Chagyrskaya 8 be associated with MIS 5 deposits in the cave that have since been removed by erosion? The dated sediment samples were collected from two separate parts of the cave (Fig. S2A), neither of which retain traces of MIS 5 deposits, but we cannot rule out the possibility that such deposits were once present and may be preserved elsewhere in the cave. Alternatively, the DNA-based age for Chagyrskaya 8 may be an overestimate if Neanderthals had a higher mutation rate than modern humans, as is the case with the great apes (Besenbacher et al., 2019), since it is based on the assumption that the mutation rate was the same in Neanderthals as in present-day humans. Other uncertainties may also affect this age estimate, such as the effective population size and generation time of Neanderthals. The latter was assumed to be 29 years, the same as in modern humans (Fenner, 2005), whereas the great apes have shorter generation intervals (Langergraber et al., 2012). Mafessoni et al. (2018) note that Chagyrskaya 8 and Denisova 3, the youngest fossil currently known from Denisova Cave, are likely to be similar in age, as they have similar proportions of ‘missing’ genetic mutations compared to present-day humans. Denisova 3 has recently been dated to 69–48 ka from optical dating of the associated sediments (Jacobs et al., 2019) and to 76.2–51.6 ka using a Bayesian modelling approach that combines chronometric (radiocarbon, uranium-series and optical ages), stratigraphic and genetic information to estimate ages for the hominin fossils at the site (Douka et al., 2019). Both of these 95.4% confidence intervals accommodate the spread of optical ages for layers 6 and 5 at Chagyrskaya Cave (63.2 ± 4.4 to 47.7 ± 3.0 ka), and suggest that Chagyrskaya 8 may have lived at around the same time at Denisova 3 or up to several millennia later.

14

Section S5. Palaeoanthropological data (B.V.)

The first Neanderthal remains from Central Asia were discovered at Teshik-Tash () in 1938 (Okladnikov, 1949). Since then, numerous discoveries, such as at Sel’ungur (Islamov et al., 1988), Obi-Rakhmat and Anghilak (Glantz et al., 2008), have shown that this region was part of the Neanderthal range. Since the 1980s, the Altai Mountains in has also yielded Neanderthal remains from several sites. The first remains were discovered at near the village of Sibiryachikha. Five teeth were described as Neanderthal by Turner (1990), although later studies (Shpakova and Derevianko, 2000; Shpakova, 2001) saw more modern affinities. The postcranial material from Okladnikov Cave is fragmentary, but shows some Neanderthal traits (Viola, 2009; Mednikova, 2011, 2013). Ancient DNA analyses of the Okladnikov 7 child humerus showed that its mitochondrial DNA (mtDNA) is similar to that of other Neanderthals (Krause et al., 2007). The second site with Neanderthal fossils in the Altai is Denisova Cave. Several fossils assigned to Neanderthals, based on their mitochondrial (Brown et al., 2016) and nuclear DNA (Prüfer et al., 2014), have been found at the site, in addition to the remains of , an Asian sister group of Neanderthals (Krause et al., 2010, Reich et al., 2010, Meyer et al., 2012; Sawyer et al., 2015; Slon et al., 2017b). Additional evidence for the presence of Neanderthals in the cave comes from analyses of sedimentary DNA, which indicate several episodes of Neanderthal occupation (Slon et al., 2017a). Neanderthals and Denisovans interacted in the area of Denisova Cave, as witnessed by the Denisova 11 individual that had Denisovan and Neanderthal parents (Slon et al., 2018). The material from Okladnikov Cave and, especially, Denisova Cave is very fragmentary (Viola, 2009), limiting its usefulness for understanding the morphology of the Altai Neanderthals. New discoveries at Chagyrskaya Cave since 2008 have changed the situation, providing a large collection (N = 74) of mostly well preserved and, in many cases, morphologically diagnostic Neanderthal remains. A detailed analysis of the human remains is still ongoing, so we will make only a few preliminary points here. Table S4 lists the material discovered up until summer 2018, along with the corresponding excavation squares and stratigraphic layers. Most of the human remains (N = 60) were found in two spatial clusters, one in squares К6, К7 and Л6 (northern cluster, N = 30; Fig. S7A) and the other in squares Н10 and Н11 (southern cluster, N = 30; Fig. S7B). The latter remains originate predominantly from subunit 6b (N = 27), while those in the northern cluster were recovered mostly from sublayers 6c/2 (N = 21) and 6c/1 (N = 5). In both clusters, elements from all anatomical regions are common, while the fossils outside these clusters are predominantly isolated teeth (Fig. S7C). The fragmentation of the material complicates estimation of the minimum number of individuals (MNI) represented, as few elements are duplicated. The most reliable estimate of the MNI is based on the dental remains, particularly the lower premolars (Fig. S8C–J and Fig. S9C). The Chagyrskaya 6 mandible, preserving the right P3 and P4, belongs to a young adult. The right lower P4 Chagyrskaya 17, which is only slightly worn, belongs to a younger individual, while the right lower P3 Chagyrskaya 41 shows stronger wear than Chagyrskaya 6 and, thus, is probably a third, older individual. Both of the two left P3s present (Chagyrskaya 12 and 50) are too worn to belong to the same individual as Chagyrskaya 17. Chagyrskaya 50 is significantly larger than Chagyrskaya 41, while Chagyrskaya 12 is much more worn than Chagyrskaya 41 and 6. Chagyrskaya 50 fits Chagyrskaya 6 in size, and the morphological and wear differences between these teeth are compatible with them belonging to the same individual. All in all, therefore, these teeth have to derive from at least four individuals: Chagyrskaya 6 (including possibly Chagyrskaya 50), 17, 41 and 12. A lower incisor (Chagyrskaya 3) and upper central incisor (Chagyrskaya 11) worn to the cervix, and a lower molar with the crown completely worn away (Chagyrskaya 64), indicate the

15 presence of another, older individual, although there remains a slight possibility that one of the more worn P3s (Chagyrskaya 12 or 41) is associated with these teeth. The four deciduous teeth are all naturally exfoliated and could belong to different children. The rather similar preservation, size and morphology of the Chagyrskaya 18 and 19 upper dm1 and dm2 could indicate that they derive from the same individual. Taking all of this into consideration, it seems prudent to assume the presence of at least five adults and one to several subadults at the site. Some of the postcranial remains are likely in association. These include the remains of a left foot from the south cluster (Fig. S7B): 3rd to 5th metatarsals (Chagyrskaya 52a,b and 53), calcaneus (Chagyrskaya 36), medial cuneiform (Chagyrskaya 27) and the distal ends of tibia and fibula (Chagyrskaya 24a,b). The C1, C2 and L5 vertebrae and several hand remains found in this area could derive from the same individual; this suggestion is tentative, however, as the teeth indicate that several different adults are represented in this cluster. The other likely associations are the remains of a right arm from the northern cluster (Fig. S7A). This includes fragments of the scapula (Chagyrskaya 48b,c), most of the humerus (Chagyrskaya 58), large portions of the radius (Chagyrskaya 39b and 47a,b) and ulna (Chagyrskaya 39a and 48a), and several hand remains (Chagyrskaya 45c,d, 60, 61 and 68). A left first metacarpal and clavicle, pedal remains and a lumbar vertebra found in close proximity could also belong to this association, but the presence of teeth from at least two adults makes this less certain. Among the postcranial remains, only one can be clearly attributed to a subadult: the atlas Chagyrskaya 2. In general, the material, though fragmented, is very well preserved. Surface preservation of the majority of the remains is exceptional. No cutmarks or impact marks are visible on any of the remains, but three teeth (Chagyrskaya 50 and 51a,b) show damage compatible with having been digested by a large carnivore. The material from Chagyrskaya Cave is morphologically relatively uniform. In cases where comparable elements from several individuals are represented (mostly dental remains), they are generally similar. Many fragments show derived Neanderthal features. The Chagyrskaya 1 deciduous canine is Neanderthal-like, with marked mesial and distal marginal ridges (Viola et al., 2011). The Chagyrskaya 6 mandible preserves the right C–M2. The canine shows marked mesial and distal marginal ridges, the P4 is asymmetrical and the molars show continuous midtrigonid crests (Fig. S9C), all features that are much more frequent in Neanderthals than in modern humans (Bailey, 2002). The morphology of the mandibular corpus, with a posteriorly placed mental foramen and an oblique mylohyoid line, is also reminescent of Neanderthals (Fig. S9A,B; Viola et al., 2012). The Chagyrskaya 13 upper central incisor shows marked mesial and distal marginal ridges, a large lingual tubercle and a strong labial curvature (Fig. S8A,B). The upper molars (Chagyrskaya 10, 59 and 63, and the M1 and M2 of Chagyrskaya 57) differ from the two known Denisovan upper molars (Denisova 4 and 8; Reich et al., 2010, Sawyer et al., 2015) in their smaller size, lack of numerous accessory cusps, and the absence of large and strongly flared roots, making their attribution to Denisovans unlikely. The postcranial remains also show traits that are similar to Neanderthals. The distal manual phalanges (Chagyrskaya 8, 56a, 56b and 61) show expanded and rounded apical tufts, a characteristic that distinguishes Neanderthals from later modern humans (Musgrave, 1971; Niewoehner, 2006). The first metacarpals (Chagyrskaya 45d and 68) are very robust, with pronounced Musculus opponens pollicis crests, and the Chagyrskaya 45c hamate has a robust and very projecting hamulus; these are features usually seen in Neanderthals (e.g., Trinkaus, 1983).

Is the genetic evidence compatible with two Neanderthal migrations to the Altai? Slon et al. (2017a) reported the recovery of Neanderthal mtDNA from sediments in sublayer 6c/1 at Chagyrskaya Cave. Interestingly, their phylogenetic trees (Slon et al., 2017a: Fig. 2 and Fig. S41) place the Chagyrskaya mtDNA sequence closer to western Eurasian Neanderthals than to the sequences recovered from fossils and sediments at Denisova Cave.

16

Further supporting evidence comes from mtDNA analyses of Neanderthal remains from Okladnikov Cave in the Altai (Krause et al., 2007) and from in the northern (Briggs et al., 2009). The lithic at Okladnikov is similar to that found at Chagyrskaya, named the Sibiryachikha techno-complex (Derevianko et al., 2013b), while the Mezmaiskaya lithic assemblage shares technological affinities with the eastern European Micoquian (Golovanova et al., 1999, 2017). Skoglund et al. (2014), Slon et al. (2017a) and Hajdinjak et al. (2018) found that the Okladnikov mtDNA sequences are closer to those of western Neanderthals than to the ‘Altai Neanderthal’ (Denisova 5), who lived ~110 ka ago (95% confidence interval: 90.9–130.0 ka; Douka et al., 2019), while Dalén et al. (2012) proposed a close phylogenetic relationship between the Okladnikov and Mezmaiskaya Neanderthals, based on their mtDNA sequences. These results should be treated with caution, however, as they are based on mtDNA data— and on incomplete mitogenomes in the case of Dalén et al. (2012) and the sedimentary DNA recovered from Chagyrskaya Cave (Slon et al., 2017a)—and mtDNA phylogenies often differ from the true phylogeny. For example, the mtDNA phylogeny places Denisovans as an outgroup to Neanderthals and modern humans (Krause et al., 2010), whereas the relationship reconstructed using the whole genome shows that Neanderthals are a sister group to Denisovans (Reich et al., 2010). Similarly, Hajdinjak et al. (2018) found differences in the position of a late Neanderthal from Les Cottés (Z4-1514) based on mitochondrial and nuclear DNA data (Hajdinjak et al., 2018: Fig. 2). Dalén et al. (2012) proposed that the mtDNA dataset indicates a population turnover within western European Neanderthals, with a local extinction followed by recolonisation from either the east or a local refugium. Interestingly, Hajdinjak et al. (2018) also proposed a possible population turnover in late Neanderthals. One of their scenarios is a population more similar to western European Neanderthals, represented by the Mezmaiskaya 2 individual, replacing the population of Mezmaiskaya 1. The other scenario is the replacement of earlier western European Neanderthals by a population similar to Mezmaiskaya 2, which is similar to the first hypothesis of Dalén et al. (2012). Ongoing nuclear DNA analyses of human remains from Chagyrskaya Cave will hopefully clarify the situation. Another hint of the possible presence of two different Neanderthal populations in the Altai region emerged from the genome of Denisova 11 (Slon et al., 2018), a young female with a Neanderthal mother and a Denisovan father who lived ~100 ka ago (95% confidence interval: 79.3– 118.1 ka; Douka et al., 2019). The Neanderthal mother has a genome closer to that of the Vindija 33.19 (a female Neanderthal who lived in northern ~48 ka cal. BP; Devièse et al., 2017) than to the high-coverage genome of Denisova 5, with an estimated population split time of ~20 ka before the time when Denisova 5 lived. This result could be explained by eastern Neanderthals spreading into western Eurasia sometime after ~100 ka and/or by western Neanderthals migrating eastward before this time and partially replacing the local population in the Altai (Slon et al., 2018). Chagyrskaya 8, whose age is constrained by a DNA-based estimate of ~80 ka (Mafessoni et al., 2018; Bokelmann et al., 2019) and the optical ages for the associated deposits (59–49 ka), has a closer genetic resemblance to the Neanderthal mother of Denisova 11 and to European Neanderthals than to Denisova 5 (Mafessoni et al., 2018). This implies an earlier separation of the latter from the ancestors of Chagyrskaya 8, Mezmaiskaya 1, Vindija 33.19 and later Neanderthals, from which Chagyrskaya 8 also differs. The archaeological evidence presented in this paper is also consistent with multiple episodes of gene flow between Neanderthals and modern humans (Wall et al., 2013; Vernot and Akey, 2014). The proposed additional pulses of introgression in East Asians (Villanea and Schraiber, 2019) could be linked to the Chagyrskaya Neanderthals, as the two introgressing Neanderthal populations (one that contributed DNA to all Eurasian populations, and the other that contributed DNA only to East Asians) were closely related (Browning et al., 2018). Further support for population replacement comes from the fact that the Denisovan father of Denisova 11 carries evidence for earlier gene flow from Neanderthals, likely dating back at least

17

300–600 generations (i.e., about 8,700–17,400 years based on a generation interval of 29 years; Fenner, 2005) before he lived (Slon et al., 2018). The high heterozygosity found in these regions of the genome indicates that this gene flow originated from a different Neanderthal population than that of Denisova 11’s mother (Slon et al., 2018). Due to the short length of these DNA segments, however, it cannot be ascertained if this Neanderthal population was the same as that to which Denisova 5 belonged.

18

Section S6. Lithic assemblage from subunits/sublayers 6a–6c/2 (V.P.C., K.A.K., A.V.S., S.V.M.)

During the earliest stages of investigation (Derevianko et al., 2013b), the Chagyrskaya assemblage was broadly perceived as technologically and typologically distinct from the other techno-complexes in the Altai, with the exception of the artefacts from Okladnikov Cave. Initially, it was defined as a -like assemblage, based on radial core reduction with a relatively high proportion of tools, including déjeté side-scrapers. Notches, denticulates, points and bifacial tools were also noted (Derevianko et al., 2013b). A total of 89,539 artefacts have been recovered from subunits 6a, 6b and 6c during the 2007– 2016 excavation seasons. At present, the technological and typological characteristics of Chagyrskaya artefacts are based on detailed studies of 4132 artefacts from subunits/sublayers 6a– 6c/2. We used Gladilin’s typology (Gladilin, 1976) – which takes into account the substantial variability among convergent/déjeté scrapers, retouched points and bifacial tools that together dominate the assemblage – and conducted a detailed attributive analysis (Chabai and Demidenko, 1998; Chabai and Uthmeier, 2017). In terms of technologically significant attributes, we used attributive analysis to reconstruct the methods used to work with the raw materials (Monigal, 2002; Chabai, 2006). Pebbles were used as raw material, with the nearby Charysh riverbed being the likely source. Petrographic analysis identified 25 types of raw material in the lithic collection, of which four were commonly used: Zasurye jasper, porphyrite, fine-grain sandstone, and hornstone. Blanks were manufactured from a wide variety of raw materials, whereas tools were produced from a limited range of raw materials. Zasurye jasper was used preferentially for manufacturing bifacial tools and convergent scrapers (Derevianko et al., 2015). The analysed assemblages from each of the subunits share multiple technological and typological characteristics. In general terms, the assemblages from subunits/sublayers 6a–6c/2 are characterised by a relatively high proportion of tools (up to 29% of the total) and a rarity of cores (0.6–0.9%). Primary flaking was focused on flake production (60–90%), with blades present in low numbers as occasional by-products (1–5%). These proportions exclude debris and chips, which account for 40–60% of the total number of pieces (Table S5). The flaking technology is based on radial (Levallois Centripetal) and orthogonal core reduction methods of flake production (Table S6 and Figs S10–S12). The assemblage contains a large number of core preparation blanks (typically different types of éclats débordant) and bifacial thinning flakes, associated with radial, orthogonal flaking and bifacial tool production (Table S7 and Figs S10, S13, S15A and S16A). The production of bifacial tools has resulted in a significant proportion of chips, including bifacial thinning chips (Table S8 and Figs S13 and S16A). Most of the flakes (N = 881) have asymmetrical trapezoidal and rectangular shapes, consistent with the morphology of the cores (Table S9). The assemblage contained significant numbers of cortical and partly cortical flakes (Table S10). The tool kit is characterised by the predominance of various scrapers (54–86%), with a preference for semi-trapezoidal, semi-crescent and semi-leaf convergent scrapers (Figs S10, S11, S14 and S17). The scrapers are accompanied by leaf-shaped and semi-trapeziodal (Mousterian) points, bifacial scrapers, bifacial points, truncations, notches, denticulates and end-scrapers (Table S11 and Figs S11, S12, S15B and S17). The largest blanks were chosen to manufacture tools inside the cave (Fig. S18). The vast majority of the bifacial tools were produced using plano-convex and plano-convex alternate methods (Figs S11, S12, S16B and S17), with Klausennischemesser and Bocksteinmesser types identified among those represented (Fig. 2). The bifacial tools were produced using numerous bone retouchers (Fig. S20). Prior to investigations at Chagyrskaya Cave, bone retouchers had not been documented for Middle Palaeolithic industries in the Altai Mountains. The technological and typological characteristics of the artefacts from subunits/sublayers 6a– 6c/2 constitute a single techno-complex. No substantial distinctions are apparent between subunits; the differences in proportions of some artefact categories mostly reflect differences in the number of

19 artefacts in each of the assemblages (e.g., 3021 and 317 artefacts for sublayers 6c/1 and 6c/2, respectively). In summary, the composition of the artefact assemblages from Chagyrskaya Cave is characterised by a relatively high percentage of tools and debitage and a low percentage of cores and bifacial tools. The large numbers of cortical flakes, a significant number of partly cortical flakes (including different varieties of débordant core-trimming elements), and the presence of bifacial thinning flakes and chips are a clear indication of on-site core reduction and tool production.

20

Section S7. Comparison with other Altai Middle Palaeolithic assemblages (K.A.K., A.I.K.)

In recent years, the Altai Mountains have become a focus of scientific discussion concerning the relationship between populations of archaic hominins and modern humans. The latest palaeogenetic and palaeoanthropological discoveries have provided evidence for the long-term coexistence of at least two hominin groups in this area – Denisovans and Neanderthals – and of gene flow between them (Prüfer et al, 2014; Kuhlwilm et al., 2016). The Middle Palaeolithic lithic assemblages in the region exhibit a large degree of variability. Technological and typological features enable to distinguish three main variants: the Denisova, Kara-Bom and Sibiryachikha techno-complexes. The Denisova and Kara-Bom techno-complexes reflect the local development of Levallois- based industries (Derevianko et al., 2013b). Linking specific hominin groups with particular techno-complexes is not straightforward, however: for example, Denisovan and Neanderthal remains have both been found in Denisova Cave and the mitochondrial DNA of both groups recovered from sediments containing indistinguishable Middle Palaeolithic industries (Reich et al., 2010; Prüfer et al., 2014; Sawyer et al., 2015; Slon et al., 2017a, 2017b). In contrast, the Sibiryachikha techno-complex has only been found in association with Neanderthals at Okladnikov and Chagyrskaya , and interpreted as evidence for a late Middle Palaeolithic migration into the Altai region (Derevianko et al., 2013b). The Denisova and Kara-Bom techno-complexes are characterised, in general, by a combination of the same flaking methods (Levallois Preferential, Levallois Convergent, radial and orthogonal cores), tool types (simple side-scrapers, notches and denticulate tools, Levallois points and rare, bi-convex bifacial tools) and similar methods of secondary treatment (Kolobova et al., 2019). These two techno-complexes have been regarded as culturally different industries, but with technological convergence resulting from several factors, such as the procurement of similar raw materials and site function (Rybin et al., 2009). The assemblages from Chagyrskaya and Okladnikova differ significantly from the Denisova and Kara-Bom techno-complexes, and have been defined as the Sibiryachikha techno-complex (Derevianko et al., 2013b; Kolobova et al., 2019). The Chagyrskaya assemblage clearly differs from the other Middle Palaeolithic industries in the Altai due to the absence of Levallois Preferential and Levallois Convergent techniques, the prevalence of radial (Levallois Centripetal) and orthogonal flaking methods and plano-convex bifacial techniques, and a high frequency of convergent side-scrapers and points. However, we recognise that some elements of the Chagyrskaya techno-complex – such as radial/orthogonal cores and low numbers of convergent scrapers and points – also appear in other Altai Middle Palaeolithic assemblages, perhaps as a result of cultural/technological adoption (diffusion), independent development (convergence) or due to palimpsests (inseparable remnants of episodes of multiple occupations of the same site by culturally different human groups). We selected the Chagyrskaya assemblage from sublayer 6c/1 to compare with other Altai Middle Palaeolithic assemblages because it is the most complete and numerous, and because the stratigraphic and micromorphological analyses indicate that this sublayer is likely an in situ deposit (see Sections S1 and S2). Archaeological data for comparison of Altai lithic assemblages was sourced from publications (Shunkov 1990; Derevianko et al., 1998a, 1998b, 2003; Derevianko and Shunkov 2002; Shalagina, 2016; Kozlikin, 2017; Krivoshapkin et al., 2018). For technological and typological comparisons, we chose stratified sites that have the largest assemblages: Denisova Cave (Entrance zone, Main and East Chambers), Kara-Bom (Middle Palaeolithic layers M2 and M1) and Ust’-Karakol-1 (layers 18 and 17–13), Strashnaya Cave (layers 10–8) and Ust’-Kanskaya Cave. Statistical analysis of these Altai Levallois-Mousterian assemblages provides technological and typological data representative of complete assemblages. These data sets and that for Chagyrskaya were examined for their techno-typological differences and similarities (Table S12). Initially, we performed a hierarchical cluster analysis to compare the technological and typological attributes of the Chagyrskaya artefacts with the Altai Levallois-Mousterian

21 assemblages, and examine the variability within and between these assemblages. Our results (Fig. S21A) demonstrate the existence of two main clusters of industries in the Altai Middle Palaeolithic: all Levallois-Mousterian assemblages assigned to the Denisova and Kara-Bom variants fall in one cluster, which is clearly separated from the Chagyrskaya assemblage in the second cluster. We did not include the artefact assemblage recovered from Okladnikov Cave in this analysis, owing to statistical incompleteness (Derevianko and Markin, 1992). Nevertheless, it has been observed that the Okladnikov collection, which is also associated with Neanderthals, shares significant typological resemblances to the Chagyrskaya assemblage (Derevianko et al., 2013b). The agglomerative coefficients support the existence of at least two main clusters, and possibly as many as five, with the Chagyrskaya assemblage clearly distinguishable from the other assemblages (Table S13). These data were subsequently imported into PAST (Hammer et al., 2001) for tests of PERMANOVA (Anderson, 2001). The resulting p-values of <0.05 for 2, 3, 4 and 5 clusters (Table S14) provide further statistical support for the existence of at least 2 clusters. We also conducted a test for the difference of means to identify those variables that have the strongest effect on cluster separation. The difference of means shows that the most influential variables are orthogonal cores, flat-faced cores, Levallois Preferential/Convergent cores, convergent/déjeté scrapers and plano-convex bifacial tools (Table S15). We performed a PERMANOVA test to assess the level of impact of these variables, and found that orthogonal cores, convergent/déjeté scrapers, simple scrapers and retouched points have a statistically significant impact (Table S16). We could not estimate the level of impact of plano-convex bifacial tools, because they occur only in the Chagyrskaya assemblage and not in the Altai Middle Palaeolithic cluster, resulting in a within-group sum of squares of zero. The absence of plano-convex bifacial tools in the latter assemblages underscores the distinction between them and the Chagyrskaya assemblage, as shown by the significant difference between the two clusters (p = 7.744E-06) indicated by the Kruskal-Wallis H test for equal medians. The results of cluster analysis can depend on the method and measure of distance used, so we also performed non-metric multidimensional scaling (nmMDS) for ordination of the Altai Middle Palaeolithic assemblages, using the same set of technological and typological variables. This non- parametric ordination method is based on computing a similarity/distance matrix and locating each item in low-dimensional space (Taguchi and Oono, 2005; Belmaker, 2017). The nmMDS plot shows a clear separation between the Chagyrskaya assemblage and the other Altai Middle Palaeolithic assemblages (Fig. S22A). Large inter-point distances signify techno-typological dissimilarities. For principal component analysis (PCA), we expanded our sample to include the following assemblages:  Two Altai Middle Palaeolithic sites without numerical age estimates: Tumechin-1 (Kara- Bom Middle Palaeolithic variant) and -2 (Denisova Middle Palaeolithic variant) (Shunkov, 1990).  Initial and early Upper Palaeolithic assemblages associated with the local development of the Altai Levallois-Mousterian: Denisova Cave (East Chamber, layer 11.1; Main Chamber, layer 11; Entrance zone, layers 7 and 6), Kara-Bom (Upper Palaeolithic levels 6–1), Ust’-Karakol-1 (layer 11) and Tumechin-4 (Derevianko et al., 1998a, 1998b; Derevianko and Shunkov, 2002).  Obirakhmatian assemblages in western Central Asia, which share many technological and typological similarities with the Kara-Bom Middle Palaeolithic variant: Obi-Rakhmat (layers 21.1–14.1) and Kulbulak (layer 23) (Kolobova et al., 2012, 2018; Krivoshapkin, 2012; Shalagina et al., 2015). We reduced the number of technological and typological variables to seven (Table S17) to generate samples of appropriate size for PCA, with a ratio of the numbers of samples to variables sufficient to yield statistically stable results (Kocovsky et al., 2009; Shaukat et al., 2016).

22

PCA was performed by projecting selected technological and typological data on to the first four principal components defined by a subset of the filtered data set. The first two principal components account for 67.9% of the variability in the data (Fig. 3B), and 90.0% is explained by the first four principal components (Table S18). The Chagyrskaya assemblage is distinct from the other Altai and Central Asian assemblages, reflecting substantial technological and typological differences. We applied the PERMANOVA test to all seven principal component scores and confirmed that the Chagyrskaya assemblage differs significantly (p-value = 0.0267) from these other lithic assemblages (Table S19). Our statistical analyses took into account mostly typological features (tool and core types), as well as several technological features (plano- and bi-convex bifacial technology), but we did not analyse the debitage for the following reasons:  The inclusion of new variables would have made the statistical results less reliable.  Only some of the Kara-Bom/Obirakhmatian assemblages have been examined using a technological approach that takes into account core preparation blanks. We did, however, find a significant difference in the attributes of debitage in the Chagyrskaya and Kara-Bom/Obirakhmatian assemblages that had been examined using a technological approach (Fig. S23 and Table S20). This outcome supports the statistical results for the other artefacts and demonstrates that there are numerous technological dissimilarities – in addition to typological differences – between the Chagyrskaya and other Altai/Central Asian Middle Palaeolithic assemblages. On the basis of three statistical analyses (hierarchical cluster analysis, non-metric multidimensional scaling, and principal component analysis), we therefore conclude that the Chagyrskaya assemblage is significantly different from the Palaeolithic assemblages recovered from other sites in the Altai and Central Asia. In particular, our results suggest that the Chagyrskaya assemblage is unique within the Altai Middle Palaeolithic, and is technologically and typologically distinct from the Levallois-Mousterian techno-complex.

23

Section S8. Comparison with European Micoquian assemblages (V.P.C., K.A.K., T.U.)

The flaking technology of the artefacts recovered from sublayer 6c/1 at Chagyrskaya Cave is based on flake radial (Levallois Centripetal) and orthogonal techniques and plano-convex/plano- convex alternate manners of bifacial tool production. The discovery of both hard stone-hammers and bone retouchers fits with the evidence observed on the blanks. The composition of the tool kit is dominated by convergent tool shapes, rather than simple shapes. There is also a preference for convergent scrapers over single- or double-edge scrapers. Trapezoidal and leaf shapes dominate the points and convergent scrapers; crescent-shaped and triangular forms also occur. Leaf-shaped points are predominant among the bifaces, but crescent-shaped, rectangular and triangular forms are also present. The Keilmesser types of bifacial scrapers were found in low numbers. All of the aforementioned typological and technological characteristics are typical of the Micoquian/Keilmessergruppen (KMG), a techno-complex based predominantly on both non- Levallois flake core reduction and a specific plano-convex method of bifacial tool production. In this study, we have paid particular attention to the presence of the Bocksteinmesser and Klausennischemesser types of bifacial tools in the Chagyrskaya assemblage, as they are diagnostic of the Micoquian (Bosinski, 1967; Richter, 1997). The earliest Micoquian sites in central and eastern are currently dated to no earlier than MIS 5d (Chabai, 2005; Richter, 2016) – that is, the last interglacial – with the latest sites in the region dated to ~30 ka (Chabai, 2013; Richter, 2016). To compare the Chagyrskaya assemblage from sublayer 6c/1 with European Micoquian assemblages, we performed the same statistical analyses as those described in Section S7. We selected the European Micoquian assemblages included in our comparison primarily on the basis that the same method of lithic analysis was used as for the Chagyrskaya assemblage (Chabai, 2005). Typological studies of the artefacts from Sesselfelsgrotte (Germany) were made by V.P.C. as part of an Alexander von Humboldt project, and all of the Crimean Micoquian collections were studied by V.P.C. Published data from techno- complexes in the Donbass-Azov region (Antonovka I and II) and the northern Caucasus (Barakaevskaya Cave) were also included in our analysis, as they were studied using a near- identical methodological approach (Gladilin, 1976; Lubin, 1994). For hierarchical cluster analysis and non-metric multidimensional scaling, we used 26 typological and technological variables (Table S21). Our data set reflects the general variability of Micoquian assemblages by including simple scrapers, déjeté scrapers and bifacial tools as key cultural and site-function markers (Jöris, 2003; Richter, 2016; Chabai and Uthmeier, 2017). We conducted a hierarchical cluster analysis to compare the technological and typological attributes of the Chagyrskaya artefacts with the European Micoquian assemblages. Hierarchical agglomerative clustering was achieved using the centroid linkage method with squared Euclidean distance; this computes the dissimilarity between the centroids of several clusters. The results of our analysis reveal that the Chagyrskaya assemblage shares many similarities with the European Micoquian assemblages. In general, the Chagyrskaya assemblage is incorporated within the Micoquian clusters (Fig. S21B). The agglomerative coefficients support the existence of four main clusters, possibly five, with the Chagyrskaya assemblage related most closely to the main cluster of numerous Micoquian assemblages. We conclude, therefore, that the Chagyrskaya and European Micoquian assemblages have a high degree of typological and technological similarity (Table S22). We tested the significance of these clusters using PERMANOVA, a non-parametric multivariate statistical test used to compare groups of objects (Anderson, 2001). The p-values calculated for 2–5 clusters are not statistically significant (all are >0.50; Table S23), which confirms the typological and technological uniformity of the Chagyrskaya and European Micoquian assemblages. We also calculated the difference of means for 2 clusters to identify those variables that have the strongest influence on cluster separation: the first cluster consists of the Chagyrskaya and European Micoquian assemblages, and the second consists of the Crimean assemblage from Kabazi II (units V and VI). The difference of means shows that the most influential variables are diagonal

24 scrapers, simple, triangular and trapezoidal points, and trapezoidal bifacial scrapers (Table S24). A PERMANOVA test indicates that core-like “Chokurcha” scrapers, leaf bifacial points and double, leaf and trapezoidal bifacial scrapers have a statistically significant impact (Table S25). The main finding from these statistical analyses is that the Chagyrskaya assemblage has a higher degree of similarity with most of the European Micoquian assemblages than do some of the Micoquian techno-complexes, such as the Crimean assemblages from Kabazi II (units V and VI), Karabai I (layer 4), Kiik-Koba (level IV) and Zaskalnaya V (unit IV). To test the validity of the cluster analysis results, we performed nmMDS scaling (Taguchi and Oono, 2005; Belmaker, 2017) for ordination of the Chagyrskaya and European Micoquian assemblages, based on the set of 26 variables in Table S21. This shows significant similarity among the Chagyrskaya and eastern European Micoquian assemblages, expressed as small inter-point distances (Fig. S22B). The Chagyrskaya assemblage clusters most closely with the Micoquian assemblages from the Crimean sites of Buran Kaya III (layer B), Starosele (level 1) and Chokurcha I, as well as with techno-complexes from the Donbass-Azov region (Antonovka I and II) and the northern Caucasus (Barakaevskaya Cave). Similarities also exist between the Chagyrskaya assemblage and the other Crimean and central European assemblages. For PCA, we merged several of the technological and typological variables to account for the significantly reduced variability among the Chagyrskaya and European Micoquian assemblages. A total of 10 variables were used for PCA (Table S26). We could not increase the number of the sites in our analysis, owing to the different methodology used to assess the other Micoquian assemblages. The first two and four principal components account for 78.1% (Fig. 3C) and 94.2% of the variability in the data, respectively (Table S27). These results provide additional statistical support for the substantial similarity of Chagyrskaya and European Micoquian techno-complexes, as do the results of the PERMANOVA test, which confirm that these assemblages are not significantly different (p-value = 0.27; Table S28). In general terms, the first principal component reflects the typological and technological uniformity of the Eurasian Micoquian, with between-assemblage variability arising mostly from differing proportions of bifacial tools. As with nmMDS scaling results, the PCA indicates that the Chagyrskaya assemblage is most similar to the Crimean Micoquian, Donbass-Azov and northern Caucasus collections. Other Micoquian assemblages contain the same morphological kits of bifacial and unifacial tools as in the Chagyrskaya assemblage, but the proportions of the various tool types are less similar. Unifacial tools with simple, trapezoidal, leaf and crescent shapes are represented in each of the analysed assemblages, as are leaf-shaped bifacial tools. These unifacial and bifacial tool shapes constitute the Micoquian ‘morphological package’. Other similarities between the Chagyrskaya and European Micoquian assemblages include the same method of primary flaking. Core-reduction strategies are represented by radial (Levallois Centripetal), orthogonal and parallel non-volumetric methods. Bifacial tool production is based on plano-convex and plano-convex-alternate methods, with bifacial scrapers and points made using the plano-convex method constituting 50–90% of the total number of bifacial tools. To check the morphological variability of bifacial tools as a probable cultural marker among European Micoquian sites and Chagyrskaya Cave, we chose assemblages from the ‘G-complex’ of Sesselfelsgrotte in Germany for geometric morphometric shape analysis. This method provides an objective and quantitative means of describing and comparing shape variability among Palaeolithic artefacts (e.g., Archer et al., 2015; Morales et al., 2015; Herzlinger et al., 2017), including Micoquian bifacial tools (Serwatka, 2014; Weiss et al., 2018). Sesselfelsgrotte is a key reference site for the Micoquian/KMG in , with one of the longest stratigraphic and cultural sequences in the region. The G-complex consists of a series of archaeological horizons containing 85,000 lithic artifacts from 13 assemblages, all of which are classified as Micoquian (Richter, 1997, 2002). Radiocarbon dating of charcoal and bone from the G-complex has yielded conventional (i.e., uncalibrated) ages of between ~48 ka BP (unit G4a/5) and ~40 ka BP (unit G2), with most of the Micoquian assemblages dating to 48–47 ka BP (Richter, 2002). A chronological position for the G-

25 complex near the end of MIS 4 and the start of MIS 3 is supported by TL ages of burnt flints from the final phase of the Micoquian/KMG at Sesselfelsgrotte and elsewhere in the region (Richter, 2000, 2002; Richter et al., 2000; Jöris, 2002). At Sesselfelsgrotte, a mean TL age (N = 4) of 56.0 ± 4.7 ka was obtained for the G-complex, which is consistent with the oldest uncalibrated 14C ages for the same layers and with a mean TL age (N = 7) of 73.2 ± 11.7 ka for the underlying Mousterian layer, unit M (Richter et al., 2000). The latter unit is separated from the overlying G-complex by archaeologically sterile deposits (units L and K) correlated with MIS 4 (Richter, 2002, 2016). Micoquian/KMG assemblages from Sesselfelsgrotte are included in our other statistical analyses (Figs 3C and 3D, Figs S21B and S22B, and Tables S21, S22, S26 and S31). To obtain a quantitative description of shape variability within and between groups of artefacts from Sesselfelsgrotte and Chagyrskaya using landmarks-based geometric morphometric shape analysis, we chose 16 and 29 bifacial tools from the Sesselfelsgrotte (units G4–G2) and Chagyrskaya (subunits/sublayers 6a–6c/2) assemblages, respectively. In our sample, we included only complete and undamaged bifacial tools, with no evidence of strong rejuvenation or knapping mistakes (such as deep negatives of unsuccessful removals with step terminations). These bifaces have been produced by means of plano-convex soft hammer bifacial flaking of chunks/pebbles, plaquettes and flake blanks. We selected bifaces with a variety of shapes, including Bocksteinmesser and Klausennischemesser types (Fig. S25). The two samples differ in terms of raw material (flint/radiolarite at Sesselfelsgrotte and jaspers/chalcedony at Chagyrskaya) and tool size. This most likely reflects distance to available sources of raw material, rather than our sampling strategy, as most raw material at Sesselfelsgrotte was transported to the site from regional sources (Richter, 1997), in contrast to the local acquisition of raw material at Chagyrskaya Cave. We first scanned each bifacial tool using structured-light 3D scanners (RangeVision PRO 5M and RangeVision Spectrum) and the raw data were processed using RangeVisionScanCenter and RangeVisionScanMerge software. The 3D images were then processed using Artifact 3D software (Grosman et al., 2008) and geometric morphometric shape analysis – including the positioning and measurement of landmarks – was performed using the Artifact GeoMorph Toolbox 3-D (AGMT3- D) software package (Herzlinger and Grosman, 2018). Bifaces were rotated on their longitudinal axis and landmarks were taken within a dense grid of 30 × 30, resulting in 1800 recorded landmarks for each biface. Principal component analysis of these data indicates a high degree of shape variability (Fig. S26), with the first two principal components accounting for 52.99% of the variability and the first four components accounting for 67.33% (Table S29). Morphological variability is greater within the Chagyrskaya sample than within the Sesselfelsgrotte sample, which lies close to the centre of the Chagyrskaya distribution. A PERMANOVA test of all PCA scores (N = 44) found no significant shape difference between the Sesselfelsgrotte and Chagyrskaya Cave bifaces (p-value = 0.0908; Table S30), which suggests a high degree of shape similarity between the two assemblages. Others factors that could explain the observed variability include differences in the raw materials used for biface production, distance to the raw material sources, and mobility patterns. Despite the geographical distance (~5000 km) between Sesselfelsgrotte and Chagyrskaya Cave, therefore, the similarities in the technology and shape of their biface assemblages strongly suggest commonalities in the conceptual design and production techniques of Micoquian/KGM bifacial tools. Links between Chagyrskaya Cave and the European Micoquian region are supported by comparisons with the genomes of Neanderthals from in Croatia and Mezmaiskaya Cave in the northern Caucasus (Prüfer et al., 2017; Mafessoni et al., 2018; Slon et al., 2018; Bokelmann et al., 2019). Chagyrskaya 8 most closely resembles Vindija 33.19 and Mezmaiskaya 1. Vindija 33.19 is from the Middle Palaeolithic (Mousterian) level G3, dated to 44,300 ± 1200 years BP, with two ages on the Vindija 33.19 bone yielding a combined estimate of 44,503 ± 1059 years BP (Devièse et al., 2017), which equates to a calibrated age range (95.4% confidence interval) of 49,940–46,130 calendar years BP. A total of 375 artefacts have been obtained from level G3. The assemblage is dominated by flake technology, with no evidence of the Levallois method. The tool kit includes side-scrapers, notches and denticulates, with “unfinished” leaf-shaped bifacial piece

26 reportedly present. The assemblage has been described as a mixture of Middle and Upper Palaeolithic elements, with the presence of bifacial technology. Unfortunately, statistical analysis of the Vindija assemblage is hindered by the low number of artefacts and evidence of post- depositional mixing of levels F, G1 and G3 (Karavanić and Smith, 1998; Devièse et al., 2017; Karavanić et al., 2018). Mezmaiskaya 1 is from the Middle Palaeolithic layer 3 at Mezmaiskaya Cave (Pinhasi et al., 2011; Prüfer et al., 2014). Middle Palaeolithic assemblages from Mezmaiskaya Cave are well documented (Golovanova et al., 1999, 2017; Golovanova and Doronichev 2003, 2017). The most numerous assemblages are from layers 3 and 2B-4, with eight associated mammal teeth dated by electron spin resonance to between 73.4 ± 5.0 and 48.5 ± 3.0 ka (Skinner et al., 2005). In general terms, the assemblages have been described as a non-Levallois flake industry, obtained mainly from recurrent flaking of single-platform cores. Two-platform, three-platform cores and bifacial multi- platform cores have also been recognised. The numerous bifacial tools have been produced using plano-convex techniques, and include Keilmesser types. The tool kits are dominated by convergent scrapers, Mousterian points and angled scrapers. Numerous simple side-scrapers have been identified, alomg with transverse, diagonal scrapers and denticulate-notched tools. The Mezmaiskaya assemblage has been regarded as a northwestern Caucasus expression of the eastern European Micoquian, together with assemblages from Ilskaya-1 and -2, Khadjokh-2, Monasheskaya, Barakaevskaya, Autlev and Matuzka Caves, and Gubs Rockshelter-1 (Golovanova and Doronichev, 2003, 2017; Golovanova et al., 2017). The Barakaevskaya Cave assemblage shares techno-typological similarities with the Chagyrskaya assemblage (Fig. 3C and Fig. S22B), as do the Mezmaiskaya assemblages from layers 3 and 2B-4 (e.g., flake-based technology with numerous plano-convex bifacial tools, convergent, angled side-scrapers and Mousterian/retouched points). For statistical comparison with the Chagyrskaya assemblage, we used the most complete data set published for the Mezmaiskaya assemblage (Golovanova et al., 1999). The typological definitions used for the latter tools mostly fit with those used for the Altai/Central Asian Middle Palaeolithic assemblages, an exception being the cores. We combined the Altai/Central Asian Middle and Upper Palaeolithic data with the European Micoquian data to compare with the Chagyrskaya and Mezmaiskaya assemblages. For PCA, we used the set of 5 common typological and technological variables listed in Table S31 (i.e., Levallois tools, simple scrapers, convergent/déjeté scrapers, retouched points and plano-convex bifacial tools). These variables were selected because they could be applied to both methodological approaches and to all assemblages, and because they are also the most informative in terms of differentiating between techno-complexes. The first two principal components account for 76.8% of the variability in the data (Fig. 3D and Table S32). All sites fall into one of two clusters, which have 95% confidence interval ellipses that are clearly separated: the Altai/Central Asian Middle and Upper Palaeolithic cluster, and the European Micoquian, Mezmaiskaya and Chagyrskaya cluster. Application of the PERMANOVA test to all 5 principal component scores confirms that the European Micoquian and Chagyrskaya cluster differs significantly (p-value = 0.0001) from the Altai/Central Asian Middle and Upper Palaeolithic assemblages (Table S33). Although this pooled analysis is based on a smaller number of variables than the original analysis, the same results are obtained, thereby supporting our previous conclusions. By comparing several techno-complexes that have most likely been produced by different populations, our analysis adds to the long-standing debate (Bordes, 1966; Binford and Binford, 1966), which is still ongoing, about the correspondence of stone tool typology to demic diffusion, trans-cultural diffusion and demography. Multiple arguments for and against such links have been proposed, drawing on data on assemblage composition, typological and technological characteristics of artefacts, environmental reconstructions and circumstantial DNA (e.g., Richter, 1997; Jöris, 2002; Soressi, 2005). Recent studies have revealed the crucial importance of cultural traditions associated with different groups of hominins (e.g., Petraglia et al., 2007; Jaubert, 2011; Slimak et al., 2011; Delagnes and Rendu, 2014; Thiébaut et al., 2014). Chagyrskaya Cave provides multiple lines of evidence (chronology, typology and genetics) in favour of the conformity of stone

27 tool typology to certain homimin groups – in this case the most recent incursion of Neanderthals into the Altai. In summary, the Chagyrskaya assemblage and European Micoquian techno-complexes (including Mezmaiskaya) share strong technological and morphological similarities, with chronological overlap between about 60 and 50 ka. The Chagyrskaya assemblage falls within the range of variation observed among the European Micoquian and is, in essence, an Altai variant of the Micoquian. Accordingly, we propose that the geographical boundary of the European Micoquian should be expanded eastward to incorporate Chagyrskaya Cave (Fig. S24). In this context, the previous easternmost Micoquian complexes at Sukhaya Mechetka (Volga River region; Kuznetsova, 1985), Khotylevo I and Betovo (Desna River region; Ocherednoi et al., 2014) and, possibly, Garchi 1 (lower layer; Pavlov et al., 2004) can now be viewed as evidence for the eastward dispersal of Neanderthals carrying Micoquian tools, with Chagyrskaya Cave providing further indications of this migration. Also, as Chagyrskaya and Okladnikov Caves comprise the Sibiryachikha Middle Palaeolithic variant in the Altai, we now consider it more appropriate to frame the Sibiryachikha variant in terms of Micoquian variability across Eurasia. Our results are supported by new DNA analyses linking the Chagyrskaya 8 with Neanderthals at Mezmaiskaya and Vindija, providing a rare example of consistency between archaeological and genomic data in a Palaeolithic context.

28

Section S9. Timing and routes of Neanderthal migration (R.G.R., N.A.R., V.P.C., B.V., K.A.K.)

In central and , the earliest manifestations of the Micoquian/KMG techno- complex are known from MIS 5 sites such as Königsaue (units A and C), Lichtenberg, Neumark- Nord, Külna (layers 7c and 9), Balver Höhle I and II, Wylotne (layers 5–8), Kabazi-II (units V and VI), Zaskalnaya V, Il’skaya-1 and Garchi 1 (Mania and Toepfer, 1973; Praslov, 1984; Valoch, 1988; Veil et al., 1994; Jöris, 2001; Chabai, 2005; Gerasimenko, 2005; Madeyska, 2006; Brühl and Laurat, 2010; Gnibidenko, 2017; contra Richter, 2016). Further development of the Micoquian/KMG took place in this region during MIS 4 and 3 (Chabai et al., 2004; Jöris, 2006; Slimak et al., 2011). The chronology and routes of Micoquian/KMG Neanderthal migration into northern Asia can be framed in terms of two hypotheses. The first hypothesis is that this group of Neanderthals initially arrived in the Altai sometime during MIS 5. This appears to be the case in central and eastern Europe, where the mild/moderate interglacial conditions of MIS 5 provided an opportunity for Neanderthals to colonise the western foothills of the central Urals, as indicated by occupation of Garchi 1 (Slimak et al., 2011; Gnibidenko, 2017; Pavlov, 2017). The second hypothesis is that this group of Neanderthals appeared in the Altai towards the end of MIS 4, and that Chagyrskaya Cave was occupied soon after their initial entry into northern Asia. The DNA-based age estimate of 87– 71 ka for Chagyrskaya 8 (Mafessoni et al., 2018; Bokelmann et al., 2019) is compatible with the first hypothesis, whereas the optical ages for the Micoquian/KMG layers at Chagyrskaya Cave (59– 49 ka) are consistent with the second hypothesis. While there is evidence for hominin occupation of the Altai Mountains during MIS 5, and for the presence of populations linked genetically to Neanderthals (Prüfer et al., 2014; Slon et al., 2017a, 2018; Jacobs et al., 2019; Douka et al., 2019), no Micoquian-like assemblages are known from this time period. Accordingly, archaeological evidence in support of the first hypothesis has yet to be found in the Altai, but it cannot be ruled out as a possibility. The second hypothesis assumes that Neanderthal populations with Micoquian/KMG artefacts colonised the Altai during the cold and harsh climate of MIS 4, when steppe or tundra-steppe conditions prevailed in the region. The ecological uniformity from west to east along the Eurasian steppe belt could have facilitated the eastward migration of Neanderthals into the Altai, perhaps assisted by the opening of additional migratory routes across the exposed shelf of the Caspian Sea in MIS 4, during the Atel regression (Svitoch, 2012; Yanina, 2014; Yanina et al., 2018). In contrast, the level of the Caspian Sea rose significantly during transgressive periods, simultaneous with the formation of the Manych spillway between the Caspian and Black Seas. At such times, the Manych spillway may have hindered the dispersal of Neanderthals into the northern part of the Greater Caucasus mountain range, where several Micoquian sites are located (Fig. S24). Late Pleistocene transgressive/regressive events in the Ponto-Caspian region were strongly controlled by climatic dynamics in the Northern Hemisphere. The Late Khazar and the Khvalynian transgressive periods were separated by the deep Atel regression, which started ~70 ka, coincident with or soon after the onset of MIS 4 (Yanina, 2014; Yanina et al., 2018). At its greatest transgressive extent, the surface area of the Caspian Sea increased by 250%, and water levels rose by 50 m, compared to the present day; in contrast, the level of the Caspian Sea fell by 140 m at the peak of the regressions. Maximum sea-level during transgressive periods was controlled by the height of the Manych threshold: this threshold was breached during the Late Khazar transgression (in MIS 5) and twice during the Early Khvalynian transgression (in MIS 3). The Atel regression associated with MIS 4 was characterised by very low water levels in the Caspian Sea, an absence of flows over the Manych spillway, and a very cold climate with tundra-steppe vegetation (Grichuk, 1954; Yanina, 2014; Yanina et al., 2018). Environmental reconstructions for the period of Neanderthal occupation of Chagyrskaya Cave (Rudaya et al., 2017) and of Sukhaya Mechetka (Dolukhanov et al., 2009) suggest that they could survive in cold, dry and treeless environments, hunting horses and bison on the cold steppe or tundra-steppe landscapes. The occurrence of herbivorous ungulates after MIS 5 on the cold steppe

29 across Eurasia – the so-called ‘tundra-steppe’ or ‘Mammoth steppe’ – and the substantial fall in the level of the Caspian Sea after ~70 ka (during the Atel regression) may have provided especially favourable conditions for the eastward expansion of Neanderthals into northern Asia.

30

Fig. S1. (A) Location map of Chagyrskaya Cave, Okladnikov Cave, Denisova Cave, Ust’-Karakol- 1 and Kara-Bom in the Altai Mountains. (B) View upstream (i.e., to the southeast) along the Charysh River valley, with the location of Chagyrskaya Cave indicated by the arrow.

31

Fig. S2. (A) Plan map of Chagyrskaya Cave showing excavation squares, locations of optical dating samples collected in 2012, 2014 and 2017 (CHAG12-1 to -10, CHAG14-2 to -12 and CHAG17-3 to -9, shown in orange) and locations of micromorphology samples collected in 2014 (MM2–4, shown in green) and 2017 (2969, 2970, 2984, 2985, 2987–2989, shown in blue). The area shaded in yellow was excavated in 2016 and 2017. (B) Stratigraphic profile along the purple line in panel A. (C) Stratigraphic profile along the red line in panel A. Both vertical scales are in cm below reference datum, the horizontal scale is in bottom right-hand corner, and the layer, subunit and sublayer numbers are circled.

32

Fig. S3. Photo montage of micromorphology samples collected by M.T.K. in 2017 (panels A–D) and by M.W.M. in 2014 (panels E–G). Sample codes and (sub)layer numbers are shown in all panels and the associated sampled profiles are indicated in Fig. S2A. The arrows in panels A–C connect the sampled profiles with a photo of the area shaded in yellow in Fig. S2A.

33

Fig. S4. Photomicrographs of key micromorphological features. (A) MM4B, layer 7. Poorly sorted coarse components, with very frequent clay aggregates, ‘rip up clasts’ reworked from elsewhere in the cave by the action of water (ppl). (B) 2984, layer 7. Angular clay aggregates suggest limited residence time in the karstic system, compared with rounded and sub-rounded clay aggregates from the same horizon, indicating multiple sources for these coarse components (ppl). (C) 2988, subunit 6d. Detail of porous nature of the sediments from this subunit, including a laminated clay crust that has been reworked into these sediments from elsewhere (ppl). (D) 2988, subunit 6d. General view of the composition of this subunit. Note coarse inclusions often coated in fine grained clays and silts (ppl). (E) 2985, sublayer 6c/2. Bone fragments, some of which have fractured in situ, in close association with sub-rounded coprolites. The general arrangements of the coarse inclusions is chaotic, with no preferential alignment of long-axes, indicative of cryoturbation of the sediments (ppl). (F) 2985, sublayer 6c/2. Thin fragment of chert with a quartz vein inclusion, the bi-product of stone tool manufacture (ppl). (G) MM4B, sublayer 6c/1. A rare coprolite fragment and well preserved small bone fragments (ppl). (H) MM4A, sublayer 6c/1. Granular microstructure of the fine sediment matrix, indicating frost action, with small bone fragments in a good state of preservation, inconsistent with having passed through the gut of a carnivore (ppl). (I) MM4A, sublayer 6c/1 in xpl to show alignment of clay minerals around mineral grains, stress features most likely related to frost heave (xpl). (J) MM3B, subunit 6b. Very fine sediment matrix with granular microstructure, indicative of cold climate processes, most likely frost heave. Especially towards the upper part of the image, fine laminated clay clasts are common, and these are likely clay crusts and cappings that are being reworked by expansion and contraction, again consistent with weak cryoturbation processes (ppl). (K) MM3B, subunit 6b. Elongate bone, crushed in situ, most likely as a result of trampling (ppl). (L) MM3B, subunit 6b. Coprolites are relatively rare in 6b (and 6c/1), but this large fragment has cracked in situ, most likely by frost action, and various quartz silt grains and small organic inclusions can be observed (ppl). (M) MM3B, subunit 6b. Fine silt and clay fabric is separating into rounded ‘peds’, with clay coatings and cappings evident on many of the mineral grains and surrounding aggregate grains, indicating disturbance, most likely by frost heave (ppl). (N) MM3A, subunit 6a. Detail of coprolite with possible hair/fur inclusion. This part of subunit 6a is much looser, possibly due to disturbance by bioturbation (ppl). (O) MM3A, subunit 6a. Detail of coprolites, showing fine grained nature of the material, with very low quantities of inclusions, consistent with hyena droppings. Note also the dense, non-porous silty clay matrix and clay cappings on some mineral grains (ppl). (P) MM2B, subunit 6a. Dense concentrations of coprolite fragments and sand-size mineral grains, primarily quartz with other metamorphic and igneous lithologies (ppl). (Q) MM2B, subunit 6a in xpl, note the coprolites are isotropic. (R) 2969, layer 5. Sub-rounded coprolite fragment with thin clay coating suggesting erosion and transport from an earlier deposit.

34

A

B CHAG12-1 CHAG12-2 CHAG12-3 CHAG12-4 = 178 = 149 = 82 = 111 = 22 = 51 = 21 = 49

CHAG12-5 CHAG12-6 CHAG12-7 CHAG12-9 = 118 = 129 = 64 = 120 = 21 = 21 = 11 = 19

CHAG12-10 CHAG14-2 CHAG14-3 CHAG14-4 = 172 = 118 = 112 = 114 = 34 = 49 = 41 = 41

CHAG14-5 CHAG14-6 CHAG14-7 CHAG14-8 = 143 = 98 = 126 = 91 = 49 = 57 = 73 = 55

CHAG14-9 CHAG14-10 CHAG14-12 CHAG17-3 = 141 = 59 = 172 = 70 = 51 = 11 = 41 = 60

35

CHAG12-1 CHAG12-2 CHAG12-3 CHAG12-4 = 178 = 149 = 82 = 111 = 22 = 51 = 21 = 49

CHAG12-5 CHAG12-6 CHAG12-7 CHAG12-9 = 118 = 129 = 64 = 120 = 21 = 21 = 11 = 19

CHAG12-10 CHAG14-2 CHAG14-3 CHAG14-4 = 172 = 118 = 112 = 114 = 34 = 49 = 41 = 41

CHAG14-5 CHAG14-6 CHAG14-7 CHAG14-8 = 143 = 98 = 126 = 91 = 49 = 57 = 73 = 55

CHAG14-9 CHAG14-10 CHAG14-12 CHAG17-3 = 141 = 59 = 172 = 70 = 51 = 11 = 41 = 60

CHAG17-5 CHAG17-6 CHAG17-7 CHAG17-8 = 130 = 80 = 86 = 144 = 10 = 27 = 16 = 24

CHAG17-9 = 74 = 28

Fig. S5. (A) K concentrations (weight %) estimated from quantitative evaluation of minerals energy-dispersive spectroscopy (QEM-EDS) measurements of 146 individual K-feldspar grains from samples CHAG12-1 (filled circles) and CHAG12-6 (open circles), plotted as a function of Tn intensity. The dashed line shows the Tn threshold intensity used for De estimation. (B) Distributions of re-normalised single-grain Ln/Tn ratios for six samples from layer 5, 17 samples from layer 6, and two samples from layer 7. Sample codes are given in each panel. The radial plots are arranged by year (starting with samples collected in 2012) and then in numerical order within each year. For 23 of these samples, the finite mixture model was used to fit the two or three Ln/Tn components in each of the single-grain distributions and to estimate the weighted mean value of each component. The major and minor components consist of re-normalised Ln/Tn ratios shown as filled circles and open triangles, respectively, and the solid lines are centred on the weighted mean value of each component. Two samples (CHAG12-9 and CHAG12-10) have Ln/Tn distributions consistent with a single component; the shaded band is centred on the weighted mean value estimated using the central age model, after rejecting grains identified as statistical outliers (open triangles) using the normalised median absolute deviation. The age listed for each of the single-grain samples in Table S3 is based on the weighted mean value for the major component (i.e., the Ln/Tn component comprised of the majority of grains, shown as filled circles) or, for CHAG12-9 and CHAG12-10, the weighted mean value of the re-normalised Ln/Tn ratios following outlier rejection (i.e., the ratios shown as filled circles).

36

A B C 250

200

150

value (Gy) value e

100 LnTn D LnTn

50

0 0 50 100 150 200 250 SGC D value (Gy) e

Fig. S6. (A) Comparison of weighted mean De values for the major Ln/Tn component of the 23 samples from layers 5 and 6, estimated using two single-grain methods: method B (standardised growth curve, SGC) and method C (LnTn). Uncertainties are shown at 1σ and the dashed line indicates the 1:1 ratio. (B) Weighted mean ages for the same samples, but calculated from the minor Ln/Tn component comprised of larger re-normalised ratios; the shaded band is centred on the pooled mean age of ~310 ka. (C) Same data as in panel B plus the six age estimates obtained for the four samples from layer 7 (open triangles), compared with the weighted mean ages of the 23 samples from layers 5 and 6 (filled circles), determined from the major Ln/Tn component (panel A); the shaded bands are centred on the respective pooled mean ages.

37

A B C

Fig. S7. Overview of the human remains from Chagyrskaya Cave. The teeth and postcranial remains are not to scale. (A) Remains from the northern cluster, squares К6, К7 and Л6. (B) Remains from the southern cluster, squares Н10 and Н11. (C) Remains from outside the two clusters.

38

A C E G I

B D F H J

Fig. S8. Selected isolated teeth from Chagyrskaya Cave. (A, B) Chagyrskaya 13 I1 in occlusal and labial view. Note the pronounced shoveling and large basal tubercle. (C, D) Chagyrskaya 12 P3 in occlusal and mesial view. (E, F) Chagyrskaya 14 P4 in occlusal and distal view. (G, H) Chagyrskaya 41 P3 in occlusal and distal view. (I, J) Chagyrskaya 50 P3 in occlusal and mesial view.

39

A B

C

Fig. S9. Chagyrskaya 6 mandible fragment preserving right C–M2. (A) Buccal view; note the relatively posterior position of the mental foramen. (B) Lingual view and (C) occlusal view.

40

Fig. S10. Core, core preparation blanks and tools from Chagyrskaya Cave (subunit 6a): straight ventral with natural back (1), overpassed bifacial thinning flake (2), bifacial thinning flake (3), semi-crescent dorsal scraper (4), sub-trapezoidal alternate scraper (5), semi-leaf dorsal, thinned base point (6), bifacial scraper straight, thinned base, naturally back (7), unidentifiable bifacial fragment (8), radial core (9), semi-leaf dorsal, thinned back scraper (10).

41

Fig. S11. Core and tools from Chagyrskaya Cave (subunit 6b): semi-leaf alternate point (1), unidentifiable convergent bifacial scraper (2), semi-crescent dorsal thinned base scraper (3), sub- trapezoidal alternate scraper (4), semi-trapezoidal point (5), semi-triangular dorsal thinned base point (6), semi-triangular dorsal point (7), sub-trapezoidal alternate scraper (8, 11), sub-trapezoidal dorsal thinned base scraper (9), orthogonal core (10).

42

Fig. S12. Cores from Chagyrskaya Cave (sublayer 6c/1): radial core (1), orthogonal cores (2, 3).

43

Fig. S13. Core preparation blanks from Chagyrskaya Cave (sublayer 6c/1): crested débordant flake (1), débordant flake from radial core (2), bifacial thinning flakes (3, 4), cortical débordant flake (5), technical flake (6), lateral débordant flake (7), débordant flake from radial core/pseudo- Levallois point (8).

44

Fig. S14. Side-scrapers from Chagyrskaya Cave (sublayer 6c/1): semi-trapezoidal dorsal scrapers (1–4), semi-trapezoidal alternate scrapers (5, 6), semi-leaf dorsal, thinned base scraper (7), semi- leaf dorsal (8, 9).

45

Fig. S15. Core preparation blanks and tools from Chagyrskaya Cave (sublayer 6c/1). (A) Core preparation blanks: technical flakes (1, 6), lateral débordant flakes (2, 4), débordant flakes from radial cores (3, 7), cortical débordant flake (5). (B) Tools: semi-leaf asymmetrical dorsal point (1, 3), semi-leaf asymmetrical alternate point (2), semi-leaf dorsal point (4), sub-leaf dorsal point (5), leaf-shape dorsal (6), denticulates (7, 8), truncation (9).

46

Fig. S16. Bifacial thinning flakes and bifacial tools from Chagyrskaya Cave (sublayer 6c/1). (A) Bifacial thinning flakes (1–8). (B) Bifacial tools: semi-leaf backed bifacial point with thinned base (1), sub-triangular backed bifacial point with thinned base (2), sub-leaf naturally backed bifacial point with thinned base (3), sub-leaf bifacial scraper (4), straight-convex bifacial scraper, distally thinned (5), sub-leaf bifacial scraper (6).

47

Fig. S17. Tools from Chagyrskaya Cave (sublayer 6c/2): semi-crescent alternate scraper (1, 3, 7), straight bifacial scraper (2), semi-trapezoidal dorsal scraper (4), semi-trapezoidal alternate scraper (5), semi-leaf alternate (6), retouched flake (8).

48

Fig. S18. Comparison of the dimensions (length and width) of the complete blanks from Chagyrskaya Cave (sublayer 6c/1), showing that the largest blanks were used preferentially for tool manufacture.

49

Fig. S19. Scar-pattern analysis of a Klausennischemesser type of bifacial tool from Chagyrskaya Cave, sublayer 6c/1. Initially, the plano (A–F)-convex (L–S) surfaces of the pre-form were shaped. The plano (H) and convex (T) surfaces of the cutting edge were then treated and, finally, the cutting edge was retouched (U). These results suggest that this tool has not been broken.

Fig. S20. Three bone retouchers (A–C) from Chagyrskaya Cave, sublayer 6c/1.

50

Fig. S21. Hierarchical cluster analysis of Altai Middle Palaeolithic and European Micoquian assemblages. (A) Dendrogram showing the clear separation of the Chagyrskaya Cave assemblage from the other Altai Middle Palaeolithic assemblages. (B) Dendrogram showing that the Chagyrskaya Cave assemblage lies within the range of variability of the European Micoquian. Both dendrograms were constructed using the centroid linkage method and squared Euclidean distance. Assemblages that are most similar have the smallest linkage distances.

51

Fig. S22. Non-metric multidimensional (3D) scaling of Altai Middle Palaeolithic and European Micoquian assemblages. (A) Plot of Altai Middle Palaeolithic assemblages by Euclidean distance matrix from frequency distributions (stress value = 0.17): 1, Chagyrskaya Cave (sublayer 6c/1); 2, Ust’- Karakol-1 (layers 17–13); 3, Ust’-Karakol-1 (layer 18); 4, Kara-Bom (layer MP2); 5, Kara-Bom (layer MP1); 6–8, Denisova Cave (Entrance zone, layers 10–8, respectively); 9–13, Denisova Cave (Main Chamber, layers 22, 21, 19, 14 and 12, respectively); 14–19, Denisova Cave (East Chamber, layers 15, 14, 12 and 11.4–11.2, respectively); 20, Strashnaya Cave; 21, Ust’-Kanskaya Cave. Central Asian Middle Palaeolithic and Altai Upper Palaeolithic (UP) assemblages are not plotted here, but correspond to the following italicised numbers in Fig. 3B: 22, Ust’-Karakol-1 UP (layer 11); 23, Kara-Bom UP (layers 6 and 5); 24, Kara-Bom UP (layers 4–1); 25, 26, Denisova Cave UP (Entrance zone, layers 7 and 6, respectively); 27, Denisova Cave UP (Main Chamber, layer 11); 28, Denisova Cave UP (East Chamber, layer 11.1); 29, Tumechin-1; 30, Tumechin-2; 31, Tumechin-4; 32–39, Obi-Rakhmat (layers 21.1, 20, 19.5–19.1 and 14.1, respectively); 40, Kulbulak, layer 23). (B) Plot of Chagyrskaya Cave and European Micoquian assemblages by Euclidean distance matrix from frequency distributions (stress value = 0.1372): 1, Chagyrskaya Cave (layer 6c/1); 2–6, Kabazi V (subunits I/4A–II/7, III/1, III/1А, III/2 and III/5, respectively); 7, Karabai I (layer 4); 8 and 9, Kabazi II (units IIA–III and V–VI, respectively); 10, Kiik-Koba (level IV); 11, Buran Kaya III (layer B); 12, Starosele (level 1); 13, Chokurcha I (unit IV); 14–20, Zaskalnaya V (units I, II, IIа, III/1–III/9-1, III/10–III/14, IIIA and IV, respectively); 21–23, Sesselfelsgrotte (units G4–G2, respectively); 24, Antonovka I; 25, Antonovka II; 26, Barakaevskaya Cave.

52

Fig. S23. Non-metric multidimensional (3D) scaling of Altai and Central Asian Middle Palaeolithic debitage assemblages. Plot of assemblages by Euclidean distance matrix from frequency distributions (stress value = 0.066): 1, Chagyrskaya Cave (sublayer 6c/1); 2, Kara-Bom (layer MP2); 3, Kara- Bom (layer MP1); 4–11, Obi-Rakhmat (layers 21.1, 20, 19.5–19.1 and 14.1, respectively); 12, Kulbulak, layer 23.

53

Fig. S24. Map of Eurasia showing the locations of the main Micoquian/Keilmessergruppen (KMG) sites in western, central and eastern Europe and northern Asia: 1, Grotte de Verpilliere I and II; 2, La Baume de Gigny; 3, Bockstein; 4, Hohler Stein Schambach; 5, Schulerioch; 6, Balve; 7, Sesselfelsgrotte; 8, Klausennische; 9, Buhlen; 10, Zalzgitter-Lebenstedt; 11, Lichfenberg; 12, Königsaue; 13, Kulna; 14, Okiennik; 15, Zwolen; 16, Wylotne; 17, Ciemna; 18, Korolevo; 19, Yezupil; 20, Ripiceni Izvor; 21, Starosele; 22, Kabazi II; 23, Kabazi V; 24, Chokurcha I; 25, Kiik-Koba; 26, Buran Kaya III; 27, Karabai I; 28, Sary Kaya; 29, Zaskalnaya V; 30, Zaskalnaya VI; 31, Prolom I; 32, Prolom II; 33, Ilskaya-1 and -2; 34, Mezmaiskaya Cave; 35, Barakaevskaya Cave; 36, Monasheskaya Cave; 37, Sukhaya Mechetka; 38, Antonovka I; 39, Antonovka II; 40, Garchi 1; 41, Chagyrskaya Cave.

54

Fig. S25. Bifacial plano-convex tools with differing morphologies from Chagyrskaya Cave and Sesselfelsgrotte (Germany): bifacial scrapers, semi-crescent, Klausennischemesser type (1, 2), bifacial scraper, sub-leaf, Klausennischemesser type (4), bifacial scrapers, sub-leaf (3, 5, 9–11), bifacial points, sub-triangular (6–8), bifacial scrapers, sub-crescent (12–15).

55

Fig. S26. Scatterplot of scores on the first two principal components obtained from geometric morphometric shape analysis of bifacial tools from Chagyrskaya Cave (subunits/sublayers 6a–6c/2) and Sesselfelsgrotte (G-complex). Centroids are indicated by crosses.

56

Table S1. Micromorphological characteristics of layers 5–7 as observed in thin section, and inferred sedimentation processes and depositional environments.

Thin Layer Sedimentation processes Micromorphological characteristics Depositional environment section number (climatic signal?)

2969 5 Loose silty clay with fine to medium rock fragments of Aeolian and colluvial (humid, Significant quantities of fragmented and rounded variable lithology (volcanic and fine-grained clastic rocks, and cold) coprolites, some of which are clay coated, suggest that isolated mineral grains, mostly quartz and hornblende). layer 5 contains reworked material from older Biological (animal coprolites) Aggregate microstructure is well developed in localised areas. subunits/sublayers (e.g., 6a, 6b, 6c/1 or 6c/2). The Small angular bone fragments are present and coprolite aggregate microstructure indicates the impact of frost fragments are very frequent, usually rounded, and in some action, with reorganisation of the matrix related to ice areas they constitute the dominant composition of the lensing and early stages of grain and aggregate rotation. sediments. Coprolites are usually yellow with dark brown or black spots (plane polarised light; ppl) and are isotropic in cross polarised light (xpl). Some coprolites contain elongated fur inclusions and others are coated with brown clay.

2970 5 Same as thin section 2969

MM2A 5 Moderately compact silty clay with fine quartz sand. Poorly Aeolian, colluvial and Microstratigraphic features are indicative of deposition in sorted, with some large (~30 mm) rock fragments, mainly of infiltration a cave floor environment, with evidence of bioturbation, volcanic/metamorphic origin (e.g., schist, marble, andesite). possibly carnivores such as cave hyena (Crocuta crocuta (humid?) Speckle-stippled b-fabric. Very frequent coprolite fragments spelaea) or wolf (Canis lupus) (Vasiliev, 2013). Large in the lowest third of the thin section. Coprolites generally quantities of small coprolite fragments, most likely hyena have a pale yellow colour and a darker rind, with a speckled (Horwitz and Goldberg, 1989), are present throughout. (stippled) appearance (ppl), and are isotropic (xpl). Coprolite Poorly sorted composition may be consistent with fragments gradually become smaller in size towards upper reworking, possibly through bioturbation. Natural region of thin section. In this upper region, pore spaces sedimentation with presence of non-hominin animals. increase in size and frequency. Occasional phosphatised bone Minor signs of diagenesis, possible presence of and dissolving limestone fragments. groundwater in the sediment.

57

MM2B 6a Densely compact and non-porous clay silt, containing coarse Aeolian, colluvial and Cave floor environment, with minor signs of reworking inclusions including occasional gravel clasts up to ~30 mm in infiltration and clay translocation. High frequencies of coprolites size. Angular to sub-angular quartz sand grains and clay suggest carnivore activity in the cave, at times particularly aggregate ‘rip up clasts’ are present in moderate numbers. (cold, dry) intensive. Mechanically cracked aggregate grains and Mineral grains are primarily of volcanic and metamorphic coprolites are consistent with dry conditions. Again, origin (e.g., granitic, schistose), with occasional sedimentary natural sedimentation processes with the presence of rocks, including chert and weathered sandstone. Very non-hominin animals, presumably hyenas based on frequent coprolite fragments, rounded to sub-angular, many coprolite morphology. of which are disintegrating in situ. Some mineral grains and rock fragments have clay coatings and cappings and phosphatised speleothem fragments are present in very low numbers. Coprolite fragments frequent in localised concentrations. Incipient separation of fine matrix into rounded ‘peds’, or granular microstructure.

MM3A 6a Subunit 6a as recorded in this thin section is very similar to Same as MM2B Very similar to the upper part of subunit 6a recorded in MM2B. Coarse fraction includes very frequent rounded MM2B, but with a much higher frequency of limpid clay coprolites, ranging from 200–500 µm in size. Fine fraction b- aggregate grains and crust fragments, which may suggest fabric is generally speckled (stippled), but with occasional reworking and fracturing of fine waterlain clays, possibly granostriated b-fabric present around rock fragments and formed during freeze–thaw processes. Granostriated b- aggregate grains. Clay aggregates often comprise small (silt- fabric and granular microstructure is consistent with frost- to fine-sand size) sub-angular to sub-rounded aggregates of heave and incipient cryoturbation processes (Van Vliet- limpid clay with no internal structure, whilst larger Vanöe, 2010). Coprolitic material indicates intensive use aggregates of darker clays with fine silt are also present in of the cave by carnivores. Generally dry, but minor inputs lower numbers. Larger clay aggregates are mechanically of water might be linked to nascent freeze–thaw action. fractured.

6b Subunit 6b is densely compact and non-porous sediment, Aeolian, colluvial and possible Evidence of cracking and fracturing of bones and clay comprising occasional to moderate quantities of quartz sand infiltration of fine clays aggregates in situ might indicate trampling of the and occasional clay aggregates in a fine silt and clay matrix. sediments by large animals, possibly hominins. Gravel clasts are also present in low numbers, ranging in size (cold, dry) Mechanical disintegration of clay aggregates most likely a up to ~30 mm. Rock fragments include both limestone and function of freeze–thaw action. The frequency of occasional pieces of sandstone. Where clay aggregates are coprolites declines dramatically in comparison with present these are smaller and in smaller numbers. Bone subunit 6a, indicating less frequent visitations by fragments and coprolites are present in low to moderate carnivores at this time, possibly linked to the presence of quantities. An elongate bone fragment (~3 cm) aligned hominins at the cave? Fine fabric microstructure is horizontally is cracked and crushed in situ. Clay aggregates consistent with cold conditions and weak cryoturbation. are often cracked and fractured in situ. Fine matrix has a Disturbance of the upper surface of this subunit, possibly granular microstructure. Sediments become increasingly by carnivores. heterogeneous and poorly sorted towards the upper limit of the thin section.

58

MM3B 6b Same as thin section MM3A

6c/1 The lower part of this thin section covers the upper part of Colluvial sheetwash, aeolian The upper part of sublayer 6c/1 is rather mixed and (upper) sublayer 6c/1. Towards the upper surface, this sublayer has a and possibly infiltration heterogeneous, possibly due to the action of burrowing very disturbed composition, possibly as a result of animals and the infilling of these voids. Freezing Bioturbation bioturbation. Also in this region there is a high concentration conditions are evident, and evidence of animals is at a of angular to sub-angular quartz sand (~40–50%), possibly the (cold and dry) minimum. coarse fill of a burrow. Stringers of desiccated clay are also present within the sand. Bone fragments and coprolites are present, but infrequent. The clay-rich matrix at the base has strongly expressed, granostriated b-fabrics surrounding coarse components.

MM4A 6c/1 This sublayer is very mixed, poorly sorted and heterogeneous Cave floor environment Disturbance of this sublayer is evident, possibly by freeze– (lower) (relative to subunits 6a and 6b). Quartz sand and clay thaw actions on the sediments and mixing by the users of aggregates are frequent. Rock fragments are common, (cold and dry) the cave (most likely hominins, given the paucity of including limestone (locally derived), volcanic and coprolites). Bone is in very good condition, supporting a metamorphic lithologies (e.g., basalt, schist) A small number cold and dry climate. Elongate bone fragment in vertical of very angular chert flakes are also present in the coarse position indicates frost heave. Sharp chert fragments fraction, as well as bone. One elongate piece of bone is probably relate to hominin use of the site. oriented vertically. Granostriated b-fabrics are evident, both around mineral grains and rock fragments, and aggregate grains. Coprolites are present in low numbers.

2987 6c/1 Similar thin section MM4A. Bone fragments are orange (ppl), Colluvial sheetwash, Aggregates of sediment from sublayer 6c/2 indicate that partially destroyed (cracked but not transported). Occurrence infiltration the sublayer was eroded and redeposited, probably by of rounded aggregates with a dense structure, clay silt sheetwash, and representing a source of material for lithology and brownish colour typical of sublayer 6c/2. (humid, then cold and dry) sublayer 6c/1. Granostriated b-fabrics are developed around large mineral grains and coprolite fragments.

59

2985 6c/2 This sublayer is a variably compacted clay silt with well- Cave floor environment, Numerous coprolites most likely reflecting occupation of expressed aggregate microstructure, a speckle-stippled b- aeolian and biogenic the cave by hyenas (Horwitz and Goldberg, 1989). fabric, and granostriated b-fabric present around coarse accumulation Aggregates of limpid clay, common in subunit 6d do not inclusions (mineral grains, coprolites and aggregates). occur here, indicating a change in source material and the Fragments of bones and coprolites are frequent; bone (cold and dry?) accumulation of allogenic sediments. Vertical and semi- fragments are rounded and cracked and some are fractured vertical orientation of flakes and elongated bone in situ forming fine angular pieces retaining close association. fragments is consistent with in situ reworking of the Coprolite fragments are yellow or yellowish brown, usually sediments and accords with the granostriated b-fabric with black spots. Some coprolites contain bones and empty and granostriated microstructure, recording cryoturbation spaces related to fur inclusions and gas bubbles. Thin and processes (Van Vliet-Vanöe, 2010). thick lithic debitage occur, oriented semi-vertically. Charcoal fragments.

2983 6c/2 Similar to thin section 2985.

2988 6d This subunit is formed of highly porous clay silt with a well- Cryoturbation Subunit 6d exhibits characteristics typical of both layer 7 developed aggregate microstructure, speckle-stippled b- and sublayer 6c/2, indicating formation by post- fabric, and a granostriated b-fabric formed around (cold and dry) depositional mixing of material near the contact zone aggregates and mineral grains. Three different components between two strata, as an effect of cryoturbation. are visible in xpl: 1) highly birefringent dense clay (sub- angular and rounded clasts), 2) birefringent compact silty clay (ovoid aggregates), and 3) loose clay silt with speckle-stippled b-fabric. Compact clay aggregates are brown in ppl, features shared with sublayer 6c/2. Highly birefringent fragments of clay aggregates or clay coatings are orange in ppl, characteristics shared with layer 7. Coarse mineral grains, bones and coprolite fragments are frequent. Elongated grains exhibit chaotic orientation, with semi-vertical orientation of some components.

MM4B 6c/1 Same as MM4A (lower)

Erosive contact Erosion

60

7 Basal layer 7 in lower part of thin section is densely compact, Karstic channelling and High frequency of clay aggregates indicate reworking of non-porous clay silt with coarse quartz silt and very frequent erosion older sediments (possibly old phreatic fills) and clay aggregates. Clay coatings of rock fragments and other incorporation into sediment matrix. Possible multiple Reworking of older phreatic inclusions (e.g., bone). Clay aggregates are rounded to sub- sources based on variable internal composition. Presence fills? angular, often comprising dark brown clay, but other of water and erosion. lithologies are present including lighter clays and silts with (humid) variable quantities of quartz silt inclusions.

2984 7 This layer is a densely compact, non-porous clay. The Reworking of older clay High frequency of clay aggregates indicate reworking of sediments are dominated by angular, sub-angular and sediments (phreatic fills?) older sediments (possibly relict phreatic fills) and rounded aggregates. Some of these aggregates are residual incorporation into younger sediment matrix. Presence of crushed clay coatings composed of orange laminated clays (humid) water and erosion. and silts. Most aggregates are clasts of non-structural orange- brown or brown limpid clay. Other lithologies occur infrequently and are represented by sub-angular clasts of highly weathered rock grains.

2989 7 The greenish subunit is a densely compact, non-porous clay Water ponds Presence of pyrite indicates anoxic conditions, related to silt with rounded sand-sized grains of weathered rock. Some saturation zone or subaqueous environment. The of these grains reveal quartz crystals (aphanitic volcanic rock (humid) greenish intercalations were deposited in temporal water or quartzite). Non-transparent square mineral crystals occur ponds, while typical facies of layer 7 is consistent with both inside and outside of the weathered grains, most drier conditions. probably pyrite.

61

Table S2. Radiocarbon (14C) ages, isotopic values, C:N ratios and % collagen yields for 20 bison remains from Chagyrskaya Cave.

MPI lab code Layer, Human % δ13C δ15N AMS lab code 14C age Calibrated age range (yr cal. BP) b Skeletal element % C % N C:N (S-EVA) horizon modification collagen (‰) a (‰) a (MAMS-) (yr BP) b 68.2% CI 95.4% CI

24479 5, 1 incisor none 9.1 –18.7 9.1 41.8 15.0 3.3 14954 33,760 ± 170 38,540–38,060 38,690–37,670 24480 5, 2 incisor none 5.9 –18.5 7.8 42.3 15.4 3.2 14955 >49,000 – – 24481 5, 5 phalanx none 0.9 –19.0 8.7 35.9 13.1 3.2 14956 4,497 ± 26 5,283–5,054 5,292–5,046 24482 6a, 1 phalanx none 7.9 –19.1 5.4 52.5 19.2 3.2 14957 >49,000 – – 24483 6b, 3 rib cutmarks 6.1 –19.3 8.8 36.2 13.2 3.2 14958 >49,000 – – 24484 6b, 4 metatarsal fragment none 8.5 –19.4 9.1 43.6 15.9 3.2 14959 >49,000 – – 23051 6b, 4 longbone fragment cutmarks 4.3 –19.8 6.8 22.5 8.4 3.1 14353 >52,000 – – possible 23052 6b, 4 longbone fragment 6.4 –18.8 9.6 23.1 8.7 3.1 14354 >52,000 – – cutmark 22314 6c/1, 1 rib fragment cutmarks 5.2 –19.2 8.9 28.7 10.8 3.1 13033 45,672 ± 481 49,740–48,630 >50,000–48,110 23053 6c/1, 1 longbone fragment cutmarks 4.5 –20.6 5.2 27.3 10.2 3.1 14355 >52,000 – – 24485 6c/1, 1 longbone fragment impact mark 6.6 –18.9 6.3 4.6 15.9 3.2 14960 >49,000 – – 22315 6c/1, 2 rib fragment none 8.2 –19.4 7.9 29.6 11.1 3.1 13034 48,724 ± 692 49,460–48,050 >50,000–47,440 22316 6c/1, 3 longbone fragment cutmarks 5.8 –19.5 7.7 38.2 14.2 3.1 13035 50,524 ± 833 – – possible 23054 6c/1, 3 longbone fragment 4.3 –18.8 8.1 24.9 9.3 3.1 14356 >52,000 – – impact mark 23055 6c/1, 3 longbone fragment cutmarks 4.3 –19.2 7.1 24.0 9.0 3.1 14357 >52,000 – – possible 23056 6c/1, 3 longbone fragment 3.1 –19.2 10.5 20.4 7.7 3.1 14358 >52,000 – – impact mark 24486 6c/1, 4 bone fragment cutmarks 2.7 –19.5 6.4 36.4 12.9 3.3 14961 >49,000 – – 24487 6c/1, 5 rib fragment cutmarks 0.8 –21.2 7.3 42.3 13.6 3.6 14962 17,630 ± 50 21,450–21,200 21,550–21,060 24488 6c/1, 5 longbone fragment impact mark 7.1 –19.2 8.0 43.8 15.7 3.2 14963 >49,000 – – 24489 6c/2 phalanx none 5.0 –19.1 7.5 41.7 15.1 3.2 14964 >49,000 – –

a δ13C values reported relative to the Vienna Pee Dee Belemnite (VPDB) standard and δ15N values reported relative to the air standard. b Ages calibrated using OxCal v4.3 (Bronk Ramsey, 2009) and the IntCal13 data set (Reimer et al., 2013), with age ranges estimated at the 68.2% and 95.4% confidence intervals.

62

Table S3. Dose rate data, De values and optical ages for sediment samples from layers 5, 6 and 7.

Dose rates (Gy/ka) b Number of Water Total dose rate d OD value g Sample code De (Gy) grains or Optical age (ka) content (%) a (Gy/ka) c (%) e Beta Gamma Cosmic aliquots f

Layer 5 CHAG12-1 18.6 1.35 ± 0.08 0.74 ± 0.04 0.03 2.92 ± 0.13 141.3 ± 4.1 41 ± 2 (23) 178 (1600) 48.3 ± (1.8, 2.8) CHAG12-2 15.9 1.25 ± 0.07 0.63 ± 0.04 0.03 2.71 ± 0.13 145.7 ± 4.7 44 ± 2 (23) 149 (1500) 53.7 ± (2.4, 3.2) CHAG14-2 18.5 1.45 ± 0.08 0.77 ± 0.05 0.03 3.05 ± 0.14 171.4 ± 9.3 57 ± 3 (24) 118 (1400) 56.3 ± (3.3, 4.1) CHAG14-3 21.1 1.28 ± 0.07 0.68 ± 0.04 0.03 2.79 ± 0.13 164.3 ± 5.4 57 ± 4 (20) 112 (1000) 58.9 ± (2.4, 3.5) CHAG14-4 24.9 1.29 ± 0.07 0.70 ± 0.04 0.03 2.82 ± 0.13 153.6 ± 5.2 44 ± 3 (21) 114 (1000) 54.5 ± (2.2, 3.3) CHAG14-5 22.8 1.32 ± 0.08 0.73 ± 0.04 0.03 2.88 ± 0.13 149.4 ± 5.6 49 ± 3 (24) 143 (1400) 51.9 ± (2.3, 3.2) Subunit 6a CHAG12-3 19.4 1.38 ± 0.08 0.76 ± 0.04 0.03 2.96 ± 0.13 158.2 ± 6.9 38 ± 3 (21) 82 (500) 53.3 ± (2.8, 3.5) CHAG12-4 17.8 1.39 ± 0.08 0.74 ± 0.04 0.03 2.96 ± 0.13 146.7 ± 6.2 42 ± 2 (24) 111 (1100) 49.6 ± (2.6, 3.2) CHAG14-6 20.5 1.43 ± 0.08 0.77 ± 0.05 0.03 3.03 ± 0.14 144.3 ± 5.6 46 ± 3 (22) 98 (1400) 47.7 ± (2.1, 3.0) CHAG14-7 20.0 1.43 ± 0.08 0.80 ± 0.05 0.03 3.06 ± 0.14 161.0 ± 5.9 50 ± 3 (22) 126 (1400) 52.6 ± (2.3, 3.2) Subunit 6b CHAG12-5 18.2 1.30 ± 0.08 0.73 ± 0.04 0.03 2.86 ± 0.13 151.8 ± 6.2 37 ± 2 (24) 118 (1000) 53.1 ± (2.5, 3.4) CHAG14-8 20.3 1.48 ± 0.08 0.87 ± 0.05 0.03 3.17 ± 0.14 162.2 ± 6.8 45 ± 3 (21) 91 (900) 51.1 ± (2.4, 3.3) Sublayer 6c/1 CHAG12-6 13.8 1.61 ± 0.09 0.76 ± 0.04 0.03 3.19 ± 0.14 186.4 ± 11.3 34 ± 2 (26) 129 (1000) 58.5 ± (3.9, 4.5) CHAG12-7 12.8 1.58 ± 0.09 0.76 ± 0.04 0.03 3.17 ± 0.14 171.1 ± 6.8 49 ± 4 (18) 64 (400) 53.9 ± (2.7, 3.4) CHAG14-9 24.8 1.38 ± 0.08 0.95 ± 0.06 0.03 3.16 ± 0.14 165.2 ± 6.6 54 ± 3 (24) 141 (1300) 52.3 ± (2.4, 3.3) CHAG17-5 17.4 1.61 ± 0.09 0.27 ± 0.02 0.04 2.72 ± 0.14 156.2 ± 5.6 30 ± 2 (30) 130 (1300) 57.5 ± (2.5, 3.7) CHAG17-8 11.8 1.51 ± 0.09 0.50 ± 0.03 0.04 2.86 ± 0.14 172.5 ± 5.9 35 ± 2 (22) 144 (1500) 60.3 ± (2.5, 3.7) CHAG17-9 15.6 1.56 ± 0.09 0.42 ± 0.02 0.04 2.82 ± 0.14 148.9 ± 6.9 42 ± 3 (22) 74 (1000) 52.8 ± (2.7, 3.7) Sublayer 6c/2 CHAG12-10 13.5 1.95 ± 0.11 0.89 ± 0.05 0.04 3.67 ± 0.16 194.0 ± 4.1 28 ± 2 (16) 172 (1000) 52.9 ± (1.6, 2.7) CHAG14-12 23.2 1.96 ± 0.11 1.02 ± 0.06 0.03 3.81 ± 0.16 202.4 ± 7.6 31 ± 2 (22) 172 (1400) 53.2 ± (2.3, 3.2) CHAG17-3 14.8 1.89 ± 0.11 1.13 ± 0.07 0.04 3.86 ± 0.16 210.8 ± 15.0 49 ± 3 (24) 70 (1400) 54.6 ± (4.0, 4.6) Subunit 6d CHAG17-6 20.1 1.67 ± 0.10 0.40 ± 0.02 0.04 2.91 ± 0.14 184.0 ± 8.4 38 ± 3 (18) 80 (700) 63.2 ± (3.3, 4.4) CHAG17-7 20.1 1.70 ± 0.10 0.47 ± 0.03 0.04 3.00 ± 0.14 178.2 ± 10.3 38 ± 3 (26) 86 (500) 59.3 ± (3.7, 4.6)

63

Layer 7

CHAG12-9 18.8 1.78 ± 0.10 0.97 ± 0.06 0.03 3.57 ± 0.15 1203 ± 80 38 ± 4 (33) 120 (600) 337 ± (23, 27)

1164 ± 59 – 6 (6) 326 ± (18, 23)

CHAG14-10 23.2 2.03 ± 0.12 1.10 ± 0.06 0.03 3.97 ± 0.17 1266 ± 347 39 ± 3 (29) 59 (400) 319 ± (88, 89)

2.15 ± 0.13 1.10 ± 0.06 0.03 3.71 ± 0.15 1145 ± 45 – 6 (6) 309 ± (16, 18)

CHAG14-11 22.1 2.14 ± 0.12 1.13 ± 0.07 0.03 4.11 ± 0.17 1332 ± 73 – 6 (6) 324 ± (19, 23)

CHAG17-4 9.5 1.98 ± 0.12 0.81 ± 0.05 0.04 3.32 ± 0.15 1151 ± 44 – 6 (6) 347 ± (15, 21)

a Measured (field) water contents, expressed as mass of water to mass of dry sample, multiplied by 100. The total dose rates and ages were calculated using a long-term water content of 20 ± 5% (at 1σ) for all samples. b Beta dose rates were measured by low-level beta counting and gamma dose rates by in situ (field) gamma spectrometry. Cosmic-ray dose rates were estimated from published equations and assigned a relative uncertainty of 15%. These external dose rate components have each been adjusted for the long-term water content. c Mean ± total uncertainty (at 1σ). Grains of 180–212 μm in diameter were measured for all samples except CHAG17-4 (125–150 μm) and CHAG14-10 (90 –125 μm, MAR procedure) . An internal dose rate of 0.80 ± 0.10 Gy/ka is included for all samples except CHAG17-4 (0.49 ± 0.05 Gy/ka) and CHAG14-10 (0.42 ± 0.05 Gy/ka, MAR procedure). d Equivalent dose (De) values, calculated using the finite mixture model (FMM) or, for CHAG12-10 and all samples from layer 7, the central age model (CAM). For samples fitted using the FMM, the De value corresponds to the the component containing the largest proportion of grains; this component was used for age determination. For the other samples, the CAM was applied after rejecting outliers using the normalised median absolute deviation. e Overdispersion (OD) is the scatter remaining in De values after taking measurement uncertainties into account. OD values (± 1σ uncertainties) are listed for the full De distributions. The OD values in brackets are the point estimates corresponding to the optimal FMM fits (estimated using a combination of maximum log likelihood and Bayes Information Criterion) or, for CHAG12-9 and CHAG12-10, the point estimates after outlier rejection. f Number of individual grains (or multi-grain aliquots for some of the samples from layer 7) accepted and used for De estimation (i.e., the number associated with the FMM component containing the largest proportion of De values or, for CHAG12-9 and CHAG12-10, the number remaining after outlier rejection). The total number of grains or aliquots measured is shown in brackets. g Mean ± (random uncertainty, total uncertainty). Both uncertainties are at 1σ. The total uncertainty includes a relative systematic error of 2% added (in quadrature) to the propagated random uncertainties to allow for any bias associated with calibration of the laboratory beta source.

64

Table S4. Human remains from Chagyrskaya Cave.

Year of Excavation Stratigraphic Specimen Anatomical element discovery square subunit/sublayer

Chagyrskaya 1 2008 Л6 6b lower left dc Chagyrskaya 2 2009 M8 6b atlas fragment, child Chagyrskaya 3 2009 Л8 6c/1 upper premolar fragment Chagyrskaya 4 2009 M8 6c/1 lower left incisor, very worn Chagyrskaya 5 2009 M8 6c/1 patella Chagyrskaya 6 2011 Н9 6b right mandible fragment with C–M2 Chagyrskaya 7 2012 Н11 6c/1 thoracal vertebral process fragment Chagyrskaya 8 2011 Н10 6b distal manual phalanx Chagyrskaya 9 2011 Н10 6a left proximal ulna fragment Chagyrskaya 10 2011 Н10 6b left UM?1 Chagyrskaya 11 2012 Н11 6b I1, very worn

Chagyrskaya 12 2012 M10 6c/1 left P3 Chagyrskaya 13 2013 O12 6b left I1

Chagyrskaya 14 2011 Н10 6b left I2

Chagyrskaya 17 2013 M12 6c/1 right P4 Chagyrskaya 18 2012 Н10, M10 6c/1 left dm1 Chagyrskaya 19 2012 O11 6a left dm2 Chagyrskaya 20 2012 M10 6c/1 right upper? dc various fragments, including tibial shaft Chagyrskaya 21 2011 Н10 6b fragment and ischial tuberosity Chagyrskaya 22 2011 Н10 6b middle phalanx of foot Chagyrskaya 23 2011 Н10 6b left 3rd metacarpal Chagyrskaya 24a 2011 Н10 6b left tibia, distal articular end Chagyrskaya 24b 2011 Н10 6b left fibula, distal articular end Chagyrskaya 26 2012 Н11 6b thoracal vertebra, T3–T5? Chagyrskaya 27 2012 Н11 6b left medial cuneiform spinous process, thoracal vertebra, probably Chagyrskaya 29 2012 Н11 6c/1 upper (T1–T7) Chagyrskaya 30 2012 Н11 6b sternum fragment Chagyrskaya 31 2012 Н10 6b middle pedal phalanx fragment Chagyrskaya 32 2012 O11 6c/1 proximal phalanx, hallux Chagyrskaya 34 2012 Н11 6b small fragments of vertebral process Chagyrskaya 35 2012 Н11 6b L5 vertebra Chagyrskaya 36 2012 Н11 6b left calcaneus Chagyrskaya 37 2012 Н11 6b C1, atlas Chagyrskaya 38 2012 Н11 6b C2, axis (fragmentary) Chagyrskaya 39a 2015 K6 6c/2 right ulna, middle half of shaft Chagyrskaya 39b 2015 K6 6c/2 right radius, proximal part

Chagyrskaya 41 2015 K6 6c/1 right P3 Chagyrskaya 42 2015 K6 6c/1 fragment of base of MC or MT Chagyrskaya 43 2015 K6 6c/1 end phalanx of foot Chagyrskaya 44 2015 K6 6c/1 end phalanx of foot

65

Chagyrskaya 45a 2015 K6 6c/2 distal fragment of (left?) ulna Chagyrskaya 45b 2015 K6 6c/2 ?4th ?left metacarpal fragment Chagyrskaya 45c 2015 K6 6c/2 right hamate Chagyrskaya 45d 2015 K6 6c/2 right MC1, distal half fragment, possibly part of MC base, or Chagyrskaya 45e 2015 K6 6c/2 phalanx base? Chagyrskaya 46 2015 K6 6c/1 right talus, very abraded Chagyrskaya 47a 2015 K6 6c/2 radius, distal articular surface Chagyrskaya 47b 2015 K6 6c/2 radius, proximal end Chagyrskaya 48a 2015 K6 6c/2 right ulna, proximal end Chagyrskaya 48b 2015 K6 6c/2 scapula fragment, base of acromion Chagyrskaya 48c 2015 K6 6c/2 scapula fragment, axillary border Chagyrskaya 50 2015 K6 6c/2 left P Chagyrskaya 51a 2015 K6 6c/2 left UM1/2 fragment Chagyrskaya 51b 2015 K6 6c/2 lower M?1 fragment Chagyrskaya 52a 2012 Н11 6b left 5th metatarsal, proximal 2/3 Chagyrskaya 52b 2012 Н11 6b left 4th metatarsal, proximal 2/3 Chagyrskaya 53 2012 Н11 6b left 3rd metatarsal, complete Chagyrskaya 54 2012 Н11 6b sternal end of rib Chagyrskaya 55a,b 2012 M11 6b right clavicle Chagyrskaya 56a 2012 Н11 6b distal thumb phalanx Chagyrskaya 56b 2012 Н11 6b distal manual phalanx, possibly II ray middle phalanx of hand, possibly II ray, fits Chagyrskaya 56c 2012 Н11 6b with 56b Chagyrskaya 57 2012 Н11 6b left maxilla fragment with M2 and M3 Chagyrskaya 58 2015 K6 6c/2 right humerus, distal 2/3 Chagyrskaya 59 2016 K7 6c/2 left upper M (2 or 3?) manual middle phalanx (V?), associated with Chagyrskaya 60 2016 K7 6c/2 Chagyrskaya 61 manual terminal phalanx (V?), associated Chagyrskaya 61 2016 K7 6c/2 with Chagyrskaya 60 Chagyrskaya 62 2016 K7 6c/2 terminal phalanx, hallux Chagyrskaya 63 2016 И7 6a left upper M2/3 germ Chagyrskaya 64 2016 Н9 6c/2 left lower M2?, crown completely worn away Chagyrskaya 65 2016 И7 6a pedal middle phalanx, IV/V? Chagyrskaya 66 2016 K7 6c/2 lumbar vertebra, not well preserved Chagyrskaya 67 2017 Л6 6c/2 lateral portion of left clavicle Chagyrskaya 68 2017 L6 6d 1st metacarpal

66

Table S5. Breakdown of lithic assemblage (Chagyrskaya Cave, subunits/sublayers 6a–6c/2).

6a 6b 6c/1 6c/2 TOTAL Type Number % % a Number % % a Number % % a Number % % a Number % % a pre-cores 0 0 0 0 0 0 3 0.10 0.21 0 0,00 0,00 3 0,07 0,14 cores 2 0.62 1.64 2 0.42 0.86 27 0.89 1.88 4 0,92 1,32 35 0,82 1,68 pre-forms 0 0 0 1 0.21 0.43 8 0.26 0.56 0 0,00 0,00 9 0,21 0,43 tools 31 9.66 25.41 61 12.87 26.29 428 14.17 29.83 69 15,94 22,85 591 13,91 28,29 flakes 83 25.86 68.03 163 34.39 70.26 874 28.93 60.91 213 49,19 70,53 1333 31,37 63,81 blades 4 1.25 3.28 3 0.63 1.29 82 2.71 5.71 15 3,46 4,97 104 2,45 4,98 unidentifiable debitage 0 0 0 0 0 0 13 0.43 0.91 1 0,23 0,33 14 0,33 0,67 chips 169 52.65 – 210 44.30 – 1409 46.64 – 76 17,55 – 1864 43,87 – chunks 30 9.35 – 34 7.17 – 177 5.86 – 55 12,70 – 296 6,97 – TOTAL 319 99.38 100 474 100 100 3021 100 100 433 100 100 4249 100 100 a Percentage when chips and chunks are omitted from the total.

67

Table S6. Breakdown of cores (Chagyrskaya Cave, subunits/sublayers 6a–6c/2).

6a 6b 6c/1 6c/2 TOTAL Type Number Number Number Number Number PRE-CORES – – 3 – 3 unidirectional, triangular, naturally convex back – – 1 – 1 bifacial, orthogonal/unidirectional – – 1 – 1 unidentifiable, ovoid, naturally flat back – – 1 – 1 CORES 2 2 27 4 35 radial, circular, naturally convex back – – 1 1 2 radial, ovoid, naturally convex back – – 1 1 2 radial, rectangular, naturally convex back 1 – 1 1 2 radial-pyramidal, ovoid, naturally convex back – – 2 – 2 convergent, unidentifiable, flattened back – – 1 – 1 unidirectional, unidentifiable, naturally concave back – – 1 – 1 unidirectional, unidentifiable – – 1 – 1 unidirectional, rectangular, naturally flat back 1 1 – 1 4 bitransversal, rectangular, naturally convex back – – 1 – 1 orthogonal, unidentifiable – – 1 – 1 semi-crossed, ovoid, naturally convex back – – 1 – 1 semi-crossed, rectangular, flattened back – – 1 – 1 semi-crossed, rectangular, naturally convex back – 1 1 – 1 crossed, rectangular, naturally convex back – – 1 – 1 bifacial, bidirectional/transverse, rectangular – – 1 – 1 bifacial, bitransversal/unidirectional, rectangular – – 1 – 1 unidentifiable – – 11 – 11 TOTAL 2 2 30 4 38

68

Table S7. Composition of blank assemblage (Chagyrskaya Cave, subunits/sublayers 6a–6c/2).

6a 6b 6c/1 6c/2 TOTAL Type Number % % a Number % % a Number % % a Number % % a Number % % a blades, regular 5 4.42 4.42 4 1.82 1.86 74 5.39 5.5 13 4,45 4,55 96 4,80 4,90 blades, cortical débordant 0 0 0 0 0 0 15 1.09 1.11 0 0,00 0,00 15 0,75 0,77 blades, lateral débordant 1 0.88 0.88 0 0 0 9 0.66 0.67 1 0,34 0,35 11 0,55 0,56 blades, crested débordant 0 0 0 0 0 0 7 0.51 0.52 1 0,34 0,35 8 0,40 0,41 blades, radial core débordant 0 0 0 0 0 0 1 0.07 0.07 1 0,34 0,35 2 0,10 0,10 blades, bifacial thinning 0 0 0 0 0 0 1 0.07 0.07 0 0,00 0,00 1 0,05 0,05 blades, primary 0 0 0 0 0 0 6 0.44 0.45 0 0,00 0,00 6 0,30 0,31 flakes 55 48.67 48.67 110 50.00 51.16 750 54.62 55.72 190 65,07 66,43 1105 55,31 56,38 flakes, cortical débordant 0 0 0 10 4.55 4.65 94 6.85 6.98 14 4,79 4,90 118 5,91 6,02 flakes, lateral débordant 10 8.85 8.85 25 11.36 11.63 70 5.1 5.2 10 3,42 3,50 115 5,76 5,87 flakes, crested débordant 1 0.88 0.88 3 1.36 1.40 21 1.53 1.56 3 1,03 1,05 28 1,40 1,43 flakes, crested 0 0 0 1 0.45 0.47 1 0.07 0.07 1 0,34 0,35 3 0,15 0,15 flakes, radial core débordant 18 15.93 15.93 30 13.64 13.95 87 6.34 6.46 13 4,45 4,55 148 7,41 7,55 flakes, technical/radial core débordant 0 0 0 0 0 0 1 0.07 0.07 0 0,00 0,00 1 0,05 0,05 flakes, technical 2 1.77 1.77 4 1.82 1.86 42 3.06 3.12 1 0,34 0,35 49 2,45 2,50 flakes, bifacial thinning 3 2.65 2.65 4 1.82 1.86 29 2.11 2.15 4 1,37 1,40 40 2,00 2,04 flakes, bifacial thinning, overpassed 1 0.88 0.88 0 0 0 5 0.36 0.37 0 0,00 0,00 6 0,30 0,31 flakes, primary 17 15.04 15.04 24 10.91 11.16 133 9.69 9.88 34 11,64 11,89 208 10,41 10,61 unidentifiable debitage 0 0 – 5 2.27 – 27 1.97 – 6 2,05 – 38 1,90 – TOTAL 113 100 100 220 100 100 1373 100 100 292 100 100 1998 100 100 a Percentage when unidentifiable debitage is omitted from the total.

69

Table S8. Composition of chip assemblage by size (Chagyrskaya Cave, subunits/sublayers 6a–6c/2).

< 1.0 cm 1.1–2.0 cm 2.1–2.9 cm TOTAL Type 6a 6b 6c/1 6c/2 6a 6b 6c/1 6c/2 6a 6b 6c/1 6c/2 Number % a bifacial thinning 0 – 1 0 3 4 43 3 6 15 83 4 162 16.91023 regular 1 4 17 0 25 29 186 7 50 68 380 29 796 83.08977 unidentifiable 2 2 28 0 38 47 279 9 44 41 392 24 906 – TOTAL 3 6 46 0 66 80 508 19 100 119 855 57 1859 100 a Percentage when unidentifiable chips are omitted from the total.

70

Table S9. Shape of blanks (Chagyrskaya Cave, subunits/sublayers 6a–6c/2). This table continues on the next page.

Rectangular Trapezoidal Triangular Rectangular Trapezoidal Triangular elongated elongated elongated Type

6a 6a 6a 6a 6a 6a

6b 6b 6b 6b 6b 6b

6c/1 6c/2 6c/1 6c/2 6c/1 6c/2 6c/1 6c/2 6c/1 6c/2 6c/1 6c/2 blades, regular – – – 3 – – 42 – – – – – – – 8 1 1 – – – 3 1 7 3 blades, cortical débordant – – – – – – 10 – – – – – – – 1 – – – – – – – 1 blades, lateral débordant – – – – 1 – 5 1 – – – – – – 1 – – – – – – – – blades, crested débordant – – – – – – – – – – – – – – – – – – – – – – 3 1 blades, radial core débordant – – – – – – – – – – – – – – – – – – – – – – – 1 blades, bifacial thinning – – – – – – 1 – – – – – – – – – – – – – – – – blades, primary – – – – – 2 – – – – – – – – – – – – – – – – flakes 5 8 99 15 – – 22 4 16 30 174 35 2 – 35 5 3 10 30 20 1 2 6 8 flakes, cortical débordant – 2 12 1 – 1 5 – – – 25 6 – – 6 – – 2 5 1 – 1 1 1 flakes, lateral débordant – 4 17 – 1 2 3 1 3 5 13 2 – 3 8 – 3 2 3 1 1 1 1 1 flakes, crested débordant – – 2 1 – – 2 1 – 2 7 – 1 – 1 – – 1 – 1 – – 1 – flakes, crested – 1 – – – – – – – – – – – – 1 – – – – 1 – – – – flakes, radial core débordant – 1 7 – – – 4 – 11 19 48 9 – – 4 – – 2 6 – – 1 – – flakes, technical/ radial core débordant – – – – – – – – – – 1 – – – – – – – – – – – – – flakes, technical – – 8 – – – 1 – 1 4 29 1 – – – – – – – – – – – – flakes, bifacial thinning 1 4 – – – – – 2 4 18 2 – – – – – – – 1 – – – flakes, bifacial thinning, overpassed – – – – – – – – 1 – 1 – – – – – – – 1 – – – – – flakes, primary – 4 11 6 2 – 2 – 3 5 32 5 – – 5 – 1 – 2 – – – – – unidentifiable debitage – – – – – – 1 – – – 1 – – – – – – – – – – – – – TOTAL 6 20 160 26 4 3 100 7 37 69 349 60 3 3 70 6 8 17 47 25 5 6 20 15 % a 0.4 1.4 11.4 1.9 0.3 0.2 7.1 0.5 2.6 4.9 24.9 4.3 0.2 0.2 5 0.4 0.6 1.2 3.4 1.8 0.4 0.4 1.4 1.1 a Percentage when blanks of unidentifiable shape are omitted from the total.

71

Table S9 continued.

Crescent Leaf-shaped Ovoid Irregular Unidentifiable Type TOTAL

6a 6a 6a 6a 6a

6b 6b 6b 6b 6b

6c/1 6c/2 6c/1 6c/2 6c/1 6c/2 6c/1 6c/2 6c/1 6c/2

blades, regular – 1 1 – – – 3 – – 1 – – – – 4 1 1 1 9 5 96 blades, cortical débordant – – 1 – – – – – – – – – – – – – – – 2 – 15 blades, lateral débordant – – – – – – 1 – – – – – – – 1 – – – 1 – 11 blades, crested débordant – – – – – – – – – – – – – – 3 – – – 1 – 8 blades, radial core débordant – – – – – – – – – – – – – – 1 – – – – – 2 blades, bifacial thinning – – – – – – – – – – – – – – – – – – – – 1 blades, primary – – – – – – 3 – – – 1 – – – – – – – – 6

flakes 2 2 23 5 – 2 35 2 4 2 29 10 4 12 33 17 18 42 264 69 1105 flakes, cortical débordant – – 3 2 – – 2 – – 1 3 – – 1 4 1 – 2 28 2 118 flakes, lateral débordant – 2 4 2 – – – – – – 1 – – 1 4 – 2 5 16 3 115 flakes, crested débordant – – 2 – – – – – – – – – – – 1 – – – 5 – 28 flakes, crested – – – – – – – – – – – – – – – – – – 3

flakes, radial core débordant – 3 1 1 – – 3 – – 1 7 – 2 1 3 1 5 2 4 2 148 flakes, technical/ radial core débordant – – – – – – – – – – – – – – – – – – – – 1 flakes, technical – – 2 – – – – – 1 – 2 – – – – – – – – – 49 flakes, bifacial thinning – – – – – – – – – – 3 1 – – 3 – – – 1 – 40 flakes, bifacial thinning, overpassed – – – – – – – – – – – – – – – – – – 3 – 6 flakes, primary – – 1 1 – – 9 – 3 7 25 5 1 4 1 4 7 4 45 13 208 unidentifiable debitage – – – – – – – – – – 1 – – – – – – 5 24 6 38 TOTAL 2 8 38 11 0 2 56 2 8 12 72 16 7 19 58 24 33 61 403 100 1998 % a 0.1 0.6 2.7 0.8 0 0.1 4 0.1 0.6 0.9 5.1 1.1 0.5 1.4 4.1 1.7 – – – – 100 a Percentage when blanks of unidentifiable shape are omitted from the total.

72

Table S10. Proportion of cortex surface by blank type (Chagyrskaya Cave, subunits/sublayers 6a–6c/2).

0% 1–25% 26–50% 51–75% 76–100%

Type TOTAL

6a 6a 6a 6a 6a

6b 6b 6b 6b 6b

6c/1 6c/2 6c/1 6c/2 6c/1 6c/2 6c/1 6c/2 6c/1 6c/2

blades, regular 5 3 42 12 – – 6 – – – 23 1 – 1 3 – – – – – 96 blades, cortical débordant – – – – – – 8 – – – 7 – – – – – – – – – 15 blades, lateral débordant – – 6 – – – 3 1 1 – – – – – – – – – – – 11 blades, crested débordant – – 5 1 – – – – – – 2 – – – – – – – – – 8 blades, radial core 2 – – – 1 – – 1 – – – – – – – – – – – – – débordant blades, bifacial thinning – – – – – – – – – – – – – – 1 – – – – – 1 blades, primary – – – – – – – – – – – – – – – – – – 6 – 6 flakes 44 88 535 142 7 10 102 22 4 9 85 21 – 3 28 3 – – – 2 1105 flakes, cortical débordant – – 3 – – 4 72 12 – 5 15 2 – 1 2 – – – 2 – 118 flakes, lateral débordant 5 15 52 7 4 6 11 3 1 1 3 – – 3 2 – – – 2 – 115 flakes, crested débordant – 3 16 3 1 – 3 – – – 1 – – – – – – – 1 – 28 flakes, crested – 1 1 1 – – – – – – – – – – – – – – – – 3 flakes, radial core 148 12 19 61 7 5 6 13 4 – 3 4 – – 1 4 – 1 1 5 2 débordant flakes, technical/radial – – – – – – – – – – 1 – – – – – – – – – 1 core débordant flakes, technical – – 10 – 1 2 4 – 1 1 15 1 – 1 9 – – – 4 – 49 flakes, bifacial thinning 3 4 21 4 – – 6 1 – – – – – – – – – – 2 – 40 flakes, bifacial thinning, 6 1 – 2 – – – 1 – – – – – – – 1 – – – 1 – overpassed flakes, primary – – – – – – – – – – – – – – – 2 17 24 133 32 208 unidentifiable debitage – 5 22 5 – – 2 1 – – 1 – – – 1 – – – 1 – 38 TOTAL 70 138 776 183 18 28 232 44 7 19 157 24 0 10 51 5 18 25 157 36 1998 % 62.0 62.7 56.5 62.7 15.9 12.7 16.9 15.1 6.2 8.6 11.4 8.2 0 4.6 3.7 1.7 15.9 11.7 11.4 12.3

73

Table S11. Breakdown of tool kit (Chagyrskaya Сave, subunits/sublayers 6a–6c/2).

6a 6b 6c/1 6c/2 TOTAL Type Number % % a Number % % a Number % % a Number % % a Number % % a

POINTS 3 9.7 12.5 7 11 17 33 7.7 14 3 4.3 7.3 46 7.8 13 distal, dorsal – – – 2 – – 4 – – – – – 6

sub-triangular, dorsal 1 – – – – – 3 – – 1 – – 5

sub-trapezoidal, dorsal 1 – – – – – – – – – – – 1

sub-leaf, dorsal – – – – – – 1 – – 1 – – 2

semi-triangular, dorsal – – – 2 – – – – – – – – 2

semi-trapezoidal, dorsal – – – 1 – – 6 – – – – – 7

semi-crescent, dorsal – – – – – – 3 – – – – – 3

semi-crescent, alternate – – – – – – – – – – – – 0

semi-leaf, dorsal – – – 1 – – 7 – – – – – 8

semi-leaf, alternate 1 – – 1 – – 1 – – – – – 3

semi-leaf asymmetrical, dorsal – – – – – – 5 – – – – – 5

leaf-shaped, dorsal – – – – – – – – – 1 – – 1 – –

leaf-shaped asymmetrical, dorsal/bifacial – – – – – – 1 – – – – – 1

unidentifiable, dorsal – – – – – – 1 – – – – – 1

unidentifiable, alternate – – – – – – 1 – – – – – 1

SCRAPERS 14 45 58.3 22 36 54 169 39 72 35 51 85 240 41 70 transverse-straight, dorsal – – – 3 – – 2 – – 3 – – 8

transverse-convex, dorsal – – – 2 – – 6 – – 3 – – 11

transverse-convex, ventral – – – – – – 1 – – 1 – – 2

diagonal-straight, dorsal – – – 1 – – 9 – – 1 – – 11

diagonal-straight, ventral – – – – – – 1 – – – – – 1

diagonal-convex, dorsal 1 – – 2 – – 12 – – – – – 15

diagonal-convex, ventral – – – – – – 2 – – – – – 2

74 straight, dorsal 4 – – 3 – – 21 – – 4 – – 32 straight, ventral 2 – – – – – 1 – – – – – 3 convex, dorsal 2 – – – – – 26 – – 6 – – 34 convex, ventral – – – – – – 2 – – 1 – – 3 convex, alternating – – – – – – 1 – – – – – 1 convex, bifacially retouched – – – – – – 1 – – – – – 1 wavy, dorsal – – – – – – 3 – – – – – 3 double-straight, dorsal/bifacial – – – – – – 1 – – – – – 1 straight-convex, dorsal – – – – – – 2 – – – – – 2 straight-convex, alternate – – – – – – 1 – – – – – 1 double-convex, dorsal – – – – – – 2 – – – – – 2 sub-triangular, dorsal – – – – – – 1 – – – – – 1 sub-trapezoidal, dorsal 1 – – 4 – – 7 – – 4 – – 16 sub-trapezoidal, alternate – – – 2 – – – – – – – – 2 sub-crescent, dorsal – – – – – – 4 – – 1 – – 5 sub-crescent, alternate – – – 1 – – 1 – – – – – 2 sub-leaf, dorsal – – – 1 – – 2 – – 3 – – 6 sub-leaf, alternate – – – – – – 1 – – – – – 1 semi-trapezoidal, dorsal 2 – – 1 – – 24 – – 2 – – 29 semi-trapezoidal, ventral – – – – – – 2 – – – – – 2 semi-trapezoidal, alternate – – – – – – 6 – – – – – 6 semi-recrangular, dorsal – – – 1 – – 4 – – – – – 5 semi-rectangular, ventral – – – – – – 1 – – – – – 1 semi-crescent, dorsal 1 – – 1 – – 5 – – 2 – – 9 semi-crescent, ventral – – – – – – 1 – – – – – 1 semi-crescent, alternate – – – – – – – – – – – – 0 semi-leaf, dorsal 1 – – – – – 9 – – 1 – – 11 semi-leaf, alternate – – – – – – 1 – – – – – 1 semi-ovoid, alternating – – – – – – 1 – – 1 – – 1 triangular, dorsal – – – – – – 2 – – – – – 1

75 convergent, dorsal – – – – – – 3 – – 1 – – 2 crescent, dorsal – – – – – – – – – 1 – – 1 – –

DENTICULATES – – – 2 3.3 4.9 4 0.9 1.7 – – – 6 1.1 1.7 transverse-convex, dorsal – – – – – – 1 – – – – – 1 straight, dorsal – – – 2 – – 1 – – – – – 3 convex, dorsal – – – – – – 2 – – – – – 2

NOTCHES – – – 2 3.3 4.9 4 0.9 1.7 – – – 6 1.1 1.7 lateral, dorsal – – – 2 – – 1 – – – – – 3 lateral, ventral – – – – – – 3 – – – – – 3

TRUNCATIONS – – – – – – 9 2.1 3.8 – – – 9 1.5 2.6 END-SCRAPERS – – – – – – 2 0.5 0.9 – – – 2 0.3 0.6 BIFACIAL POINTS – – – 1 1.6 2.4 6 1.4 2.5 1 1.4– 2.4 8 1.4 2.3 sub-triangular, thinned base, retouched back – – – – – – 1 – – – – – 1 semi-trapezoidal, naturally backed – – – – – – 1 – – – – – 1 semi-leaf – – – – – – 1 – – – – – 1 semi-leaf, naturally backed – – – – – – 1 – – – – – 1 semi-leaf, unidentifiable – – – – – – 1 – – – – – 1 sub-leaf, thinned base, backed – – – – – – 1 – – – – – 1 sub-trapezoidal, naturally back – – – 1 – – – – – – – – 1 crescent – – – – – – – – – 1 – – 1

BIFACIAL SCRAPERS 7 23 29.2 7 11 17 10 2.3 4.2 2 2.9 4.9 26 4.4 7.6 straight, reutilised – – – – – – 1 – – – – – 1 straight, distally thinned – – – – – – 1 – – – – – 1 straight, thinned base, naturally back 2 – – – – – – – – – – – 2 convex, backed 1 – – 1 – – – – – 1 – – 2 convex, thinned base – – – – – – – – – 1 – – 1 convex, reutilised 1 – – – – – 1 – – – – – 1 straight-convex, distally thinned – – – – – – 1 – – – – – 2 semi-leaf, thinned base – – – 1 – – 2 – – – – – 1

76

semi-crescent, naturally back – – – 1 – – – – – – – – 3

semi-trapezoidal – – – 1 – – – – – – – – 1

sub-leaf, distally thinned, reutilised – – – – – – 1 – – – – – 1

sub-leaf, naturally back – – – 2 – – – – – – – – 1

sub-trapezoidal 1 – – – – – – – – – – – 2

sub-crescent – – – – – – 1 – – – – – 1

crescent – – – – – – 1 – – – – – 1

convergent, unidentifiable 2 – – 1 – – 1 – – – – – 1

RETOUCHED PIECES 4 13 – 14 23 – 104 24 – 11 16 – 133 23 – blade, dorsal – – – – – – 7 – – – – – 7

blade, ventral – – – – – – 1 – – – – – 1

flake, dorsal 4 – – 13 – – 65 – – 11 – – 93

flake, ventral – – – 1 – – 21 – – – – – 22

flake, alternating – – – – – – 4 – – – – – 4

flake, alternate – – – – – – 4 – – – – – 4

unidentifiable debitage – – – – – – 2 – – – – – 2

UNIDENTIFIABLE TOOLS 3 9.7 – 6 9.8 – 87 20 – 17 25 – 113 19 – distal, dorsal 1 – – 2 – – 2 – – – – – 5

lateral, dorsal 2 – – 3 – – 50 – – 5 – – 60

lateral, alternating – – – – – – 5 – – – – – 5

lateral, ventral – – – – – – 6 – – 1 – – 7

lateral, bifacially retouched – – – – – – 1 – – – – – 1

lateral/distal, bifacial/ventral – – – – – – 1 – – – – – 1

bilateral, dorsal – – – 1 – – 13 – – 4 – – 18

bilateral, alternate – – – – – – 1 – – 1 – – 2

bifacial – – – – – – 8 – – 6 – – 14

TOTAL 31 100 100 61 100 100 428 100 100 69 100 100 589 100 100 a Percentage when unidentifiable tools are omitted from the total.

77

Table S12. Technological and typological variability of Altai Middle Palaeolithic assemblages. Values in percentages (%) a.

/

scrapers

Assemblage déjeté

oints

/

p

convex bifacial tools

-

scrapers

faced cores

vergent

-

-

convex bifacial tools

n

-

mple scrapers

at

rthogonal cores

adial coresadial etouched

o lano

nd

R O Fl PreferentialLevallois Convergent cores Si C P R toolsLevallois E Bi

Chagyrskaya Cave, sublayer 6c/1 31.3 37.5 0 0 39.4 32.2 6.8 13.9 0 0.8 0 Ust’-Karakol-1, layers 17–13 25.0 0 75.0 0 0 0 0 0 11.3 1.6 0 Ust’-Karakol-1, layer 18 0 0 28.5 71.4 4.1 0 0 0 35.1 1.0 0 Kara-Bom, MP2 0 0 58.8 35.2 1.8 0 0 0 30.6 0 0.9 Kara-Bom, MP1 0 0 0 66.7 0 0 0 4.0 17.0 4.0 0 Denisova Cave (Entrance zone), layer 10 25.0 0 50.0 25.0 15.8 1.6 0 0 22.2 0 1.6 Denisova Cave (Entrance zone), layer 9 50.0 0 50.0 0 8.3 3.3 0 0 22.5 1.5 0.8 Denisova Cave (Entrance zone), layer 8 18.2 0 54.5 27.3 10.4 4.8 0 3.2 4.0 9.6 0.8 Denisova Cave (Main Chamber), layer 22 0 0 40.0 30.0 22.5 7.5 0 0 12.5 0 0 Denisova Cave (Main Chamber), layer 21 18.2 0 9.1 0 12.5 0 0 0 0 4.1 0 Denisova Cave (Main Chamber), layer 19 23.8 9.5 33.3 14.2 15.5 1.6 0 1.1 9.1 2.1 0.8 Denisova Cave (Main Chamber), layer 14 16.6 25.0 33.3 25.0 17.7 4.9 0 1.2 12.2 3.1 0 Denisova Cave (Main Chamber), layer 12 16.6 0 61.1 11.1 14.7 2.1 0 0.7 6.7 3.2 0 Denisova Cave (East Chamber), layer 15 50.0 0 0 0 8.7 0 0 0 0 0 0 Denisova Cave (East Chamber), layer 14 45.2 0 0 5.5 13.8 4.9 0 0 0 0 0 Denisova Cave (East Chamber), layer 12 27.2 0 20.0 11.4 14.9 0.5 0 0 5.2 0 0 Denisova Cave (East Chamber), layer 11.4 19.5 0 46.3 4.8 15.2 2.2 0 1.1 6.0 1.1 0 Denisova Cave (East Chamber), layer 11.3 27.0 0 46.0 5.4 18.1 1.7 0 2.3 5.1 4.0 0 Denisova Cave (East Chamber), layer 11.2 8.0 0 48.0 8.0 19.8 1.8 0 0.6 3.1 5.0 0 Strashnaya Cave 20.0 0 20.0 20.0 5.5 0 0 5.5 0 0 0 Ust’-Kanskaya Cave 0 0 0 71.4 16.1 2.2 0 2.2 6.5 1.1 1.1 a Percentages of core types and tool types are calculated relative to the total number of cores and tools, respectively, in each assemblage.

78

Table S13. Hierarchical cluster analysis results for Altai Middle Palaeolithic assemblages, subdivided into 2–5 clusters.

Assemblage 2 clusters 3 clusters 4 clusters 5 clusters Chagyrskaya Cave, sublayer 6c/1 1 1 1 1 Ust’-Karakol-1, layers 17–13 2 2 2 2 Ust’-Karakol-1, layer 18 2 3 3 3 Kara-Bom, MP2 2 2 2 2 Kara-Bom, MP1 2 3 3 3 Denisova Cave (Entrance zone), layer 10 2 2 2 2 Denisova Cave (Entrance zone), layer 9 2 2 2 2 Denisova Cave (Entrance zone), layer 8 2 2 4 4 Denisova Cave (Main Chamber), layer 22 2 2 2 2 Denisova Cave (Main Chamber), layer 21 2 2 2 2 Denisova Cave (Main Chamber), layer 19 2 2 2 2 Denisova Cave (Main Chamber), layer 14 2 2 2 2 Denisova Cave (Main Chamber), layer 12 2 2 2 2 Denisova Cave (East Chamber), layer 15 2 2 2 2 Denisova Cave (East Chamber), layer 14 2 2 2 2 Denisova Cave (East Chamber), layer 12 2 2 2 2 Denisova Cave (East Chamber), layer 11.4 2 2 2 2 Denisova Cave (East Chamber), layer 11.3 2 2 2 2 Denisova Cave (East Chamber), layer 11.2 2 2 2 2 Strashnaya Cave 2 2 2 2 Ust’-Kanskaya Cave 2 3 3 5

Table S14. PERMANOVA test results for 2–5 clusters (Altai Middle Palaeolithic assemblages).

2 clusters 3 clusters 4 clusters 5 clusters

Number of permutations 9999 9999 9999 9999 Total sum of squares 3.54E+04 3.54E+04 3.54E+04 3.54E+04 Within-group sum of squares 2.91E+04 1.71E+04 1.64E+04 1.59E+04 F 4.103 9.639 6.524 4.893 p 0.0451 0.0002 0.0002 0.0001

79

Table S15. Difference of means for 2 clusters (1, Chagyrskaya Cave; 2, all other Altai Middle Palaeolithic assemblages).

Levallois Convergent Plano- Bi-convex Radial Orthogonal Flat-faced Simple Retouched Levallois End- Cluster Preferential / / déjeté convex bifacial cores cores cores scrapers points tools scrapers Convergent cores scrapers bifacial tools tools 1 31.25 37.50 0 0 39.4 32.2 6.8 13.9 0 0.84 0 2 21.18 1.73 37.03 18.05 11.53 1.88 0 1.03 10.25 2.50 0.33

Table S16. PERMANOVA test results for 2 clusters (1, Chagyrskaya Cave; 2, all other Altai Middle Palaeolithic assemblages).

/

oints

/ déjeté scrapers

p

convex bifacial tools

-

scrapers

faced cores

vergent

-

-

convex bifacial tools

n

-

rthogonal cores

adial coresadial i

o

nd

imple scrapers

lat

R O F PreferentialLevallois Convergent cores S C Plano Retouched toolsLevallois E B

Number of permutations 9999 9999 9999 9999 9999 9999 – 9999 9999 9999 9999 Total sum of squares 4902 1875 1.124E+04 1.088E+04 1564 958.5 – 203.5 2092 114.3 4.786 Within-group sum of squares 4769 655.7 1.016E+04 1.043E+04 836.5 87.35 – 47.35 1988 112.8 4.7 F 0.5269 35.32 2.023 0.8108 16.51 189.5 – 62.66 0.9951 0.2588 0.3465 p 0.4832 0.047 0.0922 0.2348 0.0462 0.0446 – 0.0466 0.4332 0.6177 1

80

Table S17. Technological and typological variability of Altai and Central Asian Palaeolithic assemblages. Values in percentages (%) a.

Levallois Orthogonal Flat-faced Preferential / Simple Convergent / Retouched Plano-convex Assemblage cores cores Convergent cores scrapers déjeté scrapers points bifacial tools & Levallois tools Chagyrskaya Cave, sublayer 6c/1 37.5 0 0 39.4 32.2 13.9 6.8 Ust’-Karakol-1, layer 11 9.1 72.7 3.4 0 0 1.6 0 Ust’-Karakol-1, layers 17–13 0 75.0 11.3 0 0 0 0 Ust’-Karakol-1, layer 18 0 28.5 106.5 4.1 0 0 0 Kara-Bom, layer MP2 0 58.8 65.8 1.8 0 0 0 Kara-Bom, layer MP1 0 0 83.7 0 0 4.0 0 Kara-Bom, layers UP6 and 5 0 47.8 11.4 1.6 0 4.9 0 Kara-Bom, layers UP4–1 0 50.0 0 6.7 1.0 6.7 0 Denisova Cave (Entrance zone), layer 10 0 50.0 47.2 15.8 1.6 0 0 Denisova Cave (Entrance zone), layer 9 0 50.0 22.5 8.3 3.3 0 0 Denisova Cave (Entrance zone), layer 8 0 54.5 31.3 10.4 4.8 3.2 0 Denisova Cave (Entrance zone), layer 7 0 100.0 4.7 9.3 1.2 0 0 Denisova Cave (Entrance zone), layer 6 0 0 0 8.0 1.3 2.7 0 Denisova Cave (Main Chamber), layer 22 0 40.0 42.5 22.5 7.5 0 0 Denisova Cave (Main Chamber), layer 21 0 9.1 0 12.5 0 0 0 Denisova Cave (Main Chamber), layer 19 9.5 33.3 23.3 15.5 1.6 1.1 0 Denisova Cave (Main Chamber), layer 14 25.0 33.3 37.2 17.7 4.9 1.2 0 Denisova Cave (Main Chamber), layer 12 0 61.1 17.8 14.7 2.1 0.7 0 Denisova Cave (Main Chamber), layer 11 0 66.6 2.5 11.3 0.8 0.8 0 Denisova Cave (East Chamber), layer 15 0 0 0 8.7 0 0 0 Denisova Cave (East Chamber), layer 14 0 0 5.5 13.8 4.9 0 0 Denisova Cave (East Chamber), layer 12 0 20.0 16.6 14.9 0.5 0 0 Denisova Cave (East Chamber), layer 11.4 0 46.3 10.8 15.2 2.2 1.1 0 Denisova Cave (East Chamber), layer 11.3 0 46.0 10.5 18.1 1.7 2.3 0 Denisova Cave (East Chamber), layer 11.2 0 48.0 11.1 19.8 1.8 0.6 0 Denisova Cave (East Chamber), layer 11.1 0 0 108.5 28.8 5.1 0 0 Strashnaya Cave 0 20.0 20.0 5.5 0 5.5 0 Tumechin-1 0 0 94.2 21.7 5.6 0.6 0 Tumechin-2 0 0 0 12.9 3.2 0 0 Tumechin-4 0 0 16.7 3.6 0 0 0 81

Ust’-Kanskaya Cave 0 0 77.9 16.1 2.2 2.2 0 Obi-Rakhmat, layer 21.1 0 61.2 2.5 17.1 1.9 12.4 0 Obi-Rakhmat, layer 20 0 55.6 0 18.1 0 29.3 0 Obi-Rakhmat, layer 19.5 0 35.7 9.8 19.1 0 24.5 0 Obi-Rakhmat, layer 19.4 0 60.8 28.6 20.1 0 18.8 0 Obi-Rakhmat, layer 19.3 0 74.1 6.1 21.1 0 25.1 0 Obi-Rakhmat, layer 19.2 0 72.7 20.1 22.1 0 58.8 0 Obi-Rakhmat, layer 19.1 0 50.0 22.0 23.1 0 27.7 0 Obi-Rakhmat, layer 14.1 0 62.5 0 24.1 2.3 4.7 0 Kulbulak, layer 23 0 60.0 0 25.1 0 4.2 0 a Percentages of core types and tool types are calculated relative to the total number of cores and tools, respectively, in each assemblage.

Table S18. Summary output from principal component analysis (Altai and Central Asian Palaeolithic assemblages).

Principal component Eigenvalue % variance 1 3.1738 45.3 2 1.58124 22.6 3 0.968347 13.8 4 0.580815 8.30 5 0.444861 6.36 6 0.232638 3.32 7 0.0182967 0.26

Table S19. PERMANOVA test of principal component scores for two groups of assemblages (1, Chagyrskaya Cave; 2, all other Altai and Central Asian sites).

2 groups

Number of permutations 9999 Total sum of squares 273 Within-group sum of squares 159.2 F 27.18 p 0.0267 82

Table S20. Technological and typological variability of core preparation blanks in the Chagyrskaya, Kara-Bom, Obi-Rakhmat and Kulbulak assemblages. Values in percentages (%) a.

akes

l

”

tablets

“

bordant

bordant

é

é

d

d

radial core

bordant

bordant

bordant Assemblage bordant

bordant

é /

é

bordant

é

é

é

d

d

é

d

d

thinning, overpassedthinning,

d

d

faced cores faced

points

-

bordant

é

Blades, cortical cortical Blades, lateral Blades, crested Blades, radial core Blades, thinning bifacial Blades, cortical Flakes, lateral Flakes, crested Flakes, crested Flakes, radial core Flakes, technical Flakes, d technical Flakes, bifacial thinning Flakes, bifacial Flakes, f rejuvenation platform Striking cores: from prismatic flakes rejuvenation platform Striking from flat Levallois

Chagyrskaya Cave, sublayer 6c/1 3.9 2.3 1.8 0.3 0.3 24.5 18.3 5.5 0.3 22.7 0.3 11.0 7.6 1.3 0 0 0 Kara-Bom, layer MP2 0 19.2 1.9 0 0 0 9.6 0 0 0 0 0 0 0 0 3.8 65.4 Kara-Bom, layer MP1 0 16.7 16.7 0 0 0 0 0 0 0 0 0 0 0 0 0 66.7 Obi-Rakhmat, layer 21.1 0 20.0 36.0 0 0 0 0 0 0 0 0 0 0 0 30.0 2.0 12.0 Obi-Rakhmat, layer 20 0 0 81.0 0 0 0 0 0 0 0 0 0 0 0 0 19.0 0 Obi-Rakhmat, layer 19.5 0 34.1 22.7 0 0 0 0 0 0 0 0 0 0 0 6.8 9.1 27.3 Obi-Rakhmat, layer 19.4 0 40.7 0 0 0 0 0 0 0 0 0 0 0 0 7.4 0 51.9 Obi-Rakhmat, layer 19.3 0 76.0 4.0 0 0 0 0 0 0 0 0 0 0 0 0 4.0 16.0 Obi-Rakhmat, layer 19.2 0 75.0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 25.0 Obi-Rakhmat, layer 19.1 0 36.4 0 0 0 0 0 0 0 0 0 0 0 0 0 9.1 54.5 Obi-Rakhmat, layer 14.1 0 92.8 2.4 0 0 0 0 0 0 0 0 0 0 0 1.2 3.6 0 Kulbulak, layer 23 0 45.5 9.1 0 0 0 30.3 0 0 0 0 0 0 0 3.0 12.1 0 a Percentages of core preparation blanks are calculated relative to the total number of core preparation blanks in each assemblage.

83

Table S21. Technological and typological variability of Chagyrskaya Cave and European Micoquian assemblages. Values in percentages (%) a.

"

Chokurcha

"

triangular crescent

Assemblage

crescent

like like

-

crescent

crescent

convex method

-

rapers, transverse rapers, diagonal rapers, leaf / ovoid rapers, rapers, core

ifacial leaf points, ifacial triangular points, ifacial trapezoidal points, ifacial points, ifacial scrapers, simple ifacial scrapers, double ifacial scrapers, leaf ifacial scrapers, ifacial scrapers, trapezoidal ifacial scrapers, ifacial

oints, simple oints, leaf oints, triangular oints, trapezoidal oints, oints, lano

crapers, simple crapers, double crapers, triangular crapers, rectangular/ trapezoidal crapers,

P P P P P Sc Sc S S Sc S S Sc Sc B B B B B B B B B B Keilmesser P

Chagyrskaya Cave, sublayer 6c/1 2.2 6.9 1.3 2.2 1.3 4.3 9.1 23.4 2.6 6.5 1.3 19.9 4.3 0 0.9 0 0 0 1.3 0.4 1.7 0 0 0.4 1.7 93.8 Kabazi V, subunits II/4A–II/7 0 3.9 0 3.9 9.6 7.7 0 34.6 7.7 0 0 7.7 7.7 0 1.9 1.9 0 0 0 0 1.9 0 0 0 0 66.7 Kabazi V, subunit III/1 0.5 5.2 2.6 3.1 2.1 2.6 2.6 21.8 7.3 4.2 0 14.5 8.3 0 3.1 1.0 0 2.6 0.5 0 3.6 0.5 0.5 5.7 0.5 46.3 Kabazi V, subunit III/1А 0.5 2.3 2.8 1.4 2.3 3.7 3.2 20.7 6.5 4.6 3.7 10.1 12.0 0 2.3 0 0.5 0 0 0 1.8 0 0 6.9 3.2 58.1 Kabazi V, subunit III/2 1.5 3.8 1.5 0.8 5.3 6.8 3.0 23.5 5.3 10.6 0 8.3 4.6 0 1.5 0 0 0.8 0.8 0 5.3 0 0 3.8 1.5 61.1 Kabazi V, subunit III/5 0 1.4 0.9 3.3 2.3 8.8 4.7 34.9 6.1 0.9 1.9 11.2 4.7 0 1.4 0.5 0 0.9 0.5 0 0.5 0 0.5 0.5 0.9 58.3 Karabai I, layer 4 3.2 1.6 1.6 0 0 4.8 6.4 28.6 7.9 1.6 1.6 14.3 0 1.6 4.8 1.6 0 0 3.2 1.6 0 3.2 0 0 9.5 66.7 Kabazi II, units IIA–III 1.6 1.6 0 0 0 12.5 0 29.7 15.6 0 4.7 3.1 0 0 4.7 0 0 0 0 0 3.1 0 0 4.7 3.1 62.5 Kabazi II, units V and VI 0 1.8 0 0 3.5 3.5 0 15.8 8.8 3.5 1.8 10.5 1.8 3.5 7.0 0 0 0 1.8 1.8 7.0 0 5.3 1.8 7.0 64.7 Kiik-Koba, level IV 0 4.0 6.6 19.1 5.1 5.5 2.9 17.6 4.0 2.2 2.9 11.4 2.2 0 1.5 0.7 2.2 3.3 0 0 0.4 0.7 0.7 2.9 1.1 54.1 Buran Kaya III, layer B 0.4 6.2 4.4 9.9 5.1 9.9 2.2 18.3 5.9 2.2 3.3 13.6 3.7 0 2.2 0 1.1 0.7 0.7 0.4 0.4 0 0.7 0.7 0.4 85.0 Starosele, level 1 2.7 0 9.1 0 2.7 9.1 2.7 20.0 9.1 1.8 2.7 11.8 4.6 0 0 0 0 0 0 0 6.4 0 0 0.9 0 87.5 Chokurcha I, unit IV 0 0.6 2.9 2.9 0 13.3 4.6 23.7 8.1 1.2 2.3 11.6 2.9 2.3 1.7 0.6 0 1.2 1.7 0 2.3 1.2 0 5.2 6.9 91.7 Zaskalnaya V, unit I 8.1 3.2 4.8 0 0 1.6 4.8 16.1 0 3.2 3.2 19.4 9.7 0 0 1.6 0 1.6 0 0 4.8 0 1.6 0 1.6 71.4 Zaskalnaya V, unit II 2.1 0 0 0 0 2.1 6.3 20.8 8.3 8.3 2.1 14.6 2.1 0 4.2 0 0 0 0 0 6.3 0 4.2 6.3 4.2 58.3 Zaskalnaya V, unit IIа 3.2 0 3.2 0 1.6 6.4 11.1 22.2 3.2 4.8 1.6 19.1 0 0 1.6 0 0 0 1.6 0 4.8 1.6 4.8 1.6 4.8 76.9 Zaskalnaya V, units III/1–III/9-1 1.2 1.2 0 1.2 0 6.0 4.8 27.7 4.8 6.0 1.2 4.8 2.4 0 2.4 1.2 0 1.2 0 0 6.0 3.6 2.4 3.6 13.3 53.9 Zaskalnaya V, units III/10–III/14 4.4 7.1 1.8 1.8 0 5.3 5.3 25.7 2.7 1.8 0.9 12.4 2.7 0 6.2 0.9 0.9 1.8 0 0 6.2 0 0.9 2.7 7.1 58.0 Zaskalnaya V, unit IIIA 2.6 8.8 1.6 0 2.1 3.6 7.3 22.8 2.6 4.7 2.1 9.3 4.7 0 4.7 1.0 0 0 0 0 6.2 0.5 1.0 4.2 2.6 59.0

84

Zaskalnaya V, unit IV 0 0 5.3 2.3 0.8 6.1 4.6 14.5 6.9 5.3 6.9 22.9 0.8 0 0.8 3.1 0 0 0.8 0.8 1.5 1.5 0.8 3.1 3.1 55.0 Sesselfelsgrotte, unit G4 0.3 1.9 1.0 0.6 0.6 4.4 3.8 21.1 3.2 7.9 1.0 18.3 0.6 0.3 0.3 0 0 0.3 1.9 0.3 2.8 0 1.3 1.0 3.5 66.3 Sesselfelsgrotte, unit G3 0.7 1.8 0.4 0 1.1 2.2 5.1 22.6 3.7 7.7 2.2 18.3 1.1 0 1.1 0 0 0.4 0 0 2.6 0.4 0.7 2.2 5.8 76.1 Sesselfelsgrotte, unit G2 0.9 2.3 0.9 0 0.5 2.8 7.4 27.4 0.9 6.1 1.9 15.4 1.9 0 1.9 0 0 0 0 0.5 5.1 0.9 1.9 1.9 7.4 72.4 Antonovka I 0 0.5 0 0.2 0.2 3.6 0 20.7 1.9 9.6 8.9 2.2 8.0 0 0.7 0 0 0 0 0 3.9 0.2 0 3.1 0.5 100.0 Antonovka II 0 0.2 0.8 0.8 0.6 3.4 0 33.9 5.1 5.1 8.4 4.0 4.8 0 0.4 0.2 0 0 0 0 2.5 1.7 0 1.9 0.4 100.0 Barakaevskaya Cave 1.6 1.3 3.2 0.5 0.8 3.2 0.8 35.8 6.4 6.1 1.3 6.9 1.3 0 0 0.8 0 0 0.3 0 0.3 0 0 0 0.5 100.0 a Percentages of core types and tool types are calculated relative to the total number of cores and tools, respectively, in each assemblage.

85

Table S22. Hierarchical cluster analysis results for Chagyrskaya Cave and European Micoquian assemblages, subdivided into 2–5 clusters.

Assemblage 2 clusters 3 clusters 4 clusters 5 clusters Chagyrskaya Cave, sublayer 6с/1 1 1 1 1 Kabazi V, subunits II/4A–II/7 1 1 1 1 Kabazi V, subunit III/1 1 1 1 1 Kabazi V, subunit III/1А 1 1 1 1 Kabazi V, subunit III/2 1 1 1 1 Kabazi V, subunit III/5 1 1 1 1 Karabai I, layer 4 1 2 2 2 Kabazi II, units IIA–III 1 1 1 1 Kabazi II, units V and VI 2 3 3 3 Kiik-Koba, level IV 1 1 4 4 Buran Kaya III, layer B 1 1 1 1 Starosele, level 1 1 1 1 1 Chokurcha I, unit IV 1 1 1 1 Zaskalnaya V, unit I 1 1 1 1 Zaskalnaya V, unit II 1 1 1 1 Zaskalnaya V, unit IIа 1 1 1 1 Zaskalnaya V, levels III/1–III/9-1 1 1 1 1 Zaskalnaya V, levels III/10–III/14 1 1 1 1 Zaskalnaya V, unit IIIA 1 1 1 1 Zaskalnaya V, unit IV 1 1 1 5 Sesselfelsgrotte, unit G4 1 1 1 1 Sesselfelsgrotte, unit G3 1 1 1 1 Sesselfelsgrotte, unit G2 1 1 1 1 Antonovka I 1 1 1 1 Antonovka II 1 1 1 1 Barakaevskaya Cave 1 1 1 1

Table S23. PERMANOVA test results for 2–5 clusters (Chagyrskaya Cave and European Micoquian assemblages).

2 clusters 3 clusters 4 clusters 5 clusters

Number of permutations 9999 9999 9999 9999 Total sum of squares 1.12E+04 1.12E+04 1.12E+04 1.12E+04 Within-group sum of squares 1.10E+04 1.08E+04 1.00E+04 9442 F 0.5495 0.4596 0.8438 0.9752 p 0.7357 0.8914 0.5826 0.5003

86

Table S24. Difference of means for 2 clusters (1, Chagyrskaya Cave and European Micoquian; 2, Kabazi II, units V and VI).

"

Chokurcha

"

crescent

Cluster

crescent

like like

-

simple

crescent

convex method

-

rapers, transverse rapers, diagonal rapers, leaf / ovoid rapers, crescent rapers, core

ifacial leaf points, ifacial triangular points, ifacial trapezoidal points, ifacial points, ifacial scrapers, simple ifacial scrapers, double ifacial scrapers, leaf ifacial scrapers, triangular ifacial scrapers, trapezoidal ifacial scrapers, ifacial

oints, simple oints, leaf oints, triangular oints, trapezoidal oints, oints, lano

crapers, crapers, double crapers, triangular crapers, rectangular/ trapezoidal crapers,

P P P P P Sc Sc S S Sc S S Sc Sc B B B B B B B B B B Keilmesser P

1 1.50 2.63 2.26 2.15 1.77 5.59 4.11 24.33 5.42 4.49 2.64 12.19 3.79 0.17 2.01 0.60 0.19 0.59 0.53 0.16 3.22 0.64 0.88 2.52 3.35 71.15 2 0 1.75 0 0 3.51 3.51 0 15.79 8.77 3.51 1.75 10.53 1.75 3.51 7.02 0 0 0 1.75 1.75 7.02 0 5.26 1.75 7.02 64.71

Table S25. PERMANOVA test results for 2 clusters (1, Chagyrskaya Cave and European Micoquian; 2, Kabazi II, units V and VI).

"

Chokurcha

crescent

crescent

" like

-

scrapers, double

crescent

convex method

-

rapers, transverse rapers, diagonal rapers, leaf / ovoid rapers, crescent rapers, core

ifacial leaf points, ifacial triangular points, ifacial trapezoidal points, ifacial points, ifacial scrapers, simple ifacial ifacial scrapers, leaf ifacial scrapers, triangular ifacial scrapers, trapezoidal ifacial scrapers, ifacial

oints, simple oints, leaf oints, triangular oints, trapezoidal oints, oints, lano

crapers, simple crapers, double crapers, triangular crapers, rectangular/ trapezoidal crapers,

P P P P P Sc Sc S S Sc S S Sc Sc B B B B B B B B B B Keilmesser P

Number of 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 9999 permutations Total sum of squares 84.94 152.6 132.4 413.5 131.8 241.7 215.3 912.2 264.3 212.6 133.8 761.5 252.3 17.91 94.05 15.6 6.26 20.1 17.82 5.806 125.5 24.49 58.14 106.4 286.8 6517 Within-group sum of 82.76 151.9 127.5 409 128.9 237.5 199 842.3 253.4 211.7 133.1 758.7 248.4 7.234 70.13 15.25 6.226 19.76 16.27 3.22 111.7 24.1 39.36 105.9 273.9 6477 squares F 0.634 0.105 0.93 0.263 0.539 0.423 1.957 1.991 1.033 0.108 0.123 0.088 0.373 35.41 8.187 0.552 0.131 0.409 2.28 19.28 2.958 0.392 11.45 0.116 1.126 0.148 p 0.35 0.848 0.305 0.202 0.459 0.539 0.223 0.228 0.228 0.774 0.613 0.766 0.499 0.039 0.037 0.382 1 0.463 0.116 0.043 0.042 0.542 0.0414 0.724 0.188 0.726

87

Table S26. Technological and typological variability of Chagyrskaya Cave and European Micoquian assemblages (merged set of variables). Values in percentages (%) a.

Scrapers Points, leaf- Bifacial points, Bifacial Bifacial Bifacial transverse, shaped, Points, Points, leaf-shaped, points, points, scrapers, Plano- diagonal, crescent & triangular & trapezoidal crescent & triangular & trapezoidal & simple & Assemblage convex Keilmesser simple, double scrapers, leaf- scrapers, & scrapers, bifacial scrapers, bifacial bifacial bifacial method & points, shaped, ovoid, triangular trapezoidal leaf-shaped, scrapers, scrapers, scrapers, simple crescent crescent triangular trapezoidal double

Chagyrskaya Cave, sublayer 6c/1 93.8 41.6 19.0 2.6 22.1 3.0 0 0 1.7 1.7 Kabazi V, subunits II/4A–II/7 66.7 50.0 21.2 0 11.5 3.8 1.9 0 0 0 Kabazi V, subunit III/1 46.3 34.7 19.7 2.6 17.6 15.0 1.6 0.5 0.5 0.5 Kabazi V, subunit III/1А 58.1 34.6 21.2 6.5 11.5 11.1 0 0.5 0 3.2 Kabazi V, subunit III/2 61.1 40.2 24.2 1.5 9.1 11.4 0 0 0.8 1.5 Kabazi V, subunit III/5 58.3 54.4 9.3 2.8 14.4 3.3 0.5 0.5 0.5 0.9 Karabai I, layer 4 66.7 50.8 3.2 3.2 14.3 4.8 4.8 0 4.8 9.5 Kabazi II, units IIA–III 62.5 59.4 1.6 4.7 3.1 12.5 0 0 0 3.1 Kabazi II, units V and VI 64.7 28.1 10.5 1.8 10.5 15.8 0 5.3 3.5 7.0 Kiik-Koba, level IV 54.1 30.0 13.6 9.5 30.4 8.1 1.5 2.9 0 1.1 Buran Kaya III, layer B 85.0 36.6 17.2 7.7 23.4 4.0 0 1.8 1.1 0.4 Starosele, level 1 87.5 43.6 9.1 11.8 11.8 7.3 0 0 0 0 Chokurcha I, unit IV 91.7 49.7 4.6 5.2 14.5 10.4 1.7 0 1.7 6.9 Zaskalnaya V, unit I 71.4 30.6 16.1 8.1 19.4 6.5 1.6 1.6 0 1.6 Zaskalnaya V, unit II 58.3 39.6 10.4 2.1 14.6 16.7 0 4.2 0 4.2 Zaskalnaya V, unit IIа 76.9 46.0 6.3 4.8 19.0 7.9 1.6 4.8 1.6 4.8 Zaskalnaya V, units III/1–III/9-1 53.9 44.6 9.6 1.2 6.0 13.3 4.8 2.4 0 13.3 Zaskalnaya V, units III/10–III/14 58.0 43.4 11.5 2.7 14.2 16.8 0.9 1.8 0 7.1 Zaskalnaya V, unit IIIA 59.0 38.9 20.2 3.6 9.3 15.0 1.6 1.0 0 2.6 Zaskalnaya V, unit IV 55.0 32.1 6.9 12.2 25.2 5.3 4.6 0.8 1.5 3.1 Sesselfelsgrotte, unit G4 66.3 32.8 11.0 1.9 18.9 4.4 0 1.3 2.2 3.5 Sesselfelsgrotte, unit G3 76.1 34.3 11.7 2.6 18.2 6.2 0.4 0.7 0 5.8 Sesselfelsgrotte, unit G2 72.4 39.5 10.7 2.8 15.3 8.8 0.9 1.9 0.5 7.4 Antonovka I 100.0 26.3 18.3 8.9 2.4 7.7 0.2 0 0 0.5 Antonovka II 100.0 42.3 10.7 9.3 4.8 4.8 1.9 0 0 0.4 Barakaevskaya Cave 100.0 47.7 9.5 4.5 7.4 0.3 0.8 0 0.3 0.5 a Percentages of core types and tool types are calculated relative to the total number of cores and tools, respectively, in each assemblage. 88

Table S27. Summary output from principal component analysis (Chagyrskaya Cave and European Micoquian assemblages).

Principal component Eigenvalue % variance 1 273.671 58.6 2 90.8009 19.5 3 46.5159 9.97 4 28.5615 6.12 5 12.2732 2.63 6 8.49675 1.83 7 3.69853 0.79 8 1.08487 0.23 9 0.963017 0.21 10 0.616972 0.13

Table S28. PERMANOVA test of principal component scores for two groups of assemblages (1, Chagyrskaya Cave; 2, European Micoquian).

2 groups

Number of permutations 9999 Total sum of squares 1.167E+04 Within-group sum of squares 1.097E+04 F 1.524 p 0.2698

89

Table S29. Summary output from principal component analysis resulting from geometric- morphometric shape analysis of Chagyrskaya Cave and Sesselfelsgrotte bifacial tools.

Principal component Eigenvalue % variance Principal component Eigenvalue % variance 1 6656.2737 36.25 23 75.0681 0.41 2 3073.4361 16.74 24 63.9403 0.35 3 1554.3843 8.47 25 56.9992 0.31 4 1078.2323 5.87 26 56.0011 0.31 5 874.6680 4.76 27 51.8302 0.28 6 672.8161 3.66 28 46.9504 0.26 7 492.8403 2.68 29 45.5894 0.25 8 466.3421 2.54 30 42.8193 0.23 9 411.2240 2.24 31 36.0344 0.20 10 375.5355 2.05 32 34.8675 0.19 11 332.6369 1.81 33 31.7330 0.17 12 240.0253 1.31 34 29.8207 0.16 13 209.2771 1.14 35 27.3043 0.15 14 190.6298 1.04 36 25.5218 0.14 15 164.3116 0.89 37 22.8619 0.12 16 150.4231 0.82 38 21.6744 0.12 17 145.2950 0.79 39 20.9382 0.11 18 129.8246 0.71 40 17.6239 0.10 19 106.2122 0.58 41 17.5411 0.10 20 99.1700 0.54 42 15.1771 0.08 21 88.3743 0.48 43 14.0369 0.08 22 82.0862 0.45 44 11.8820 0.06

Table S30. PERMANOVA test of principal component scores for bifacial tools from Chagyrskaya Cave and Sesselfelsgrotte.

2 groups

Number of permutations 9999 Total sum of squares 8.084E+05 Within-group sum of squares 7.76E+05 F 1.792 p 0.0908

90

Table S31. Technological and typological variability of assemblages from Chagyrskaya Cave, other Altai sites (Denisova Cave, Kara-Bom, Ust’-Karakol-1), Obi-Rakhmat in Central Asia, and Micoquian sites in eastern Europe (, Donbass-Azoz, Caucasus) and central Europe. Values in percentages (%) a.

Levallois Simple Convergent / Retouched Plano-convex Assemblage tools scrapers déjeté scrapers points bifacial tools

Chagyrskaya Cave, sublayer 6c/1 0 39.4 32.2 13.9 6.8 Ust’-Karakol-1, layer 11 3.4 0 0 1.6 0 Ust’-Karakol-1, layers 17–13 11.3 0 0 0 0 Ust’-Karakol-1, layer 18 35.1 4.1 0 0 0 Kara-Bom, layer MP2 30.6 1.8 0 0 0 Kara-Bom, layer MP1 17.0 0 0 4.0 0 Kara-Bom, layers UP6 and 5 7.1 1.6 0 4.9 0 Kara-Bom, layers UP4–1 0 6.7 1.0 6.7 0 Denisova Cave (Entrance zone), layer 10 22.2 15.8 1.6 0 0 Denisova Cave (Entrance zone), layer 9 22.5 8.3 3.3 0 0 Denisova Cave (Entrance zone), layer 8 4.0 10.4 4.8 3.2 0 Denisova Cave (Entrance zone), layer 7 4.7 9.3 1.2 0 0 Denisova Cave (Entrance zone), layer 6 0 8.0 1.3 2.7 0 Denisova Cave (Main Chamber), layer 22 12.5 22.5 7.5 0 0 Denisova Cave (Main Chamber), layer 21 0 12.5 0 0 0 Denisova Cave (Main Chamber), layer 19 9.1 15.5 1.6 1.1 0 Denisova Cave (Main Chamber), layer 14 12.2 17.7 4.9 1.2 0 Denisova Cave (Main Chamber), layer 12 6.7 14.7 2.1 0.7 0 Denisova Cave (Main Chamber), layer 11 2.5 11.3 0.8 0.8 0 Denisova Cave (East Chamber), layer 15 0 8.7 0 0 0 Denisova Cave (East Chamber), layer 14 0 13.8 4.9 0 0 Denisova Cave (East Chamber), layer 12 5.2 14.9 0.5 0 0 Denisova Cave (East Chamber), layer 11.4 6.0 15.2 2.2 1.1 0 Denisova Cave (East Chamber), layer 11.3 5.1 18.1 1.7 2.3 0 Denisova Cave (East Chamber), layer 11.2 3.1 19.8 1.8 0.6 0 Denisova Cave (East Chamber), layer 11.1 8.5 28.8 5.1 0 0 Strashnaya Cave 0 5.5 0 5.5 0 Tumechin-1 24.2 21.7 5.6 0.6 0 Tumechin-2 0 12.9 3.2 0 0 Tumechin-4 16.7 3.6 0 0 0 Ust’-Kanskaya Cave 6.5 16.1 2.2 2.2 0 Obi-Rakhmat, layer 21.1 2.5 17.1 1.9 12.4 0 Obi-Rakhmat, layer 20 0 18.1 0 29.3 0 Obi-Rakhmat, layer 19.5 9.8 19.1 0 24.5 0 Obi-Rakhmat, layer 19.4 11.3 20.1 0 18.8 0 Obi-Rakhmat, layer 19.3 2.4 21.1 0 25.1 0 Obi-Rakhmat, layer 19.2 2.0 22.1 0 58.8 0 Obi-Rakhmat, layer 19.1 8.4 23.1 0 27.7 0

91

Obi-Rakhmat, layer 14.1 0 24.1 2.3 4.7 0 Kulbulak, layer 23 0 25.1 0 4.2 0 Kabazi V, subunits II/4A–II/7 0 50.0 15.4 17.4 5.7 Kabazi V, subunit III/1 0 34.3 27.0 13.5 18.0 Kabazi V, subunit III/1А 0 34.1 30.4 9.3 14.7 Kabazi V, subunit III/2 0 38.6 23.5 12.9 13.7 Kabazi V, subunit III/5 0 54.5 18.7 7.9 5.7 Karabai I, layer 4 0 47.7 19.1 6.4 23.9 Kabazi II, units IIA–III 0 57.8 7.8 3.2 15.6 Kabazi II, units V and VI 0 28.1 21.1 5.3 31.7 Kiik-Koba, level IV 0 30.0 18.7 34.8 13.5 Buran Kaya III, layer B 0 36.3 22.8 26.0 7.3 Starosele, level 1 0 40.9 20.9 14.5 7.3 Chokurcha I, unit IV 0 49.7 20.3 6.4 20.8 Zaskalnaya V, unit I 0 22.5 35.5 16.1 11.2 Zaskalnaya V, unit II 0 37.5 27.1 2.1 25.2 Zaskalnaya V, unit IIа 0 42.9 25.5 8.0 20.8 Zaskalnaya V, units III/1–III/9-1 0 43.3 14.4 3.6 33.7 Zaskalnaya V, units III/10–III/14 0 39.0 17.8 15.1 26.7 Zaskalnaya V, unit IIIA 0 36.3 20.8 15.1 20.2 Zaskalnaya V, unit IV 0 32.1 35.9 8.4 15.5 Sesselfelsgrotte, unit G4 0 32.5 28.1 4.4 11.4 Sesselfelsgrotte, unit G3 0 33.6 29.3 4.0 13.2 Sesselfelsgrotte, unit G2 0 38.5 25.3 4.6 19.6 Antonovka I 0 26.2 28.7 0.9 8.4 Antonovka II 0 42.4 22.3 2.4 7.1 Barakaevskaya Cave 0 46.2 15.6 7.4 1.9 Mezmaiskaya Cave, layer 2B-4 0 45.8 18.5 8.8 10.6 Mezmaiskaya Cave, layer 3 0 41.2 13.0 7.1 12.6 a Percentages of core types and tool types are calculated relative to the total number of tools, respectively, in each assemblage.

92

Table S32. Summary output from principal component analysis (Chagyrskaya Cave, Denisova Cave and Kara-Bom variants (Altai), Obirakhmatian (Central Asia), and Micoquian sites in eastern and central Europe, including Mezmaiskaya Cave).

Principal component Eigenvalue % variance 1 2.83563 56.7 2 1.00378 20.1 3 0.597631 12.0 4 0.299524 5.99 5 0.263439 5.27

Table S33. PERMANOVA test of principal component scores for two groups of assemblages (1, European Micoquian, Mezmaiskaya Cave and Chagyrskaya Cave; 2, Altai and Central Asian Middle and Upper Palaeolithic).

2 groups

Number of permutations 9999 Total sum of squares 330 Within-group sum of squares 163.1 F 66.5 p 0.0001

93

References

Aitken, M. J., 1998. An Introduction to Optical Dating: the dating of Quaternary sediments by the use of photon-stimulated luminescence (Oxford University Press, Oxford). Anderson, M. J., 2001. A new method for non-parametric multivariate analysis of variance. Austral Ecology 26, 32–46. Archer, W., Gunz, P., van Niekerk, K. L., Henshilwood, C. S. & McPherron, S. P., 2015. Diachronic change within the Still Bay at , . PLoS ONE 10, e0132428. Bailey, S. E., 2002. A closer look at Neanderthal postcanine dental morphology: the mandibular dentition. Anatomical Record 269, 148–156. Belmaker, M., 2017. The Southern Levant during the Last Glacial and zooarchaeological evidence for the effects of climate-forcing on hominin population dynamics, in Climate Change and Human Responses: a zooarchaeological perspective (ed. Monks, G.G.) 7–25 (Springer, Dordrecht). Besenbacher, S., Hvilsom, C., Marques-Bonet, T., Mailund, T. & Schierup, M. H., 2019. Direct estimation of mutations in great apes reconciles phylogenetic dating. Nature Ecology and Evolution 3, 286–292. Binford, L. R. & Binford, S. R., 1966. A preliminary analysis of functional variability in the Mousterian of Levallois facies. American Anthropologist 68, 238–295. Blegen, N., Tryon, C. A., Faith, J. T., Peppe, D. J., Beverly, E. J., Li, B. & Jacobs, Z., 2015. Distal tephras of the eastern Lake Victoria basin, equatorial East Africa: correlations, chronology and a context for early modern humans. Quaternary Science Reviews 122, 89– 111. Bokelmann, L., Hajdinjak, M., Peyrégne, S., Brace, S., Essel, E., de Filippo, C., Glocke, I., Grote, S., Mafessoni, F., Nagel, S., Kelso, J., Prüfer, K., Vernot, B., Barnes, I., Pääbo, S., Meyer, M. & Stringer, C., 2019. A genetic analysis of the Neanderthals. Proceedings of the National Academy of Sciences of the U.S.A. 116, 15610–15615. Bordes, F. H., 1966. cultures in southwest , in Studies in Prehistory: Robert Bruce Foote memorial volume (eds Sen, D. & Ghosh, A. K.) 49–57 (Mukhopadhyay, Calcutta). Bosinski, G., 1967. Die Mittelpaläolithischen Funde im Westlichen Mitteleuropa (Böhlau- Verlag, Köln). Bøtter-Jensen, L. & Mejdahl, V., 1988. Assessment of beta dose-rate using a GM multicounter system. Nuclear Tracks and Radiation Measurements 14, 187–191. Bøtter-Jensen, L., Andersen, C. E., Duller, G. A. T. & Murray, A. S., 2003. Developments in radiation, stimulation and observation facilities in luminescence measurements. Radiation Measurements 37, 535–541. Briggs, A. W., Good, J. M., Green, R. E., Krause, J., Maricic, T., Stenzel, U., Lalueza-Fox, C., Rudan, P., Brajković, D., Kućan, Ž., Gušic, I., Schmitz, R., Doronichev, V. B., Golovanova, L. V., de la Rasilla, M., Fortea, J., Rosas, A. & Pääbo, S., 2009. Targeted retrieval and analysis of five Neandertal mtDNA genomes. Science 325, 318–321. Brock, F., Geoghegan, V., Thomas, B., Jurkschat, K. & Higham, T. F. G., 2013. Analysis of bone “collagen” extraction products for radiocarbon dating. Radiocarbon 55, 445–463. Bronk Ramsey, C., 2009. Bayesian analysis of radiocarbon dates. Radiocarbon 51, 337–360. Brown, S., Higham, T., Slon, V., Pääbo, S., Meyer, M., Douka, K., Brock, F., Comeskey, D., Procopio, N., Shunkov, M., Derevianko, A. & Buckley, M., 2016. Identification of a new hominin bone from Denisova Cave, Siberia using collagen fingerprinting and mitochondrial DNA analysis. Scientific Reports 6, 23559. Brown, T. A., Nelson, D. E., Vogel, J. S. & Southon, J. R., 1988. Improved collagen extraction by modified Longin method. Radiocarbon 30, 171–177.

94

Browning, S. R., Browning, B. L., Zhou, Y., Tucci, S. & Akey, J. M., 2018. Analysis of human sequence data reveals two pulses of archaic Denisovan admixture. Cell 173, 53–61. Brühl, E. & Laurat, T., 2010. The Middle Palaeolithic at the Geisel Valley – recent excavations at the fossil lake Neumark-Nord 2 (Sachen-Anhalt, Germany). Acta Universitatis Wratislaviensis 3207, 2–36. Carrión, J. S., Scott, L., Arribas, A., Fuentes, N., Gil‐Romera, G. & Montoya, E., 2007. Pleistocene landscapes in central Iberia inferred from pollen analysis of hyena coprolites. Journal of Quaternary Science 22, 191–202. Chabai, V. P., 2004. The Middle of Crimea [in Russian] (Simferopol, Ukraine). Chabai, V. P., 2005. Kabazi II, Units V and VI: artefacts, in Palaeolithic Sites of Crimea, Volume 1, Kabazi II: Last Interglacial Occupation, Environment & Subsistence (eds Chabai, V., Richter, J. & Uthmeier, T.) 99–132 (Simferopol–Cologne, Shlyakh). Chabai, V. P. & Demidenko, Y. E., 1998. The classification of flint artifacts, in The Paleolithic of Crimea, Volume 1. The of Western Crimea (eds Marks, A. E. & Chabai, V. P.) 31–51 (Études et Recherches Archéologiques de l’Université de Liège 84, Liège). Chabai, V. P. & Uthmeier, T., 2017. New excavations at the Middle Paleolithic site Zaskalnaya V, Crimea. The 2012 and 2013 field seasons: a preliminary report. Quartär 64, 27–71. Chabai, V. P., Richter, J. & Uthmeier, T. (eds), 2006. Palaeolithic Sites of Crimea, Volume 2, Kabazi II: the 70,000 years since the Last Interglacial (Simferopol–Cologne, Shlyakh). Cremaschi, M. & Van Vliet-Lanoë, B., 1990. Traces of frost activity and ice segregation in Pleistocene loess deposits and till of northern : deep seasonal freezing or permafrost? Quaternary International 5, 39–48. Dalén, L., Orlando, L., Shapiro, B., Brandström-Durling, M., Quam, R., Gilbert, M. T. P., Díez Fernández-Lomana, J. C., Willerslev, E., Arsuaga, J. L. & Götherström, A., 2012. Partial genetic turnover in Neandertals: continuity in the east and population replacement in the west. Molecular Biology and Evolution 29, 1893–1897 Delagnes, A. & Rendu, W., 2011. Shifts in Neandertal mobility, technology and subsistence strategies in western France. Journal of Archaeological Science 38, 1771–1783. Derevianko, A. P. & Markin, S. V., 1992. Mousterian of Gorny Altai [in Russian] (Nauka, Novosibirsk). Derevianko, A. P. & Shunkov, M. V., 2002. Middle Paleolithic industries with foliate bifaces in Gorny Altai. Archaeology, Ethnology and Anthropology of Eurasia 9(1), 16–42. Derevianko, A. P., Petrin, V. T., Rybin, E. P. & Chevalkov, L. M., 1998a. Paleolithic Complexes of the Stratified Part of the Kara-Bom Site [in Russian] (Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Derevianko, A. P., Glinskiy, S. V., Dergacheva, M. I., Dupal, T. A., Efremov, S. A., Zenin, A. N., Krivoshapkin, A. I., Kulikov, O. A., Malaeva, E. M., Markin, S. V., Nikolaev, S. V., Nohrina, T. I., Petrin, V. T., Pozdnyakov, A. A., Popova, S. M., Rybin, E. P., Simonov, Y. G., Fedeneva, I. N., Chevalkov, L. M. & Shunkov, M. V., 1998b. Problems of Paleoecology, Geology and Archaeology of Paleolithic of Altay [in Russian] (Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Derevianko, A. P., Shunkov, M. V., Agajanian, A. K., Baryshnikov, G. F., Malaeva, E. M., Uliyanov, V. A., Kulik, N. A., Postnov, A. V. & Anoikin, A. A., 2003. Paleoenviroment and Paleolithic Human Occupation of Gorny Altai (Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Derevianko, A. P., Markin, S. V., Zykin, V. S., Zykina, V. S., Zazhigin, V. S., Sizikova, A. O., Solotchina, E. P., Smolyaninova, L. G. & Antipov, A. S., 2013a. Chagyrskaya Cave: a

95

Middle Paleolithic site in the Altai. Archaeology, Ethnology and Anthropology of Eurasia 41(1), 2–27. Derevianko, A. P., Markin, S. V. & Shunkov, M. V., 2013b. The Sibiryachikha facies of the Middle Paleolithic of the Altai. Archaeology, Ethnology and Anthropology of Eurasia 41(1), 89–103. Derevianko, A. P., Markin, S. V., Kulik, N. A. & Kolobova, K. A., 2015. Lithic raw material exploitation in the Sibiryachikha facies, the Middle Paleolithic of Altai. Archaeology, Ethnology and Anthropology of Eurasia 43(3), 3–16. Derevianko, A. P., Markin, S. V., Kolobova, K. A., Chabai, V. P., Rudaya, N. A., Viola, B., Buzhilova, A. P., Mednikova, M. B., Vasiliev, S. K., Zykin, V. S., Zykina, V. S., Zazhigin, V. S., Volvakh, A. O., Roberts, R. G., Jacobs, Z. & Li, B., 2018. Multidisciplinary Studies of Chagyrskaya Cave – A Middle Paleolithic site in Altai [in Russian] (Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Devièse, T., Karavanić, I., Comeskey, D., Kubiak, C., Korlević, P., Hajdinjak, M., Radović, S., Procopio, N., Buckley, M., Pääbo, S. & Higham, T., 2017. Direct dating of Neanderthal remains from the site of Vindija Cave and implications for the Middle to transition. Proceedings of the National Academy of Sciences of the U.S.A. 114, 10606– 10611. Dolukhanov, P. M., Chepalyga, A. L. & Lavrentiev, N. V., 2010. The Khvalynian transgressions and early human settlement in the Caspian basin. Quaternary International 225, 152–159. Douka, K., Slon, V., Jacobs, Z., Bronk Ramsey, C., Shunkov, M. V., Derevianko, A. P., Mafessoni, F., Kozlikin, M. B., Li, B., Grün, R., Comeskey, D., Devièse, T., Brown, S., Viola, B., Kinsley, L., Buckley, M., Meyer, M., Roberts, R.G., Pääbo, S., Kelso J. & Higham, T., 2019. Age estimates for hominin fossils and the onset of the Upper Palaeolithic at Denisova Cave. Nature 565, 640–644. Duller, G. A. T., 2008. Single-grain optical dating of Quaternary sediments: why aliquot size matters in luminescence dating. Boreas 37, 589–612. Fenner, J. N., 2005. Cross-cultural estimation of the human generation interval for use in genetics-based population divergence studies. American Journal of Physical Anthropology 128, 415–423. FitzPatrick, E. A., 1993. Soil Microscopy and Micromorphology (Wiley, Chichester & New York). Galbraith, R. F. & Roberts, R. G., 2012. Statistical aspects of equivalent dose and error calculation and display in OSL dating: an overview and some recommendations. Quaternary Geochronology 11, 1–27. Galbraith, R. F., Roberts, R. G., Laslett, G. M., Yoshida, H. & Olley, J. M., 1999. Optical dating of single and multiple grains of quartz from Jinmium rock shelter, northern : Part I, experimental design and statistical models. Archaeometry 41, 339–364. Gerasimenko, N., 2005. Vegetation evolution of the Kabazi II site, in Palaeolithic Sites of Crimea, Volume 1, Kabazi II: Last Interglacial Occupation, Environment & Subsistence (eds Chabai, V., Richter, J. & Uthmeier, T.) 25–49 (Simferopol–Cologne, Shlyakh). Gladilin, V. N., 1976. Eastern Europe Upper Paleolithic Problems [in Russian] (Naukova dumka, Kiev). Glantz, M., Viola, B., Wrinn, P., Chikisheva, T., Derevianko, A., Krivoshapkin, A., Islamov, U., Suleimanov, R. & Ritzman, T., 2008. New hominin remains from Uzbekistan. Journal of Human Evolution 55, 223–237. Glantz, M., Van Arsdale, A., Temirbekov, S. & Beeton, T., 2018. How to survive the glacial apocalypse: hominin mobility strategies in late Pleistocene Central Asia. Quaternary International 466, 82–92.

96

Gnibidenko, Y. N., 2017. Environments of the northeastern East European Plain at the time of the initial human colonization, in Human Colonization of the Arctic: the interaction between early migration and the paleoenvironment (eds Kotlyakov, V. M., Velichko, A. A. & Vasil’ev, S. A.) 105–133 (Elsevier/Academic Press). Golovanova, L. V. & Doronichev, V. B., 2003. The Middle Paleolithic of the Caucasus. Journal of World Prehistory 18, 71–140. Golovanova, L. V. & Doronichev, V. B., 2017. The dynamics of stone industry transformation at the interface of Lower and Middle Paleolithic in the northwestern Caucasus. Quaternary International 428, 26–48. Golovanova, L. V., Hoffecker, J. F., Kharitonov, V. M. & Romanova, G. P., 1999. Mezmaiskaya Cave: Neanderthal occupation in the northern Caucasus. Current Anthropology 40, 77–86. Golovanova, L. V., Doronicheva, E. V., Doronichev, V. B. & Shirobokov, I. G., 2017. Bifacial scraper-knives in the Micoquian sites in the north-western Caucasus: typology, technology, and reduction. Quaternary International 428, 49–65. Goldberg, P. & Berna, F., 2010. Micromorphology and context. Quaternary International 214, 56–62. Grichuk, V. P., 1954. Materials to the paleobotany characteristic of the Quaternary and deposits of the northern-western part of the Precaspian lowland, in Materials on Geomorphology and Paleogeography of the USSR [in Russian] 11, 31–83 (Nauka, Moscow). Grosman, L., Smikt, O. & Smilansky, U., 2008. On the application of 3-D scanning technology for the documentation and typology of lithic artifacts. Journal of Archaeological Science 35, 3101–3110. Hajdinjak, M., Fu, Q., Hübner, A., Petr, M., Mafessoni, F., Grote, S., Skoglund, P., Narasimham, V., Rougier, H., Crevecoeur, I., Semal, P., Soressi, M., Talamo, S., Hublin, J.- J., Gušić, I., Kućan, Ž., Rudan, P., Golovanova, L. V., Doronichev, V. B., Posth, C., Krause, J., Korlević, P., Nagel, S., Nickel, B., Slatkin, M., Patterson, N., Reich, D., Prüfer, K., Meyer, M., Pääbo, S. & Kelso, J., 2018. Reconstructing the genetic history of late Neanderthals. Nature 555, 652–656. Hammer, Ø., Harper, D. A. T. & Ryan, P. D., 2001. PAST: paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4/1, article 4. Herzlinger, G. & Grosman, L., 2018. AGMT3-D: a software for 3-D landmarks-based geometric morphometric shape analysis of archaeological artifacts. PLoS ONE 13, e0207890. Herzlinger, G., Goren-Inbar, N. & Grosman L., 2017. A new method for 3D geometric morphometric shape analysis: the case study of handaxe knapping skill. Journal of Archaeological Science: Reports 14, 163–173. Horwitz, L. K. & Goldberg, P., 1989. A study of Pleistocene and Holocene hyaena coprolites. Journal of Archaeological Science 16, 71–94. Huntley, D. J. & Hancock, R. G. V., 2001. The Rb contents of the K-feldspars being measured in optical dating. Ancient TL 19, 43–46. Huntley, D. J., Godfrey-Smith, D. I. & Thewalt, M. L. W., 1985. Optical dating of sediments. Nature 313, 105–107. Hütt, G., Jaek, I. & Tchonka, J., 1988. Optical dating: K-feldspars optical response stimulation spectra. Quaternary Science Reviews 7, 381–385. Islamov, U. I., Zubov, A. A. & Kharitonov, V., 1988. Paleoliticheskaya stoyanka Sel’ungur v Ferganskoy doline (The Paleolithic site of Sel’ungur in the Fergana Valley) [in Russian]. Voprosy Antropologii 80, 38–49. Jacobs, Z. & Roberts, R. G., 2007. Advances in optically stimulated luminescence dating of individual grains of quartz from archeological deposits. Evolutionary Anthropology 16, 210–223.

97

Jacobs, Z. & Roberts, R. G., 2015. An improved single grain OSL chronology for the sedimentary deposits from Diepkloof Rockshelter, Western Cape, South Africa. Journal of Archaeological Science 63, 175–192. Jacobs, Z., Duller, G. A. T. & Wintle, A. G., 2006. Interpretation of single grain De distributions and calculation of De. Radiation Measurements 41, 264–277. Jacobs, Z., Li, B., Shunkov, M. V., Kozlikin, M. B., Bolikhovskaya, N. S., Agadjanian, A. K., Uliyanov, V. A., Vasiliev, S. K., O’Gorman, K., Derevianko, A. P. & Roberts, R. G., 2019. Timing of archaic hominin occupation of Denisova Cave in southern Siberia. Nature 565, 594–599. Jaubert, J., 2011. Les archéo-séquences du Paléolithique moyen du sud-ouest de la France: quel bilan un quart de siècle après François Bordes?, in François Bordes et al Préhistoire: Colloque International François Bordes (eds Delpech, F. & Jaubert, J.), 235–253 (Éditions du Comité des travaux historiques et scientifiques, Paris). Jongerius, A. (ed.), 1964. Soil Micromorphology (Elsevier, Amsterdam). Jöris, O., 2001. Der spätmittelpaläolithische Fundplatz Buhlen (Grabungen 1966–69), Stratigraphie, Steinartefakte und Fauna des Oberen Fundplatzes. Universitätsforschungen zur prähistorischen Archäologie 73 [in German] (Dr Rudolf Habelt GmbH, Bonn). Jöris, O. 2002. Out of the cold. On late Néandertal population dynamics in central Europe. Notae Praehistoricae 22, 33–45. Jöris, O., 2003. On the chronostratigraphic position of the late Middle Palaeolithic Keilmessergruppen. The attempt of a cultural-geographical demarcation of a Middle Palaeolithic group of forms in their European context, in Bericht der Römisch- Germanischen Kommission [in German] 84, 49–153 (German Archaeological Institute, Frankfurt am Main). Jöris, O., 2006. Bifacially backed knives in the Central European Middle Palaeolithic, in Axe Age. Acheulian tool-making from quarry to discard (eds Goren-Inbar, N. & Sharon, G.) 287–310 (Equinox, London). Karavanić, I. & Smith, F. H., 1998. The Middle/Upper Paleolithic interface and the relationship of Neanderthals and early modern humans in the Hrvatsko Zagorje, Croatia. Journal of Human Evolution 34, 223–248. Karavanić, I., Vukosavljević, N., Janković, I., Ahern, J. C. M. & Smith, F. H., 2018. Paleolithic hominins and settlement in Croatia from MIS 6 to MIS 3: research history and current interpretations. Quaternary International 494, 152–166. Kocovsky, P. M., Adams, J. V. & Bronte, C. R., 2009. The effect of sample size on the stability of principal components analysis of truss-based fish morphometrics. Transactions of the American Fisheries Society 138, 487–496. Kolobova, K. A., Krivoshapkin, A. I., Pavlenok, K. K., Flas, D., Derevianko, A. P. & Islamov, U. I., 2012. The denticulate Mousterian as a supposedly distinct facies in western Central Asia. Archaeology, Ethnology and Anthropology of Eurasia 40(1), 11–23. Kolobova, K. A., Flas, D., Krivoshapkin, A. I., Pavlenok, K. K., Vandenberghe, D. & De Dapper, M., 2018. Reassessment of the (Acheulean) presence in the western Tien Shan. Archaeological and Anthropological Sciences 10, 615–630. Kolobova, K. A., Shalagina, A. V., Chabai, V. P., Markin S. V. & Krivoshapkin, A. I., 2019. The significance of bifacial technologies in Altai Middle Paleolithic. L’Anthropologie 123, 276–288. Kozlikin, M. B., 2017. The Paleolithic Complexes from the Eastern Chamber of Denisova Cave [in Russian] (PhD thesis, Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Krajcarz, M. & Krajcarz, M. T., 2019. Post-depositional bone destruction in cave sediments: a micromorphological study of the MIS 5a–d cave bear strata of Biśnik Cave, . Journal of Quaternary Science 34, 138–152.

98

Krause, J., Orlando, L., Serre, D., Viola, B., Prüfer, K., Richards, M. P., Hublin, J.-J., Hänni, C., Derevianko, A. P. & Pääbo, S., 2007. Neanderthals in central Asia and Siberia. Nature 449, 902–904. Krause, J., Fu, Q., Good, J. M., Viola, B., Shunkov, M. V., Derevianko, A. P. & Pääbo, S., 2010. The complete mitochondrial DNA genome of an unknown hominin from southern Siberia. Nature 464, 894–897. Kreutzer, S., Schmidt, C., Fuchs, M.C., Dietze, M., Fischer, M. & Fuchs, M., 2012. Introducing an R package for luminescence dating analysis. Ancient TL 30, 1–8. Krivoshapkin, A. I., 2012. Obirakhmatian Middle to Upper Paleolithic Transitional Variant in Central Asia [in Russian] (PhD thesis, Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Krivoshapkin, A. I., Shalagina, A., Baumann, M., Shnaider, S. & Kolobova, K., 2018. Between Denisovans and Neanderthals: Strashnaya Cave in the Altai Mountains. Antiquity 92(365), e1. Kromer, B., Lindauer, S., Synal, H.-A. & Wacker, L., 2013. MAMS – a new AMS facility at the Curt-Engelhorn-Centre for Archaeometry, Mannheim, Germany. Nuclear Instruments and Methods in Physics Research B 294, 11–13. Kuhlwilm, M., Gronau, I., Hubisz, M. J., de Filippo, C., Prado-Martinez, J., Kircher, M., Fu, Q., Burbano, H. A., Lalueza-Fox, C., de la Rasilla, M., Rosas, A., Rudan, P., Brajkovic, D., Kucan, Ž., Gušic, I., Marques-Bonet, T., Andrés, A. M., 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. Kuznetsova, L. V., 1985. Paleolithic of the Middle and Lower Volga Region [in Russian] (PhD thesis, Institute for the History of Material Culture, Russian Academy of Sciences, Leningrad). Langergraber, K. E., Prüfer, K., Rowney, C., Boesch, C., Crockford, C., Fawcett, K., Inoue, E., Inoue-Muruyama, M., Mitani, J. C., Muller, M. N., Robbins, M. M., Schubert, G., Stoinski, T. S., Viola, B., Watts, D., Wittig, R. M., Wrangham, R. W., Zuberbühler, K., Pääbo, S. & Vigilant, L., 2012. Generation times in wild chimpanzees and gorillas suggest earlier divergence times in great ape and human evolution. Proceedings of the National Academy of Sciences of the U.S.A. 109, 15716–15721. Larkin, N. R., Alexander, J. & Lewis, M. D., 2000. Using experimental studies of recent faecal material to examine hyaena coprolites from the West Runton Freshwater Bed, Norfolk, UK. Journal of Archaeological Science 27, 19–31. Li, B., Roberts, R. G., Jacobs, Z. & Li, S.-H., 2014. A single-aliquot luminescence dating procedure for K-feldspar based on the dose-dependent MET-pIRIR signal sensitivity. Quaternary Geochronology 20, 51–64. Li, B., Jacobs, Z. & Roberts, R. G., 2016. Investigation of the applicability of standardised growth curves for OSL dating of quartz from cave, . Quaternary Geochronology 35, 1–15. Li, B., Jacobs, Z., Roberts, R. G., Galbraith, R. & Peng, J., 2017a. Variability in quartz OSL signals caused by measurement uncertainties: problems and solutions. Quaternary Geochronology 41, 11–25. Li, B., Jacobs, Z. & Roberts, R. G., 2017b. An improved multiple-aliquot regenerative-dose (MAR) procedure for post-IR IRSL dating of K-feldspar. Ancient TL 35, 1–10. Li, B., Roberts, R. G., Jacobs, Z. & Li, S.-H., 2018. Single‐grain dating of potassium‐rich feldspar grains: towards a global standardised growth curve for the post‐IR IRSL signal. Quaternary Geochronology 45, 23–36. Lubin, V. P. & Autlev, P. U., 1994. Lithic assemblage from Mousterian layer, in Neandertals from Gypskiy Canyon in Northern Cacasus [in Russian] (ed. Lubin, V. P.) 99–141 (Meoty, Maycop).

99

Madeyska, T., 2006. Sediments of Wylotne Rockshelter, in Wylotne and Zwierzyniec, Paleolithic Sites in Southern Poland (ed. Kozłowski, S. K.) 51–57 (Polish Academy of Arts and Sciences, Warsaw University, Kraków). Mafessoni, F., de Filippo, C., Slon, V., Grote, S., Chintalapati, M., Peter, B., Viola, B., Markin, S. V., Vasilyev, S. K., Rudaya, N. A., Kolobova, K. A., Shunkov, M. V., Derevianko, A. P., Kelso, J., Meyer, M., Prüfer, K. & Pääbo, S., 2018. A high-coverage Neandertal genome from Chagyrskaya Cave, in The Origins of the Upper Paleolithic in Eurasia and the Evolution of the Genus Homo (Proceedings of the International Symposium, Denisova Cave, Altai, Russia, July 2–8, 2018) 51–55 (Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Mallol, C. & Mentzer, S. M., 2017. Contacts under the lens: perspectives on the role of microstratigraphy in archaeological research. Archaeological and Anthropological Sciences 9, 1645–1669. Mania, D. & Toepfer, V., 1973. Königsaue. Gliederung, Ökologie und mittelpaläolithishe Funde der letzten Eiszeit. Veröffentlichungen des Landesmuseums für Vorgeschichte in Halle 26 [in German]. Martynovich, N. V., Markin, S. V. & Kolobova, K. A., 2016. New data about birds in the Chagyrskaya Cave (northwest Altai), 2012–2015 excavations, in Problems of Archaeology, Ethnography and Anthropology of Siberia and Neighbouring Territories 22, 114–117 [in Russian] (Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Mednikova, M. B. 2011. Postkranialnaya morfologiya i taksonomiya predstavitelei roda Homo iz peschery Okladnikova na Altae [in Russian] (Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Mednikova, M. B., 2013. An archaic human ulna from Chagyrskaya Cave, Altai: morphology and taxonomy. Archaeology, Ethnology and Anthropology of Eurasia 41(1), 66–77. Mercier, N. & Falguères, C., 2007. Field gamma dose-rate measurement with a NaI(Tl) detector: re-evaluation of the “threshold” technique. Ancient TL 25, 1–4. Meyer, M., Kircher, M., Gansauge, M.-T., Li, H., Racimo, F., Mallick, S., Schraiber, J. G., Jay, F., Prüfer, K., de Filippo, C., Sudmant, P. H., Alkan, C., Fu, Q., Do, R., Rohland, N., Tandon, A., Siebauer, M., Green, R. E., Bryc, K., Briggs, A. W., Stenzel, U., Dabney, J., Shendure, J., Kitzman, J., Hammer, M. F., Shunkov, M. V., Derevianko, A. P., Patterson, N., Andrés, A. M., Eichler, E. E., Slatkin, M., Reich, D., Kelso, J. & Pääbo, S., 2012. A high-coverage genome sequence from an archaic Denisovan individual. Science 338, 222– 226 (2012). Monigal, K., 2002. The Levantine Leptolithic: blade technology from the Lower Paleolithic to the dawn of the Upper Paleolithic (PhD thesis, Southern Methodist University, Dallas). Morales, J. I., Soto, M., Lorenzo, C. & Vergès, J. M., 2015. The evolution and stability of stone tools: the effects of different mobility scenarios in tool reduction and shape features. Journal of Archaeological Science: Reports 3, 295–305. Morley, M. W., Goldberg, P., Sutikna, T., Tocheri, M. W., Prinsloo, L. C., Jatmiko, Wayhu Saptomo, E., Wasisto, S. & Roberts, R. G., 2017. Initial micromorphological results from , Flores (): site formation processes and hominin activities at the type locality of Homo floresiensis. Journal of Archaeological Science 77, 125–142. Morley, M. W., Goldberg, P., Uliyanov, V. A., Kozlikin, M. B., Shunkov, M. V., Derevianko, A. P., Jacobs, Z. & Roberts, R. G., 2019. Hominin and animal activities in the microstratigraphic record from Denisova Cave (Altai Mountains, Russia). Scientific Reports 9, 13785. Musgrave, J. H., 1971. How dextrous was Neanderthal man? Nature 233, 538–541. Niewoehner, W. A., 2006. Neanderthal hands in their proper perspective, in Neanderthals Revisited: new approaches and perspectives (eds Harvati, K. & Harrison, T.) 157–190 (Springer, Dordrecht).

100

Ocherednoi, A., Salnaya N., Voskresenskaya, E. & Vishnyatsky, L., 2014. New geoarcheological studies at the Middle Paleolithic sites of Khotylevo I and Betovo (Bryansk oblast, Russia): some preliminary results. Quaternary International 326–327, 250–260. Okladnikov, A. P., 1949. Issledovanye musterskoy stoyanki i pogrebeniya neandertaltsa v grote Teshik-Tash, Iuzhny Uzbekistan (Srednaya Asiya), in Teshik-Tash – Paleolitichesky chelovek [in Russian] (eds Gremyatskiy, M. A. & Nesturkh, M. F.) 7–85 (Izdatelstvo Moskovskogo Gosudarstvennogo Universiteta (Moscow State University Press), Moscow). Pavlov, P. Y., Roebroeks, W. & Svendsen, J. I., 2004. The Pleistocene colonization of northeastern Europe: a report on recent research. Journal of Human Evolution 47, 3–17. Pavlov, P. Y., 2017. Middle and Upper Paleolithic sites in the northeast of the East European Plain, in Human Colonization of the Arctic: the interaction between early migration and the paleoenvironment (eds Kotlyakov, V. M., Velichko, A. A. & Vasil’ev, S. A.) 137–150 (Elsevier/Academic Press). Peng, J., Dong, Z., Han, F., Long, H. & Liu, X., 2013. R package numOSL: numeric routines for optically stimulated luminescence dating. Ancient TL 31, 41–48. Peng, J. & Li, B., 2017. Single-aliquot regenerative-dose (SAR) and standardised growth curve (SGC) equivalent dose determination in a batch model using the R package ‘numOSL’. Ancient TL 35, 32–53. Petraglia, M., Korisettar, R., Boivin, N., Clarkson, C., Ditchfield, P., Jones, S., Koshy, J., Lahr, M. M., Oppenheimer, C., Pyle, D., Roberts, R., Schwenninger, J.-L., Arnold, L. & White, K., 2007. Middle Paleolithic assemblages from the Indian subcontinent before and after the Toba super-eruption. Science 317, 114–116. Pinhasi, R., Higham, T. F. G., Golovanova, L. V. & Doronichev, V. B., 2011. Revised age of late Neanderthal occupation and the end of the Middle Paleolithic in the northern Caucasus. Proceedings of the National Academy of Sciences of the U.S.A. 108, 8611–8616. Powell, R., Hergt, J. & Woodhead, J., 2002. Improving isochron calculations with robust statistics and the bootstrap. Chemical Geology 185, 191–204. Praslov, N. D., 1984. The geological and paleogeographical limits of Paleolithic. The evolution on nature on the territory of the USSR and questions of chronology and periodization of Paleolithic, in The Paleolithic of the USSR [in Russian] (ed. Boriskovski, P. I.) 17–40 (Nauka, Moscow). Prescott, J. R. & Hutton, J. T., 1994. Cosmic ray contributions to dose rates for luminescence and ESR dating: large depths and long-term time variations. Radiation Measurements 23, 497–500. Prüfer, K., Racimo, F., Patterson, N., Jay, F., Sankararaman, S., Sawyer, S., Heinze, A., Renaud, G., Sudmant, P. H., de Filippo, C., Li, H., Mallick, S., Dannemann, M., Fu, Q., Kircher, M., Kuhlwilm, M., Lachmann, M., Meyer, M., Ongyerth, M., Siebauer, M., Theunert, C., Tandon, A., Moorjani, P., Pickrell, J., Mullikin, J. C., Vohr, S. H., Green, R. E., Hellmann, I., Johnson, P. L., Blanche, H., Cann, H., Kitzman, J. O., Shendure, J., Eichler, E. E., Lein, E. S., Bakken, T. E., Golovanova, L. V., Doronichev, V. B., Shunkov, M. V., Derevianko, A. P., Viola, B., Slatkin, M., Reich, D., Kelso, J. & Pääbo, S., 2014. The complete genome sequence of a Neanderthal from the Altai Mountains. Nature 505, 43–49. Prüfer, K., de Filippo, C., Grote, S., Mafessoni, F., Korlević, P., Hajdinjak, M., Vernot, B., Skov, L., Hsieh, P., Peyrégne, S., Reher, D., Hopfe, C., Nagel, S., Maricic, T., Fu, Q., Theunert, C., Rogers, R., Skoglund, P., Chintalapati, M., Dannemann, M., Nelson, B. J., Key, F. M., Rudan, P., Kućan, Z., Gušić, I., Golovanova, L. V., Doronichev, V. B., Patterson, N., Reich, D., Eichler, E. E., Slatkin, M., Schierup, M. H., Andrés, A., Kelso, J., Meyer, M. & Pääbo, S., 2017. A high-coverage Neandertal genome from Vindija Cave in Croatia. Science 358, 655–658. Reich, D., Green, R. E., Kircher, M., Krause, J., Patterson, N., Durand, E. Y., Viola, B., Briggs, A. W., Stenzel, U., Johnson, P. L. F., Maricic, T., Good, J. M., Marques-Bonet, T.,

101

Alkan, C., Fu, Q., Mallick, S., Li, H., Meyer, M., Eichler, E. E., Stoneking, M., Richards, M., Talamo, S., Shunkov, M. V., Derevianko, A. P., Hublin, J.-J., Kelso, J., Slatkin, M. & Pääbo, S., 2010. Genetic history of an archaic hominin group from Denisova Cave in Siberia. Nature 468, 1053–1060. Reimer, P. J., Bard, E., Bayliss, A., Beck, J. W., Blackwell, P. G., Bronk Ramsey, C., Buck, C. E., Cheng, H., Edwards, R. L., Friedrich, M., Grootes, P. M., Guilderson, T. P., Haflidason, H., Hajdas, I., Hatté, C., Heaton, T. J., Hoffmann, D. L., Hogg, A. G., Hughen, K. A., Kaiser, K. F., Kromer, B., Manning, S. W., Niu, M., Reimer, R. W., Richards, D. A., Scott, E. M., Southon, J. R., Staff, R. A., Turney, C. S. M., Van der Plicht, J., 2013. IntCal13 and Marine13 radiocarbon age calibration curves 0–50,000 years cal BP. Radiocarbon 55, 1869–1887. Rhodes, E. J. & Schwenninger, J.-L., 2007. Dose rates and radioisotopes in the concrete calibration blocks at Oxford. Ancient TL 25, 5–8. Richter, D., Mauz, B., Böhner, U., Weissmüller, W., Wagner, G. A., Freund, G., Rink, W. J. & Richter, J., 2000. Luminescence dating of the Middle/Upper Paleolithic sites ‘Sesselfelsgrotte’ and ‘Abri I am Schulerloch’, Altmühltal, Bavaria, in Neanderthals and Modern Humans – Discussing the Transition: central and eastern Europe from 50.000– 30.000 B.P. (eds Orschiedt, J. & Weniger G.-C.) 30–41 (Neanderthal Museum, Mettmann). Richter, J., 1997. Sesselfelsgrotte III: Der G-Schichten-Komplex der Sesselfelsgrotte. Zum Verständnis des . Quartär-Bibliothek 7 (Saarbrücken). Richter, J., 2000. Social memory among late Neanderthals, in Neanderthals and Modern Humans – Discussing the Transition: central and eastern Europe from 50.000–30.000 B.P. (eds Orschiedt, J. & Weniger G.-C.) 123–132 (Neanderthal Museum, Mettmann). Richter, J., 2002. Die 14C-daten aus der Sesselfelsgrotte und die Zeitstellung des Micoquien/ M.M.O. Germania 80, 1–22. Richter, J., 2016. Leave at the height of the party: a critical review of the Middle Paleolithic in Western Central Europe from its beginnings to its rapid decline. Quaternary International 411, 107–128. Roberts, R. G. & Jacobs, Z., 2018. Timelines for human evolution and dispersals. Elements 14, 27–32. Roberts, R. G., Galbraith, R. F., Yoshida, H., Laslett, G. M. & Olley, J. M., 2000. Distinguishing dose populations in sediment mixtures: a test of single-grain optical dating procedures using mixtures of laboratory-dosed quartz. Radiation Measurements 32, 459– 465. Roberts, R. G., Jacobs, Z., Li, B., Jankowski, N. R., Cunningham, A. C. & Rosenfeld, A. B., 2015. Optical dating in archaeology: thirty years in retrospect and grand challenges for the future. Journal of Archaeological Science 56, 41–60. Roberts, R. G., Jacobs, Z. & Li, B., 2018. Optical dating of sediment samples from Chagyrskaya Cave, in Multidisciplinary Studies of Chagyrskaya Cave – A Middle Paleolithic Site in Altai (Derevianko, A. P. et al.) 121–150 [in Russian] and 353–369 (Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences Press, Novosibirsk). Rousseeuw, P. J. & Croux, C., 1993. Alternatives to the median absolute deviation. Journal of the American Statistical Association 88, 1273–1283. Rousseeuw, P. J., Debruyne, M., Engelen, S. & Hubert, M., 2006. Robustness and outlier detection in chemometrics. Critical Reviews in Analytical Chemistry 36, 221–242. Rudaya, N. A., 2013. Environmental conditions during the early human settlement of Chagyrskaya Cave (Altai). Archaeology, Ethnology and Anthropology of Eurasia 41(1), 45–54. Rudaya, N., Vasiliev, S., Viola, B., Talamo, S. & Markin, S., 2017. Palaeoenvironments during the period of the Neanderthals settlement in Chagyrskaya Cave (Altai Mountains, Russia). Palaeogeography, Palaeoclimatology, Palaeoecology 467, 265–276.

102

Sawyer, S., Renaud, G., Viola, B., Hublin, J.-J., Gansauge, M.-T., Shunkov, M. V., Derevianko, A. P., Prüfer, K., Kelso, J. & Pääbo, S., 2015. Nuclear and mitochondrial DNA sequences from two Denisovan individuals. Proceedings of the National Academy of Sciences of the U.S.A. 112, 15696–15700. Schwartz, G., 1978. Estimating the dimensions of a model. Annals of Statistics 6, 461–464. Serwatka, K., 2014. Shape variation of Middle Palaeolithic bifacial tools from southern Poland: a geometric morphometric approach to Keilmessergruppen handaxes and backed knives. Lithics 35, 18−32. Shpakova, E. G., 2001. Paleolithic human dental remains from Siberia. Archaeology, Ethnology and Anthropology of Eurasia 4(8), 64–76. Shpakova, E. G. & Derevianko, A. P., 2000. The interpretation of odontological features of Pleistocene human remains from the Altai. Archaeology, Ethnology and Anthropology of Eurasia 1(1), 125–138. Shalagina, A. V., 2016. The Middle Paleolithic industries of Strashnaya Cave, in Siberian Archeology and Ethnography: the contribution of young researchers [in Russian] (ed. Konstantinov, A. V.) 8–69 (Transbaikal State University, Chita). Shalagina, A. V., Krivoshapkin, A. I. & Kolobova, K. A., 2015. Truncated-faceted pieces in the Paleolithic of northern Asia. Archaeology, Ethnology and Anthropology of Eurasia 43(4), 33–45. Shaukat, S. S., Rao, T. A. & Khan, M. A., 2016. Impact of sample size on principal component analysis ordination of an environmental data set: effects on eigenstructure. Ekológia (Bratislava) 35, 173–190. Shunkov, M. V., 1990. Mouserian Sites of Central Altai Intermontane Basins (Nauka, Novosibirsk). Skinner, A. R., Blackwell, B. A. B., Martin, S., Ortega, A., Blickstein, J. I. B., Golovanova, L. V. & Doronichev, V. B., 2005. ESR dating at Mezmaiskaya Cave, Russia. Applied Radiation and Isotopes 62, 219–224. Skoglund, P., Northoff, B. H., Shunkov, M. V., Derevianko, A. P., Pääbo, S., Krause, J. & Jakobsson, M., 2014. Separating endogenous ancient DNA from modern day contamination in a Siberian Neanderthal. Proceedings of the National Academy of Sciences of the U.S.A. 111, 2229–2234. Slimak, L., Svendsen, J. I., Mangerud, J., Plisson, H., Heggen, H. P., Brugere, A. & Pavlov, P. Y., 2011. Late Mousterian persistence near the Arctic Circle. Science 332, 841–845. Slon, V., Hopfe, C., Weiß, C. L., Mafessoni, F., de la Rasilla, M., Lalueza-Fox, C., Rosas, A., Soressi, M., Knul, M. V., Miller, R., Stewart, J. R., Derevianko, A. P., Jacobs, Z., Li, B., Roberts, R. G., Shunkov, M. V., de Lumley, H., Perrenoud, C., Gušić, I., Kućan, Ž., Rudan, P., Aximu-Petri, A., Essel, E., Nagel, S., Nickel, B., Schmidt, A., Prüfer, K., Kelso, J., Burbano, H. A., Pääbo, S. & Meyer, M., 2017a. Neandertal and Denisovan DNA from Pleistocene sediments. Science 356, 605–608. Slon, V., Viola, B., Renaud, G., Gansauge, M.-T., Benazzi, S., Sawyer, S., Hublin, J.-J., Shunkov, M. V., Derevianko, A. P., Kelso, J., Prüfer, K., Meyer, M., Pääbo, S., 2017b. A fourth Denisovan individual. Science Advances 3, e1700186. Slon, V., Mafessoni, F., Vernot, B., de Filippo, C., Grote, S., Viola, B., Hajdinjak, M., Peyrégne, S., Nagel, S., Brown, S., Douka, K., Higham, T., Kozlikin, M. B., Shunkov, M. V., Derevianko, A. P., Kelso, J., Meyer, M., Prüfer, K. & Pääbo, S., 2018. The genome of the offspring of a Neanderthal mother and a Denisovan father. Nature 561, 113–116. Smedley, R. K., Duller, G. A. T., Pearce, N. J. G. & Roberts, H. M., 2012. Determining the K- content of single-grains of feldspar for luminescence dating. Radiation Measurements 47, 790–796. Soressi, M., 2005. Late Mousterian lithic technology: its implications for the pace of the emergence of behavioural modernity and the relationship between behavioural modernity

103

and biological modernity, in From Tools to Symbols: from early hominids to modern humans (eds d’Errico, F. & Backwell, L.) 389–417 (Wits University Press, Johannesburg). Stoops, G., 2003. Guidelines for Analysis and Description of Soil and Regolith Thin Sections (Soil Science Society of America Inc., Wisconsin). Svitoch, A. A., 2012. The Caspian Sea shelf during the Pleistocene regressive epochs. Oceanology 52, 526–539. Taguchi, Y.-H. & Oono, Y., 2005. Relational patterns of gene expression via non-metric multidimensional scaling analysis. Bioinformatics 21, 730–740. Talamo, S. & Richards, M., 2011. A comparison of bone pretreatment methods for AMS dating of samples >30,000 BP. Radiocarbon 53, 443–449. Thiébaut, C., Claud, É., Deschamps, M., Discamps, E., Soulier, M.-C., Mussini, C., Costamagno, S., Rendu, W., Brenet, M., Colonge, D., Coudenneau, A., Gerbe, M., Guibert, P., Jaubert, J., Laroulandie, V., Maureille, B., Mourre, V. & Santos, F., 2014. Diversité des productions lithiques du Paléolithique moyen récent (OIS 4–OIS 3): enquête sur le rôle des facteurs environnementaux, fonctionnels et culturels, in Transitions, Ruptures et Continuité en Préhistoire (eds Jaubert, J., Fourment, N. & Depaepe, P.) 281–298 (Société Préhistorique Française, Paris). Trinkaus, E., 1983. The Shanidar Neanderthals (Academic Press, New York). Turner II, C. G., 1990. Paleolithic teeth of the Central Siberian Altai Mountains, in Chronostratigraphy of the Paleolithic in North, Central, East Asia and America (ed. Derevianko, A. P.) 239–243 (Institute of History, Philology and Philosophy, Siberian Branch of the USSR Academy of Sciences, Novosibirsk). Valoch, K., 1988. Die Erforschung der Külna-Höhle 1961–1976. Anthropos 24 (Brno). Van Klinken, G. J., 1999. Bone collagen quality indicators for palaeodietary and radiocarbon measurements. Journal of Archaeological Science 26, 687–695. Van Vliet-Lanöe, B., 2010. Frost action, in Interpretation of Micromorphological Features of Soils and Regoliths (eds Stoops, G., Marcelino, V. & Mees, F.) 81–108 (Elsevier, Amsterdam). Vasiliev, S. K., 2013. Large mammal fauna from the Pleistocene deposits of Chagyrskaya Cave, northwestern Altai (based on 2007–2011 excavations). Archaeology, Ethnology and Anthropology of Eurasia 41(1), 28–44. Veil, S., Breest, K., Höfle, H.-C., Meyer, H.-H., Plisson, H., Urban-Küttel, B., Wagner, G. A. & Zöller, L., 1994. Ein mittelpaläolithischer Fundplatz aus der Weichsel-Kaltzeit bei Lichtenberg, Lkr. Lüchow-Dannenberg. Zwischenbericht über die archäologischen und geowissenschaftlichen Untersuchungen 1987–1992 [in German]. Germania 72, 1–66. Vernot, B. & Akey, J. M., 2014. Resurrecting surviving Neandertal lineages from modern human genomes. Science 343, 1017–1021. Villanea, F. A. & Schraiber, J. G., 2019. Multiple episodes of interbreeding between Neanderthal and modern humans. Nature Ecology and Evolution 3, 39–44. Viola, T. B., 2009. New Hominid Remains from Central Asia and Siberia: the easternmost Neanderthals? (PhD thesis, University of Vienna, Vienna). Viola, T. B., Markin, S. V., Zenin, A., Shunkov, M. V. & Derevianko, A. P., 2011. Late Pleistocene hominins from the Altai Mountains, Russia, in Characteristic Features of the Middle to Upper Palaeolithic Transition in Eurasia (eds Derevianko, A. P. & Shunkov, M. V.) 207–213 (Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences, Novosibirsk). Viola, T. B., Markin, S. V., Buzhilova, A. P., Mednikova, M. B., Dobrovolskaya, M. V., Le Cabec, A., Shunkov, M. V., Derevianko, A. P. & Hublin, J.-J., 2012. New Neanderthal remains from Chagyrskaya Cave (Altai Mountains, Russian Federation). American Journal of Physical Anthropology 147, supplement 54, 293–294.

104

Wall, J. D., Yang, M. A., Jay, F., Kim, S. K., Durand, E. Y., Stevison, L. S., Gignoux, C., Woerner, A., Hammer, M. F. & Slatkin, M., 2013. Higher levels of Neanderthal ancestry in East Asians than in Europeans. Genetics 194, 199–209. Weiss, M., Lauer, T., Wimmer, R. & Pop, C. M., 2018. The variability of the Keilmesser- concept: a case study from central Germany. Journal of Paleolithic Archaeology 1, 202– 246. Wood, R., 2015. From revolution to convention: the past, present and future of radiocarbon dating. Journal of Archaeological Science 56, 61–72. Yanina, T., 2014. The Ponto-Caspian region: environmental consequences of climate change during the late Pleistocene. Quaternary International 345, 88–99. Yanina, T., Sorokin, V., Bezrodnykh, Y. & Romanyuk, B., 2018. Late Pleistocene climatic events reflected in the Caspian Sea geological history (based on drilling data). Quaternary International 465, 130–141.

105