<figure><img src="image_26.png" /><figcaption>**Fig. 25. Kara-Bom: archaeological stratigraphy and dating.** Site plan and excavation grid (after Rybin et al., 2023, modified), with positioning of the jewellery kit and of Goebel et al.'s (1993) samples onto (a) the latter's Z>I stratigraphic profile and (b) the distribution in columns Z and I of Slavinsky et al.'s (2016) UP1 and UP2 clusters of refit links (caption: "1. vertical profile of rock along line Z; 2. distribution of artifacts made from various groups of raw materials in UP1; 3. distribution of artifacts made from various groups of raw materials of UP2; 4. distribution of combustion structures").</figcaption></figure>

coeval, similar activity.

According to Slavinski et al., "the technique of subprismatic bidi-
rectional detachment of medium sized and large laminar blanks was
determinative for the industries of both horizons." If anything, this
continuity in technology suggests that both horizons are close in time
and that the significantly earlier age obtained for the hearth feature that
Goebel et al. excavated below UP2 is but a *terminus post quem* for the
overlying scatters of refitted stone tools. What exactly does that hearth
stand for? The answer may lie in the stone tools found in basal stratum 6
that did not make it into the UP2 refit sets (which, for UP1 and UP2
combined, involve no more than 294 artefacts out of a total of 2010).
Until information is provided on what such material looks like, we will
not be able to know. What we do know — not unexpectedly, given the
deposit's colluvial nature and periglacial context — is that the com-
mingling of charcoal of at least three different ages is demonstrated in
stratum 5, and, so, that there is every reason to expect stratum 6 to also

be heterogeneous in content.

The new results reported by Rybin et al. (2023) do not change the
picture. Five are on piece-plotted bones from the 1991 excavation,
which Rybin et al. plot against Derevianko and Rybin's décapage plan
for grid units Z-M (Cyrillic 3–M)/7–9. The finds in the latter, however,
"were recovered from the lower portion of lithological layer 6 consisting
of grey sand and loam impregnated with gravel and schist plates," and
"the difference in relative levels of the finds within this layer is not more
than 38 cm: from –352 cm (from zero datum) in the south-western
corner of the study area to –314 cm in the unit's north-eastern corner;
the average levels vary from –316 to –340 cm, which corresponds to
the general dip of the excavation unit's floor." Yet the 1991 samples that
Rybin et al. dated come from between 1.0 m and 1.5 m above that plan's
elevations. With regard to the five 2016 bone samples taken in the M>N
(Cyrillic M>H) profile, they come from a deposit that lies on a rock ledge
forming a terrace in the eastern side of the Trench 1 excavation, where,
as Rybin et al. (2023) do acknowledge, “lithological Layers 1–6, con-
taining UP assemblages, [and] the underlying sediments with MP as-
semblages, were not found.” Accounting for the significant dip to west of
the deposit lying on the ledge (well apparent in the photo of the 1991
excavations published by Belousova et al., 2020) does not change the
fact that those samples come from well above the deposit that yielded
the site’s UP assemblages. Yet, they are statistically the same as the two
charcoal dates reported by Goebel et al. for the sediments found below
Slavinski et al.’s UP2 cluster of refit links (Fig. 26).

The meaning of these observations is self-evident to us: an animal
bone component of Middle Palaeolithic age, dating in the range of 40–55
ka ago, is scattered through Kara-Bom’s post-Middle Palaeolithic de-
posit. That each of the units differentiated in the latter contains inherited
material of a diverse chronology is otherwise corroborated by GX-

17592, Goebel et al.'s stratigraphically anomalous charcoal result. Such background noise probably reflects the gradual incorporation of remains derived by erosion from soils and natural thanatocoenoses originally formed upslope to the east — ones that gradually gravitated, or were washed down, all through the accumulation of the aeolian material forming lithological strata 1–6 of Derevianko and Rybin's layout. These conclusions are fully consistent with the fact that, except for the tool dated to 29,110 ± 320 BP (OxA-36907), none of Rybin et al.'s other bone samples bear cut marks or are otherwise humanly modified.

To sum up: the stone tool assemblages in the UP1 and UP2 horizons
of Kara-Bom are technologically similar and fully Upper Palaeolithic in
nature. Based on the charcoal ages associated with UP1, they are likely
to date to the time range of the Early Aurignacian of western Eurasia,

<figure><img src="image_27.png" /><figcaption>**Fig. 26.** **Kara-Bom:** stratigraphic distribution of the OxA bone dates. Top: frontal view of the rock face and the excavation trench (after Belousova et al., 2020, modified); the profiles from 1991 (Goebel et al., 1993; Derevianko and Rybin, 2003) and 2016 (Rybin et al., 2023) are indicated. Bottom: projection of the positions of the pit with the jewellery and of Rybin et al.'s (2023) samples across a schematic east-west section running along the separation between rows 8 and 9 of the grid.</figcaption></figure>
~39 ka ago. Albeit widely quoted on the basis of Goebel et al.'s results, the ~46 ka horizon for the regional emergence of this technology is in fact a *terminus post quem*. Otherwise, Rybin et al.'s Bayesian model is a good example of how a statistically robust methodology can be misapplied. By claiming to date a deposit based on the age of samples that, even though not collected in it, are posited to relate to it, they effectively engage in circular reasoning, affirming the consequent and proving nothing that they had not assumed to begin with. Finally, the stratigraphic relationship between the clusters of refit links and the symbolic artefacts in the small depression found in grid unit L8 vindicates Zilhão's (2007) proposition that it be seen as a cache related to the upper, not the lower find horizon.

### 5.4.2. Tolbor

At this extensive Mongolian site, unit 3c of the Tolbor 16 locality is claimed to document an Upper Palaeolithic occupation dated to within the 41.2–46.7 ka cal BP interval (Zwyns et al., 2019). The claim is based on Bayesian modelling of the radiocarbon ages obtained from the bone fragments (in total, 19 unidentifiable and >20 mm, plus 49 from sieving and <20 mm) therein associated with an 825-strong stone tool assemblage (362-strong, excluding the <20 mm sieve finds). However:

* Unit 3c is a solifluction lobe: "With a few exceptions, most of the samples from unit 3 show particle size distribution curves (...) consistent with reworking (...) or soil development (...). Although loess is the parent material, the dominant depositional process within unit 3 seems to be sheet erosion."

* The preservation of the bones is extremely poor, and no cut marks or other evidence of human modification could be observed: "When bones are preserved, they are fragmented, and the cortical surface is missing."

* The luminescence dating of unit 3c sediment yielded ages of 26.6 ± 2.6 ka (quartz OSL) and 35.3 ± 3.2 ka (pIR-IRSL290 polymineral OSL). When the estimates obtained for the same sample do not agree, the site's luminescence study argues that the quartz age should be retained. For unit 3c, doing so implies deposition no earlier than 31.8 ka, 10–15 millennia later than indicated by the radiocarbon-dated bone samples (even if we posit that the polymineral estimate should be preferred, a deposition age in the 28.9–41.7 ka interval, also much later than indicated by the radiocarbon ages, is nonetheless implied).

Zwyns et al.'s reasoning would seem to be that (a) the OSL results date the sediments' last exposure to light and therefore a remobilisation event, and (b) the time of deposition of the displaced human occupation remains found within the sediments must predate that event and be given by the age of the bones. The latter's poor preservation, however, is not due to chemical processes, given the good collagen content implied by the successful dating of 32 % (6 out of 19) of the >20 mm bone fragments from unit 3c, and the fact that 23 out of 24 samples submitted to ZooMS analysis could be taxonomically identified. Moreover, even if rarely so, bone fragments are also found in units that contain no lithics. Given this, the parsimonious interpretation of the bones' poor physical condition is that they were subject to processes of mechanical weathering, ones that do not seem to have affected the lithics. These observations contradict the notion that the bones must relate to the human activity represented by the artefacts, and, in any case, co-occurrence in a reworked deposit is no proof of contemporaneity.

Further evidence that the bone component of unit 3c's soliflucted lobes cannot be taken as a proxy for human activity is provided by the infinite date obtained for basal level 7 of Pit 1 (AA-93134, >45,400 BP; Zwyns et al., 2014). Zwyns et al. (2019) refer to the bone sample that this result derives from as follows: "its association with the archaeology was not clear." Therefore, either the rationale that the dated bones must all be associated with the archaeology is invalid, or, if the sample was associated with the archaeology, then the soliflucted accumulations'

faunal content is at least in part made up of residual material derived from the erosion of earlier deposits. AA-93134 therefore corroborates that Tolbor 16's radiocarbon ages provide no more than *terminii post quem* for the Upper Palaeolithic stone tools found alongside the dated bone samples.

The same is the case elsewhere at Tolbor. For instance, at Tolbor 4, Occupation Horizons OH5 and OH6 yielded an assemblage like that in unit 3c of Tolbor 16, which lies <10 km downstream (Derevianko et al., 2013; Zwyns et al., 2014). On ostrich eggshell, the Tolbor 4 dates are 31,210 ± 410 BP (AA-93140) and 35,230 ± 680 BP (AA-93141) for OH5 and OH6, respectively; on bone, the corresponding results are >41,050 BP (AA-79326) and 37,400 ± 2600 BP (AA-79314). These bone dates are stratigraphically inverted and corroborate the residuality of the poorly preserved faunal assemblage apparently associated with the lithics. The ostrich eggshell dates are stratigraphically consistent, but the older one is four to six millennia younger than the ~45 ka time horizon for the emergence of the IUP at Boker Tachtit.

Another example is Tolbor-21 (Rigaud et al., 2023; Rybin et al., 2020), where the surface condition of the faunal remains is unambiguously described as follows: "The frequency of cracking (26 % of NR) and exfoliation (42 % of NR) are consistent with a slow embedding of the bone assemblage. This finds an interesting echo in the high impact of the root etching observable on 62 % of the remains, attesting a protracted exposure of the remains to vegetal activity. In consequence, most of the cortical surface of bones is usually destroyed; indeed, 52 % of the remains have lost more than 75 % of their cortical surfaces." The contemporaneity of these fragmentary and heavily weathered remains (and, hence, of the dated samples) with the Upper Palaeolithic stone tools and symbolic artefacts of the site's AH (Archaeological Horizon) 4 is, therefore, undemonstrated. This is the more so given the nature — "continually accreting low- to moderate-energy slope deposits" — of the encasing sediment (Rybin et al., 2020). At best, thus, the dated bones from AH4 of Tolbor-21 can only provide a *terminus post quem* of 41–42 ka cal BP for the human occupation that left the artefacts behind.

### 5.4.3. Kamenka

Like Kara-Bom, Kamenka, an open-air site in the Transbaikal region of Siberia, is a slope deposit. Here, layer 6 of the stratigraphic succession is the unit of interest. It yielded two archaeological horizons at a depth of 7–9 m below surface. Based on the advanced stage of weathering of the small number of faunal remains and the wide scattering of the rare artefacts, the upper horizon, Complex B, is considered to be in derived position. The lower horizon, Complex A, is reported to contain hearth features tightly associated with clusters of faunal remains, lithic artefacts, and bone tools (awls, retouchers, and a decorated "whistle"); i.e., Complex A does contain reliable indicators of on-site, well-preserved human activity (Germonpré and Lbova, 1996; Zwyns, 2021; Zwyns and Lbova, 2019).

The stone tools found in Complex A look unambiguously Upper Palaeolithic; production is based on the same core reduction techniques documented by the Kara-Bom refits. It is from this observation and the age of the stratigraphically associated, AMS-dated bone samples that a claim is derived for Complex A to represent evidence for IUP-like assemblages to be found that far to the east as early as ~45 ka. The argument is predicated on the notion that the artefacts, the features, and the faunal remains are coeval, and, therefore, that the dates are a reliable measure of the age of the human activity. This assumption, however, does not stand up to scrutiny, as it overlooks the fact, well documented by Germonpré and Lbova, that Complex A's faunal assemblage consists of two components. One, formed of remains of horse and Mongolian gazelle, represents 90 % of the NISP, is relatively fresh, and bears cut marks and other evidence of human modification. The other, representing 10 % of the NISP, is weathered, bears no evidence of human modification, includes carnivore-gnawed specimens, and, alongside horse and gazelle, is mostly comprised of taxa not found among the fresh material, namely: bison, woolly rhino, giant deer,
mountain sheep, camel, and an extinct antelope.

This second component of Kamenka's Complex A fauna is, therefore, residual: environmental background material that the deposit incorporated penecontemporaneously with the human activity, for which it provides no more than a *terminus post quem*. Yet, out of the four AMS dates that Zwyns & Lbova take as reliable, two — OxA-12116 (39,290 ± 350 BP), and OxA-12117 (41,350 ± 450 BP) — are on such residual material, specifically, on giant deer bones: a fragment of humerus, and a shed antler (Hughes et al., 2006). A third date (Vasiliev et al., 2002), AA-26,743 (40,500 ± 3800 BP), is on a bone unidentified to taxon and comes with an uncertainty interval too large for practical use in the assessment of the age of Complex A. According to Hughes et al., the fourth date — GrA-5435 (37,350 ± 310/-490 BP) — comes from a cut-marked tibia of gazelle, probably the bone photographically illustrated in Germonpré and Lbova's Fig. 8, as that is the only such tibia that the latter identified in the assemblage.

Assuming the reliability of the zooarchaeologist's classification of the mark, GrA-5435 would be the only dated sample from Kamenka that one can securely associate with human activity. Whether such activity relates in turn to the artefact assemblage depends on whether the latter is homogeneous and reflects a single episode — and that, as we have seen in the case of Kara-Bom, is assuming a lot. Such an assumption provides the best scenario for the Upper Palaeolithic in Complex A to be of an early age — albeit one no older than the Ahmarian/Protoaurignacian. Thus, at Kamenka, what is of the same age as the IUP of the Near East, perhaps even older, is not "modern humans" bearing a comparable material culture but a giant deer fauna that significantly preceded any human presence at the site.

### 5.4.4. Tolbaga

Another Transbaikal example of the site formation issues raised by the Siberian open-air sites is Tolbaga (Izuho et al., 2019). Here, two distinct cultural layers have been recognised within the topsoil and another two, layers 3 and 4, within the underlying, ~60 cm-thick, muddy gravel that overlies another soil horizon. Most of the finds come from layer 4 and, according to Izuho et al., they provide a good illustration of the region's earliest Upper Palaeolithic, characterised by "blade-based lithic toolkits with Levallois elements, rich osseous tools produced through systematic procedures, and mobile art such as pendants or sculptures (...), found alongside stone-lined dwellings, stone-lined hearths, and storage pits."

Layer 4 of Tolbaga yielded no less than 12 dates on ultrafiltrated bone collagen, none of which would seem to bear cut marks or other evidence of human modification, even though many such bones have reportedly been identified in the faunal assemblage. Most of these layer 4 results fall in the 36–42 ka cal BP range, but one is as young as 24,070 ± 150 BP (UCIAMS-143239; 27,841–28,630 cal BP), and another is as old as 43,900 ± 1800 BP (UCIAMS-143220; 43,276–49,939 cal BP). As the succession formed via colluvial slope wash with intermittent soil formation, residuality issues must be of import here as much as at Kara-Bom, Kamenka, and Tolbor. There is therefore every reason to dismiss as background noise the single sample from the archaeological horizon that dates in the 45–50 ka cal BP range.

### 5.5. Denisova

Having clarified the overall Eurasian context, we are now in a position to assess the evidence from Denisova, a key Altai site where the preservation of aDNA is excellent and samples therefrom contributed major breakthroughs to the palaeogenetics of the Upper Pleistocene. While the Siberian localities we just reviewed are all open-air sites, Denisova is a cave in the study of which it is especially important to bear in mind the lessons of Turner et al.'s (2013) taphonomic revision of Palaeolithic Siberia's faunal assemblages.

Turner et al.'s key conclusion was that, whenever evidence of hyaena denning exists in a cave site, the association between artefacts and bones

ought to be considered suspect by default because of the species' burrowing behaviour: if, alongside solifluction, frost-wedging, and the other disturbance processes that ubiquitously affect cave deposits formed under periglacial conditions, hyaena burrowing is found to be a significant contributor to syn- and post-depositional disturbance, then the presumption that individual stratigraphic units must be homogeneous in content is therefore even less warranted than usually. This caveat is the more important to consider when assessing cultural change through the Middle and early Upper Palaeolithic of Siberia because both predate the regional extinction of hyaenas and because it applies to open-air sites as well. At Kara-Bom, for instance, Turner et al. found evidence that hyaenas had been involved in the accumulation of the faunal assemblage. This factor may additionally explain the issues of sample association that afflict that site's dating record.

Regrettably, Turner et al.'s advice has seldom been heeded. Denisova, where hyaenas were highly active, is a major case in point. Significant post-depositional disturbance is corroborated by Morley et al.'s (2019) soil micromorphological work, which, as argued by Kuzmin and Keates (2020) and d'Errico et al. (2024), suffices to explain the patently inconsistent dating results obtained for the site (Douka et al., 2019; Jacobs et al., 2019b; Reich et al., 2010). In fact, the Bayes-derived chronological frameworks such results have been forced into are only as good as the underlying assumptions, which, in our view and as we now proceed to argue, are questionable.

#### 5.5.1. Stratigraphy and dating

Given the issues with stratigraphic integrity, the bottom line at Denisova is that dating by association is not possible: only comparative typology, supplemented by the direct dating of diagnostic human remains and of artefacts made on bone or tooth, is amenable to providing reliable age estimates. A clear-cut illustration of the problems is provided by the discrepancy between the direct dating results obtained for bone fragments genetically assigned to the so-called Denisovan population and the ages obtained for samples with which those fossils were found in stratigraphic association. Based on the number of mutations observed in their mitochondrial genomes, all of the site's Denisovan individuals have been dated to >51.6 ka ago, which agrees with the >49.9 ka ago radiocarbon age for a bone representing the first generation offspring of an episode of Neandertal/Denisovan interbreeding (Douka et al., 2019; Slon et al., 2018). Yet the remains of such individuals were retrieved in levels for which ages up to at least 30,000 years younger have been obtained on both bone and charcoal samples. For instance, the Denisova 4 fossil (the upper permanent molar of an adult Denisovan male) comes from level 11.1 of the South Chamber, where a charcoal sample collected at the same elevation <2 m away is as young as 19,990 ± 61 BP (Essel et al., 2023) (Fig. 27).

In addition, the site's Pleistocene sequence is exposed in three different chambers; the trenches therein excavated are not contiguous, their lateral correlation is not straightforward, and problems exist even within a single trench. Take for instance the stratigraphic G>D (Cyrillic Γ>Δ)/6–5 profile of the South Chamber. Fig. 28, where the compared images are taken from Jacobs et al. (2019b), leaves no doubt that the two photos record the same profile — as it remained after the 2003 excavations, on one hand, and after its 2016 sampling, on the other. Yet the deposit that OSL samples 1–3 of 2012 and 1–3 of 2016 are reported to provide an age for is designated level 11 in the "after 2003" photo and associated drawing but said to be the provenience of the OSL samples from level 12 in the 2016 photo. Given that all the remains genetically assigned to either Neandertals or Denisovans were excavated between 1984 and 2012, and seven out of eight come with either a level 11 (five) or level 12 (two) label (Douka et al., 2019), one cannot but wonder about their reported association with the ages published post-2016 for samples carrying the same level labels, i.e., "11" and "12" (and much the same applies to the site's directly dated animal tooth pendants): Do the finds and the samples come indeed from the same stratigraphic units, or do they simply carry the same labels, ones whose stratigraphic meaning
<figure><img src="image_28.png" /><figcaption>**Fig. 27.** Denisova Cave: site plan and spatial distributions. Provenience of (a) the human remains that have been directly dated or have provided genetic evidence allowing assignment to (or relation with) a specific human lineage, (b) the stratigraphically provenanced and directly dated animal tooth pendants and bone tools, and (c) the charcoal dates for the South Chamber (caption: a. sector number; b. dripline; c. Holocene surface; d. top of exposed Pleistocene deposit; e. excavation trenches). The area excavated in the South Chamber since 2017 has been added. After Douka et al. (2019) and Essel et al. (2023), modified.</figcaption></figure>

changed through time, as the excavation of the site progressed?

Using comparative typology, we can nonetheless conclude that the stone tools from basal level 11 of *Derevianko and Shunkov's (2003)* monograph point to a human occupation of the site taking place in the range suggested by (a) the charcoal dates associated with Kara-Bom's UP1 refit cluster, (b) the cut-marked gazelle bone from Kamenka, and (c) most of the dates for layer 4 of Tolbaga. That range is the 39–42 ka ago interval during which the Early Aurignacian and the Ahmarian/Protoaurignacian are to be found in Europe and the Near East. With regard to the pendants and the bone tools, however, *Douka et al. (2019)*, based on direct dating results, have argued that, at Denisova, the production of such types of artefacts began 43–49 ka ago and may well have been the work of neither Neandertals nor Moderns but of Denisovans. An alternative interpretation is *Zilhão's (2007)* argument, based on *Derevianko and Shunkov's (2003)* rendering of the site's stratigraphic layout, that (a) "a major discontinuity separates (...) layer 11 of Denisova from the immediately overlying (...) level 9," that (b) "the contact between the two is significantly disturbed," and, therefore, that, (c) "because the range of ornaments from level 11 is identical to that found in both level 9 and the pockets containing level 9 lithics that penetrated deeply into level 11 (...), their association with the IUP is questionable."

The implication of this reasoning was that most, if not all the pendants and bone tools from *Derevianko and Shunkov's (2003)* level 11 are of a later Upper Palaeolithic age — probably in the range suggested by the artefacts in their level 9, i.e., in the range of the Gravettian of Europe and western Russia.

In making their case, *Douka et al.* would seem to have overlooked the significance of the charcoal ages that they also obtained for the 2016 excavation of “level 11” of the Main Chamber. They report that the samples come from freshly excavated profiles (mostly from the south-east wall of the 2016 excavation trench) and “from locations that were deemed during excavation to be undisturbed.” The 14 determinations made on 10 different samples using a range of pretreatment techniques, including the more robust ABOx-SC method, are consistent and therefore can be used to define the time span during which the deposit was laid down: between 32,530 ± 260 BP (OxA-34721) and 36,300 ± 900 BP (OxA-X-2695-23), i.e., within the 36–42 ka cal BP interval. Against this background, any older dates for artefacts and bone samples must reflect residuality, the more so if the stratigraphic integrity of the area of the cave (or the part of the deposit) that they come from cannot be ascertained. That residuality is an issue here is otherwise demonstrated by the faunal dates for subunit 11.4 listed in *Douka et al.'s* set of 2016
<figure><img src="image_29.png" /><figcaption>**Fig. 28.** Denisova Cave: The G>D/5–6 profile of the South Chamber. Left: drawing, after the 2003 excavation. Middle: photo, after the 2003 excavation. Right: photo, in 2016, after sampling for OSL dating (the blue dots correspond to samples collected in 2012, the yellow dots to samples collected in 2016). After Jacobs et al. (2019b), modified. The dashed lines indicate points common to both photos, added to facilitate the comparison. The rectangle in the 2016 photo bounds the area containing the samples that Supplementary Table 12 of Jacobs et al. places in layer 12, at an elevation that is consistent with their assignment of samples 4 and 5 to layer 14, and of sample 6 to layer 22. Until at least 2003, this area was considered to belong in layer 11, and it remains so in the geological description found in Jacob et al.’s Supplementary Table 18.</figcaption></figure>

samples from the same chamber. Whereas results in full agreement with those on charcoal were obtained for the four bones of taxa in the size range that, based on Kamenka, one might expect to have been the main target of the human hunters (*Capra* sp. and *Ovis/Capra*), infinite ages, >50,300 BP, were obtained for two bones of *Bos/Bison* that, based on Turner et al., one might expect to reflect the activity of Middle Palaeolithic hyaenas.

### 5.5.2. *The artefacts’ direct dates*

Two of Douka et al.’s (2019) artefact dates are for bone points from “level 11” of the Main Chamber. Both are at odds with expectations derived from the dates on the charcoal and the bones of the caprines from the same level and chamber: one, for an item made on a horse bone and retrieved in 2016, is too old by some five millennia; the other, for an item made on a bison bone and retrieved in 1997, is too old by some ten millennia (Fig. 27). As Kuzmin and Keates (2020) have remarked, it remains to be demonstrated that these are not just naturally pointed, hyaena-digested shaft fragments, which are known to sometimes mimic the morphologies produced by human modification (Villa and d’Errico, 2001). But, if artefacts indeed, these two objects are akin to the minimally modified bone tools of the European Middle Palaeolithic. They could derive from the underlying strata, or they could simply reflect that the raw material was procured by the cave’s Upper Palaeolithic occupants from among the abundant bones scattered across the coeval cave floor, which must have included material of Middle Palaeolithic age displaced by hyaena burrowing.

The directly dated tooth pendants are four: one is a perforated tooth of moose (*Alces alces*) from the Main Chamber, was collected while profile cleaning, is of uncertain stratigraphic provenience, and was dated to 28,390 ± 330 BP (OxA-34729); another perforated moose tooth is from the East Chamber, comes from “level 11” of 2014, and was dated to 41,300 ± 2400 BP (OxA-30963); the other two pendants are perforated red deer teeth from “level 11” of the East Chamber, were collected in 2007 and 2006, and have been dated to 27,820 ± 340 BP (OxA-30006) and to 35,400 ± 900 BP (OxA-30005), respectively. These four results are on non-ultrafiltrated collagen but there is no reason to

suspect that they are minimum ages only because Douka et al. (2019) obtained statistically identical results when dating, with and without ultrafiltration, sub-samples of the same bone sample. Zilhão’s (2007) suggestion that most of Denisova’s “level 11” ornaments relate to human use of the cave in post-IUP, most likely Gravettian times, would therefore seem to be vindicated; indeed, such is also the range indicated by the charcoal dates that, in the South Chamber, have been suggested to provide an age bracket for the perforated red deer tooth recently published by Essel et al. (2023) (Fig. 27).

With regard to the two older results for the tooth pendants, OxA-30005 is within the range to be expected based on the charcoal dates for the parts of “level 11” of the Main Chamber that are claimed to be undisturbed; considering its 95.4 % probability interval, so is OxA-30963. Note, however, that, according to Jacobs et al. (2019b), the number of moose remains in Denisova’s identified mammal bone assemblages is, all layers combined and excluding small animals, (a) zero among the Main Chamber’s 1412 remains, and (b) four among the East Chamber’s 8003 remains (all four coming from different subunits of “level 11;” no body part given, presumably including at least the perforated tooth from subunit 11.2 that was directly dated). With regard to the South Chamber, moose is only represented, and by a single find, in level 16 of the species list published by Vasiliev et al. (2021) for the recent, 2019–2020 excavations (totalling >100,000 bones, of which >6000 identified), while the other taxon whose perforated teeth were directly dated, red deer, is absent from level 11.

Given these numbers, the possibility must be entertained that OxA-30005 and OxA-30963 reflect Gravettian-age usage of scavenged material, either obtained on-site from hyaena-scattered remains of an earlier age or available in the cave’s environment (d’Errico et al., 2024). Bearing in mind the dramatic oscillations that the climates of the northern hemisphere experienced during MIS 3, one would expect erosion and melting of the permafrost to have provided multiple opportunities for Siberian hunters to obtain rare (and thereby prized) items from well-preserved, freshly exposed carcasses of long-dead animals — much as with mammoths and mammoth ivory in the present-day. This caveat — when dating osseous material, one is dating the time of death
of the animal the bone or the tooth belonged to, not the time of manufacture of the artefact that such raw material was transformed into — is the more important in the case of Denisova because (a) the “level 11” assemblage of symbolic artefacts includes items that, typologically, are strongly suggestive of a more recent, even Holocene, age, namely, the polished green stone bracelet published by Derevianko et al. (2008), and (b) the age estimates derived from the human and deer DNA extracted from the surface of the pendant published by Essel et al. (2023) and from the sediment adhering to it do not exclude a relatively recent Holocene chronology either.

Thus, as is the case with the other Siberian sites, Denisova has yielded no reliable evidence for the Upper Palaeolithic to have emerged in central and eastern Asia prior to ~41.5 ka cal BP. Douka et al.'s (2019) speculation that the site's pendants and bone tools may have been made by Denisovans is based, presumably, on the genetic determination to that lineage of two "level 11" remains — the Denisova 4 molar and the Denisova 3 hand phalange. However, (a) neither has been directly dated, (b) the level is ambiguously defined, heterogeneous, and it also yielded the Denisova 5 foot phalange and the Denisova 15 long bone fragment, both genetically determined as Neandertal, and (c) Denisova 15 has been directly dated to >50,200 BP (OxA-36012) (Fig. 27). In all likelihood, therefore, the association that Douka et al. rely on is incidental, and the makers of Denisova's earliest symbolic items are neither Neandertals nor Denisovans but the regional equivalent of western Eurasia's post-admixture, genetically and morphologically modern or near-modern people. In eastern Asia, such people are represented by the Tianyuan 1 partial skeleton from China, directly dated to 34,430 ± 510 BP (BA-03222), i.e., to 37.9–40.7 ka cal BP (Fu et al., 2013; Shang et al., 2007).

## 6. What's to be done?

Modern/Sapiens and Archaic/Neandertal are concepts whose definition is based on the skeleton and on features fixed by natural selection and drift, ones that change over geological time, while the genomic history of the Transition is based on inferences relating to lineage and to changes in the genomes that take place over generational time. The interchangeable use of Neandertal and Modern as categories that are valid in both realms, Human Palaeontology and Genetics, assumes that the differences observed in the latter can be mapped onto differences observed in aspects of the skeleton or the behaviour. Yet, they cannot (Ackermann et al., 2019; Colagè and d'Errico, 2020). This is the fundamental paradigmatic obstacle hindering significant advances in the modelling of what happened in northern Africa and western Eurasia between 35,000 and 45,000 years ago.

The genomes of all European fossils reliably dated to this time interval are consistent with (a) their belonging in the Neandertal lineage, as indicated by skeletal morphology or, (b) if diagnosed as genetically Modern, their having had Neandertal ancestors less than ten generations back in their family history, in some cases no more than four to six (i.e., with their Neandertal ancestors having lived no more than a couple of centuries earlier). If those genetically "modern" people and their genetically "neandertal" ancestors were both alive within a 200-year bracket encompassed by the two-millennial duration of a given technocomplex, and if their descendants are of a mosaic nature, both anatomically and genetically, then discussing the Transition and the associated process of Neandertal assimilation in the dichotomic framework of the 1980s and 1990s is, by definition, a non-starter.

Elements of the traditional framework, namely the chronozo graphic structure of the Transition interval, retain validity, but others have lost the relevance that they once had. For instance, in illo tempore, demonstrating that the Châtelperronian predated anatomically modern humans in western Europe, whether the latter were identified based on diagnostic skeletal remains or based on artefactual index fossils, could and did play a major role in the falsification of models of recent human evolution such as ROA and Acculturation — ones that, as Villa and

Roebroeks (2014) aptly put it, suffered from the “modern human superiority complex.” These days, however, we know that the evidence for Neandertal symbolic thinking goes back to tens of thousands of years before the Transition. In this updated context, debating whether the Châtelperronian’s personal ornaments stand for independent invention or for the influence of coeval, putatively modern human-associated technocomplexes no longer carries implications for the Big Issue question, i.e., whether the Neandertals were a distinct species and cognitively not as modern as their African contemporaries. Yet, the notion that “behavioural modernity” and Neandertal-ness are somehow incompatible continues to inspire, explicitly or implicitly, persistent attempts at assigning to “modern humans” the authorship of the Transition’s technocomplexes, or at least the precedence of “modern humans” in coming up with the innovations that those technocomplexes stand for — and this not just with regard to the Châtelperronian but now also with regard to the Bachokirian and the Ranisian.

Yet, what the genetic evidence from Bacho Kiro and Ranis actually does is add to the obsolescence of the 19th-century mindset whereby (a) the Linnaean names given to human fossils stand for discrete entities and true biological species, (b) a difference in morphology necessarily implies a difference in cognition and behaviour, and (c) one-to-one correlations should be expected to exist between sets of skeletal features (or of genes) and a given material culture or given cultural traits. If the people of the Bachokirian and the Ranisian are eventually shown to have been anatomically modern humans of very recent non-European origin, the fact that their personal ornaments are of a Neandertal/European tradition would be an excellent illustration of the fact that, when it comes to understanding cultural variation and population dynamics over the last 100,000 years, Neandertal and Modern are categories that have outlived their use-by date. Instead, what we are seeing ever more clearly is a dynamic and fluid landscape of interaction whereby genes and cultural traits spread across the Old World regardless of the artificial boundaries created by the two bad habits of (a) using Linnaean taxonomy to frame variation among ancient human populations, and (b) taking categories of modern cartography to frame the realities of Ice Age Human Geography.

Unlike the original one, the resurrected Acculturation Hypothesis can now be assessed thanks to the progress made by Genetics over the last two decades, which makes it possible to ask about Upper Pleistocene humans the questions that truly are fundamental here: Does it make any sense to posit that an individual's cultural identity is determined by their genetic make-up and physical appearance, the more so when, such as with Denisovans, Bachokirians, and Ranisians, said physical appearance is entirely unknown? And can we really assume that the cultural self-identification of Bachokirian and Ranisian communities would have been based on how much of the genomes of each of their individual members would have been inherited from ancestral lineages?

Consider the fact that the Denisova 11 individual has been shown to be the female child of a Neandertal mother and a Denisovan father (Slon et al., 2018): Was that person a “Neandertal,” or a “Denisovan?” And even if the question had any sense in terms of lineage history, does whatever we might want to call the child have any implications for our understanding of the period’s cultural, social, and family life? Likewise, what do staunch dichotomists propose that we call the father, or the mother, of that grandson or great-grandson of a Neandertal who died in the Banat mountains of Romania some 40,000 years ago?

Furthermore, in what sense are the Neandertals not directly ancestral to the people of the Transition if, whenever we are able to peer into the latter's genomes, we realise that every single one of them had had fully Neandertal grandparents or great-grandparents? And, at a population level, what kind of an "extinction" is one whereby people of the "replaced" population/species are to be found in the immediate ancestry of every single individual of the "replacing" population/species? And, even closer to issues that lie within the realm of archaeological enquiry, can we really assume that when two populations mix, the material culture will automatically undergo change, and that said change will
necessarily consist of adopting the culture of the side whose genes eventually predominate? In fact, is there any basis to assume that the material culture of the genetically distinct populations whose admixture is documented at Oase, Bacho Kiro, and Ranis was itself also distinct in any archaeologically visible way?

More to the point, would reducing such fragmentary material as the Bacho Kiro and Ranis remains to the Neandertal/Modern dichotomy even be attempted if the classification were not considered to impact our perception of the dichotomy's poles in terms of their putatively species-specific behavioural capabilities and achievements? We posit that it would not. We posit that, absent that framework, the fossil remains of the Transition, if incomplete and fragmentary, would simply be treated as a chronologically intermediate sample useful in the assessment of (a) how aspects of skeletal and dental morphology changed between the early and the late Upper Pleistocene, and (b) what such change means in terms of the long term co-evolution of body, mind, and culture.

Clearly, the Transition was a time of complex processes involving migration, local extinction, replacement, avoidance, interbreeding, and assimilation. In western Eurasia, the start and end points of the process are clear (Holliday, 2023): prior to about 45,000 years ago, its inhabitants are of the Neandertal phenotype, and their lithic technology is of the Middle Palaeolithic kind; after 35,000 years ago, its inhabitants are of the Cro-Magon phenotype (albeit unevenly displaying, at least in south-west Europe, Neandertal-inherited features, such as the low crural indexes of the Lagar Velho child and of two of the Cro-Magon adults; Trinkaus et al., 2022; Zilhão and Trinkaus, 2002) and their lithic technology is of the Upper Palaeolithic kind. In between, many things happened, starting with the appearance in western Siberia, no later than ~45 ka ago, of people with a non-Neandertal post-cranial skeleton, as represented by the Ust'-Ishim femur (Fu et al., 2014). Namely:

* Given the implications of femoral shape for the body's proportions and bauplan, it is likely that the Ust'-Ishim people had recent ancestors in lower latitudes, e.g. the Arabian Peninsula or north-eastern Africa — where, however, skull-wise, the coeval people were not quite modern, as exemplified by the Nazlet Khater 2 individual.

* So far, no unambiguous evidence has been found as to the behavioural and material culture manifestations associated with the demic processes involved in the emergence of the Ust'-Ishim pattern — in Siberia and central Asia, the earliest Upper Palaeolithic is several millennia later and of broadly the same age as the Protoaurignacian/Ahmarian and the Early Aurignacian of Europe and the Near East.

* The genomes derived from undiagnostic, fragmentary human remains of eastern and central Europe dated to the 40–45 ka ago interval suggest increased gene flow, including migration, from populations that, genetically, were distinct from the Neandertals living in humanity's northernmost frontier (the periglacial belt adjacent to the ice caps) — but such genomes do not exclude that, morphologically, those populations could have been Neandertal-like too.

* If immigrant communities with a very recent origin in Africa are indeed involved in these processes, the consistent presence in the genomes of their descendants of a significant, recent contribution from the northern Neandertals shows that such immigrants admixed extensively with the locals, everywhere and as a rule.

* The Oase fossils show that the people we find in eastern Europe during the 40–45 ka ago interval were, skull-wise, of an archaic/modern, mosaic anatomy, but, because we still know nothing about their post-cranial skeleton and body proportions, we can't tell whether the reason lies in that (a) such was the “type” of their Nazlet Khater-like putative ancestors, (b) is what, anatomically, ought to be expected from their mixing with local Neandertals, or (c) both.

* This moment in time marks the threshold for the pattern of intermittent intercontinental gene flow of preceding times to have given rise, over a relatively short period of intense interaction, to the emergence of a fully unified, pancontinental gene reservoir, and to

the attendant merger (i.e., Assimilation) of the regionally distinct populations of the past into the “anatomically modern human” physical pattern since extant worldwide (for a concurrent view from the side of Genetics, see Iasi et al., 2024; for a largely concurrent view from the side of Archaeology, see Greenbaum et al., 2019).

* The ultimate cause of such increased levels of interaction must lie in the significant demographic growth implied by the novel ritual behaviours (burial, personal ornamentation, cave painting) seen in the archaeological record of the Last Interglacial and the early Last Glacial, which bespeak of the complexification of social relations to be expected in a scenario of rising population numbers, and this in Africa as much as in Eurasia (where additional evidence to that effect is provided by the significant increase in the number of Middle Palaeolithic sites of MIS 3 rather than MIS 4 or MIS 5 age) (Zilhão, 2015, 2020).

* Starting at about the same time, ~45 ka ago, the Levallois method is altogether abandoned, and lithic production systems that are based on the reduction of prismatic cores and target the production of slender blanks (blades and bladelets) become ubiquitous, with such a widespread process probably reflecting the rise to dominance of weaponry systems based on lighter projectiles with a longer range.

* Such lithic production systems have deep roots in the early Upper and late Middle Pleistocene of Europe, the Near East, and Africa, and neither their earlier developments nor their ~45 ka ago blooming bear specific relation to any one of the various taxonomic entities of Human Palaeontology.

* In the Levant, the emergence of new lithic production systems, exemplified by the IUP (in the sense of a “Boker Tachtit level 4 industry”), lacks clarity with regard to the identity of their creators.

* In central and eastern Europe, similar systems characterise the Bohunician and Ranisian cultures; no human remains are associated with the former, while the Ranisian of the eponymous site is associated with undiagnostic remains that, genetically, differ from those of the previous Neandertal inhabitants of Eurasia's steppe-tundra belt (who, as shown to have been the case with the Bachokirians of Bulgaria, may nonetheless have also been among the Ranisians' ancestors).

* In western Europe, the range of the Ranisian extends into southern England, encompassing Belgium, where diagnostics of the technocomplex co-occur at the site of Spy with diagnostic Neandertal remains directly dated to many centuries after the technocomplex's start date.

* In south-western Europe, the Châtelperronian is associated with Neandertal skeletal remains within the same archaeological site or stratigraphic unit, indicating, directly or indirectly, that this cultural tradition represents an indigenous Neandertal development.

* The innovations in the realm of lithic technology are parsimoniously explained by convergence, but cultural diffusion is indicated, ~41.5 ka, by the addition of beadworks crafted from small marine gastropod shells to Neandertal Europe's record of body decoration items, which is documented since at least 115,000 years ago both directly (by pendants made of large bivalve shells, raptor talons, fossils, bones, and teeth) and indirectly (by the zooarchaeological evidence for the procurement of feathers).

* The data remain consistent with the persistence, all along, of a Neandertal, Middle Palaeolithic isolat in southern and western Iberia, but, eventually, no later than 37 ka ago, the interaction dynamics of preceding millennia gave rise to a pattern of cultural and genetic homogeneity across vast swathes of Eurasia, from Portugal to Siberia, as reflected in the geographical extent of the Gravettian and the Cro-Magnon people, a.k.a., the “Hunters of the Golden Age” (Roebroeks et al., 2000).

Processes of population turnover involving the regional extinction of entire genetic lineages were involved in the neolithisation of parts of Europe and in the spread of the Bell Beaker culture to the British Isles
(Allentoft et al., 2024a; Olalde et al., 2018). For instance, Allentoft et al. found that the genetic contribution of people of the regional Mesolithic to subsequent early farmer populations was <10 % in Denmark, and zero in Sweden, i.e., inferior to the contribution of regional Neandertals to the ancestry of the Romanian early Upper Palaeolithic people represented by the Oase 1 individual. This evidence makes it clear that the Neandertals' participation in the genetic make-up of the post-admixture peoples of Europe was fully within the range of what to expect based on patterns of interaction between recent human populations with distinct ancestries but that, by definition, belonged to the same biological species. Also consider that, (a) morphologically, most of the individuals from the Transition dubbed "Homo sapiens" in the ROA sense of the concept in fact differ from recent humans in several aspects of their skeletons, and (b) genetically, their contribution to the ancestry of living people was as limited as the Neandertals' (Zeberg et al., 2024). Yet, these observations have led no one to conclude that those individuals belonged in a now-extinct species. We see no reason to view coeval Neandertals differently.

That ancient genomics has been able to shed much light on Holocene population dynamics is largely due to an abundance of samples. The processes of biological and cultural interaction that took place in recent prehistory are thereby much better understood, and ought to provide the adequate source of inspiration for modelling the Transition — the more so given the paucity and limited geographical scope of the data available to study the latter. As we hopefully have shown, it is in any case clear that the spatial and temporal patterning of the events and processes involved is irreducible to the Rapid-Replacement-of-One-Species-by-Another worldview. The latter's scenarios of competitive exclusion whereby a "for-whatever-reason-inferior" species was not just driven to but in fact doomed to extinction are mostly imaginary: individuals are reduced to one or the other of two essential types, and populations are mapped onto technocomplexes modelled as abstract, bio-behaviourally homogeneous entities. This rationale then leads to explorations of the period's complex dynamics that would resonate with Plato but not with Darwin and provide good examples of the seductive and enduring power of simplifying complexity into binary oppositions. That the practice remains widespread is otherwise a clear manifestation of the so-called anchoring problem (Kahneman, 2011) — a by-product of the entrenched framework that King's (1864) definition of the Neandertals as *Homo neanderthalensis* set for our understanding of ancestral human populations.

To be fair, we must note that a remarkable contrast exists between current attitudes to recent human evolution prevailing among archaeologists and biological anthropologists, on one hand, and geneticists, on the other. The latter were the driving force behind the rise to dominance of ROA but, these days, one hardly finds a Genetics paper that does not frame the issues in terms of populations and lineages. Yet, like so many archaeologists and biological anthropologists, most geneticists remain subject, despite the weight of their own evidence, to the "tyranny of the discontinuous mind" (Dawkins, 2017; Zilhão, 2019). Most continue to accept and operate within the dichotomic framework of the "Neandertal versus Modern" and the "Timing of the Split in the Tree" approaches; few are those (e.g., Tournebize and Chikhi, 2023) who manage to breach the shackles imposed by the categories of Human Palaeontology to embrace uneven geographical development and population structure as the foundations of their enquiry.

In the manner that the Transition is treated under ROA we can also see how History often gives way to Teleology, as is readily apparent in the tendency to approach the period not as something worthy of study in its own right but rather as a chain of events featuring present-day humans as its Aristotelian final cause. This finalist approach in turn leads to circularity: the limited contribution that Neandertals made to recent human genomes and the disappearance of their phenotype is taken as evidence of their having been a separate, extinct species; and this conclusion is then used retrospectively as the assumption upon which to ground the interpretation of what happened at the time of their

disappearance. Yet, it should go without saying that, when studying a given present, causality can only be found in the past of that present, not in its future. The point with the Transition is not how much and what did today's people inherit from the Neandertals, the point lies in (a) how much and in what manner did the phenotypically Neandertal populations of western Eurasia participate, 40–45 ka ago, in the making-up of the biology and culture of succeeding populations, and (b) what variation there was across time and space in the manner that that process unfolded in the different regions concerned.

## 7. Conclusion

The widespread adoption of the taphonomic perspective when dealing with the associations and the dating of the fossils has brought immense clarification on key issues relating to the early stages of human evolution, as exemplified by the study of the Trinil sites — a model on how to deal with old excavations — or the way deposition and redeposition are adequately dealt with in the context of the origin of stone tools in Africa (e.g., Archer et al., 2020; Hilgen et al., 2023; Pop et al., 2023). The same cannot be said about the Transition, a field where little progress has been made in the incorporation of the lessons that “The Neandertal Problem” debate ought to have taught us: definitions continue to be abused; misunderstanding of dating records remains widespread; diversion of the Bayesian approach to support exercises in the affirmation of the consequent is commonplace; treatment of stratigraphic units as the sealed museum drawers that they are not seems to be unstoppable; the genetic evidence for the Neandertal ancestry of Upper Palaeolithic humans is paid lip-service but otherwise largely ignored; and most interpretations of the archaeological record remain fixed in pre-1960s reified visions of material culture as the by-product of monolithic bio-behavioural entities. We posit that the explanation for this state of affairs lies in the obsolete dichotomic framework that continues to inspire much Transition research. The time to move on is overdue.

## Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

## Acknowledgments

JZ acknowledges funding provided by FCT (Fundação para a Ciência e Tecnologia, Portugal; grants UIDB/00698/2020 and UIDP/00698/2020). FD and WEB acknowledge the Grand Programme de Recherche 'Human Past' of the Initiative d'Excellence (IdEx) of the University of Bordeaux. FD acknowledges the European Research Council (Synergy Grant for the project "Evolution of Cognitive Tools for Quantification – QUANTA," no. 951388), the Research Council of Norway through its Centres of Excellence funding scheme, the SFF Centre for Early Sapiens Behaviour (SapiENCE), project number 262618, and the Talents Programme of the University of Bordeaux (grant number: 191022_001). Erik Trinkaus, John Willman, Lounès Chikhi, and Paul Bahn read earlier versions of the draft and provided useful comments and insight. We thank Fabio Martini and Sahra Talamo for useful information about the stratigraphy and dating of Grotta del Cavallo and Roccia San Sebastiano, and we acknowledge the editor and reviewers for their useful criticisms and suggestions, which served to improve the original submission. We also wish to acknowledge Clément Zanolli for clarification on the premises under which the Mandrin layer E tooth was studied, based on which we have been able to rectify the corresponding passage of the uncorrected on-line manuscript. As usual, any errors or omissions are our own.
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Zwyns, N., Lbova, L.V., 2019. The Initial Upper Paleolithic of Kamenka site, Zabaikal region (Siberia): A closer look at the blade technology. Archaeol. Res. Asia 17, 24-49.

Zwyns, N., Paine, C.H., Tsedendorj, B., Talamo, S., Fitzsimmons, K.E., Gantumur, A., Guumii, L., Davakhuu, O., Flas, D., Dogandžić, T., Doerschner, N., Welker, F., Gillam, J.C., Noyer, J.B., Bakhtiary, R.S., Allshouse, A.F., Smith, K.N., Khatsenovich, A.M., Rybin, E.P., Byambaa, G., Hublin, J.-J., 2019. The Northern Route for Human dispersal in Central and Northeast Asia: new evidence from the site of Tolbor-16, Mongolia. Sci. Rep. 9, 11759.

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