# Tooth in the spotlight: Exploring the integration of archaeological and genetic data to build multidisciplinary narratives of the Past

Francesco D'errico, Luc Doyon, João Zilhão, Jack Baker

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Francesco D'errico, Luc Doyon, João Zilhão, Jack Baker. Tooth in the spotlight: Exploring the integration of archaeological and genetic data to build multidisciplinary narratives of the Past. Comptes Rendus. Palevol, 2024, 23 (2), pp.31-43. 10.5852/cr-palevol2024v23a2. halshs-04410445

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Une dent sous les feux de la rampe : Explorer l'intégration des données archéologiques et génétiques pour construire des récits multidisciplinaires du passé

Tooth in the Spotlight: Exploring the Integration of
Archaeological and Genetic Data to build Multidisciplinary
Narratives of the Past

Authors

d'Errico, F.<fnref n="1" />,<fnref n="2" />, Doyon, L.<fnref n="1" />,<fnref n="3" />, Zilhão, J.<fnref n="4" />, Baker, J.<fnref n="1" />

Corresponding author:

<footnote n="1">¹ Université de Bordeaux, CNRS, MCC, PACEA, UMR5199, Pessac, 33615, France.</footnote>

<footnote n="2">² University of Bergen, Centre for Early Sapiens Behaviour (SapienCE), Department of Archaeology, History, Cultural Studies and Religion, Bergen, 5020, Norway. Email: Luc.doyon@u-bordeaux.fr</footnote>

<footnote n="3">³ Shandong University, Institute of Cultural Heritage, Qingdao, 266237, China.</footnote>

<footnote n="4">⁴ UNIARQ, Centro de Arqueologia da Universidade de Lisboa, Faculdade de Letras, Universidade de Lisboa, 1600 - 214 Lisboa, Portugal.email: joao.zilhao@gmail.com</footnote>
Résumé

Cet article aborde les défis posés par la collaboration interdisciplinaire qui a conduit au récent article de *Nature* présentant les résultats de l'extraction d'ADN humain ancien à partir d'une canine de cervidé perforée trouvée dans la grotte de Denisova. Nous proposons ici une analyse critique des données contextuelles qui a un impact direct sur l'interprétation des données génétiques et l'évaluation du potentiel de la nouvelle technique d'extraction de l'ADN pour les études futures. Nous approfondissons la discussion en particulier pour Denisova, seul site qui a produit des données paléogénétiques exploitables. A travers une synthèse des précédentes publications et des données récentes, nous mettons en évidence des incohérences de datation non résolues sur le site, qui nécessitent une explication. Nous soulignons également l'absence d'un cadre théorique approprié dans l'analyse de la dent perforée, le manque d'informations sur sa latéralisation, le sexe, l'âge de l'animal sur lequel la dent a été prélevée, la technique de perforation de la dent, l'usure et l'analyse des résidus, ainsi que la diffusion d'informations erronées sur l'assemblage faunique associé. Nous concluons que des données archéologiques de qualité et leur évaluation rigoureuse devraient compléter les données paléogénétiques afin de créer des récits unifiés, fiables et vérifiables sur le passé humain.

Abstract
The article addresses the challenges posed by the interdisciplinary collaboration that led to the recent *Nature* article presenting results of the extraction of ancient human DNA from a perforated deer tooth found in Denisova Cave. We provide a critical analysis of the contextual data that directly impacts the interpretation of the genetic data and the evaluation of the potential of the new DNA extraction technique for future studies. We deepen the discussion especially for Denisova, the only site that has produced truly exploitable palaeogenetic results. We highlight major unresolved dating inconsistencies at Denisova that need inquiry and explanation. We also emphasize the lack of an appropriate theoretical framework in the analysis of the tooth, of information on the lateralization, sex, age, perforation technique, use wear, on the ochre, and the release of erroneous information on the associated faunal assemblage. We conclude that quality archaeological data and their sound evaluation should complement paleogenetic data to create unified, reliable and verifiable narratives about human history.

## Mots-clés

Paléogénétique, perles, objets de parure, grotte de Denisova, Paléolithique supérieur, datation 14C

## Keywords

Paleogenetics, beads, personal ornaments, Denisova Cave, Upper Palaeolithic, 14C
dating

## Introduction
One can only be stunned by the technical achievement highlighted by the recent article by Essel and colleagues (2023). Palaeogenetics has advanced by great strides over the last three decades, including the extraction of DNA from isolated ancient fossil remains (Meyer et al. 2012; Pääbo 2014; Prüfer et al. 2014), the reconstitution of the genome of a fossil human population (Hajdinjak et al. 2018), the identification of hybridisation phenomena between fossil populations (Fu et al. 2015; Harris & Nielsen 2016; Kuhlwilm et al. 2016; Sankararaman et al. 2016; Hajdinjak et al. 2018), and the study of the consequences of hybridisation for the epidemiology of current human populations (Silvert et al. 2019; Koller et al. 2022; Reilly et al. 2022). Of late, achievements have concerned the extraction of DNA, including human DNA, from the sediments of caves and shelters occupied by our ancestors (Slon et al. 2017; Vernot et al. 2021; Massilani et al. 2022), and the production of models for the Upper Palaeolithic peopling of Europe (Posth et al. 2016, 2023; Villalba-Mouco et al. 2023). These developments have revealed the potential of paleogenetics to further our understanding of the evolutionary history of our lineage and the history of its adaptation to past environments. However, these advances seem to pale, at least from a technical point of view, in comparison with the performance of Essel and colleagues, who succeeded in extracting ancient human DNA from a perforated deer tooth found in Denisova cave. They attribute an age of 19,000–25,000 years to the pendant, identify the individual who would have produced or worn it as a woman and recognize in her genome affinities to a group of ancient inhabitants of northern Eurasia who lived during the same period, but who were previously found only further east, in Siberia. Based on their results, the authors conclude that their work redefines how the cultural and genetic archives can be linked to prehistoric archaeology and that it may become possible to systematically combine genetic and cultural analyses
to study the use of Palaeolithic artefacts and to ask questions about task specialization,
e.g., by sex or genetic ancestry.

These findings and statements caught our attention in several respects because they
highlight, in our opinion, certain difficulties that exist in establishing a constructive
dialogue between archaeologists and palaeogeneticists. We feel it is important to
discuss them here because they are different from the concerns raised in the context of
the debates that have animated the dialogue between these disciplines in recent years.
Such debates have focused on issues related to the assimilation of genetics, linguistic
and cultural data and the ethical and epistemological implications of such assimilations
(Pluciennik 2006; Eisenmann et al. 2018; Lebrasseur et al. 2018; Potter et al. 2018;
Hakenbeck 2019; Frieman & Hofmann 2019). A good example is the debate between
geneticists and archaeologists over the interpretation of Bell Beaker pottery as a
signature of an invasion of Europe 4500 years ago by a steppe people speaking an Indo-
European language (Otalde et al. 2018; Anthony 2019). The latest analyses temper this
scenario and show that, while there was a significant population replacement in certain
areas, such as the British Isles (Brace et al. 2019), this was much less the case in other
regions, such as the Iberian Peninsula, where there is strong genetic continuity, which
moderates the link between archaeological cultures and genetics (Szécsényi-Nagy et al.
2017). The trade-off article proposed by the geneticists, aimed at establishing criteria
for matching genetics and material culture (Eisenmann et al. 2018), has triggered a
response from some archaeologists (e.g., Riede et al. 2019) who question traditional
cultural classifications and, therefore, the relevance of linking them directly to the
results of palaeogenetic analyses. Much of the work published on this and similar issues
(Rothhammer & Dillehay 2009; Veeramah 2018; Kristiansen 2022) emphasizes the
importance of interdisciplinary collaboration and communication, and the need for
researchers to be transparent about their methods and assumptions. The authors often
call for increased dialogue between archaeologists, geneticists, and other stakeholders
to ensure that the scientific use of archaeogenetic data is responsible and respectful of
different views and interests. They further argue that archaeogenetic data should be
used in combination with other sources of information, such as archaeology, linguistics,
and historical records, to create a more complete understanding of the past.

Matters of concerns

Here we want to address another, no less sensitive but related aspect of this
interdisciplinary collaboration, which is the role of archaeologists and archaeological
data in collaborations like the one just published by Essel and colleagues. Many
archaeologists, archaeozoologists, palaeoanthropologists, and dating experts, are
among the authors of this work. What was and what should have been their contribution
to the published research? We see two areas of concern that we will develop in this
short contribution.

Logically, the participation of paleoanthropologists in a paleogenetics study is justified
primarily because they have excavated, stratigraphically assessed or dated the sites in
which the analysed objects were found, or because they are responsible for the study
of certain categories of artefacts discovered at these sites that are relevant for the
discussion of the paleogenetic evidence. In short, the archaeologists who participate in
the study should be in the best position, after years of commendable work, to propose
an informed view of the contextual data and thus enable their geneticist colleagues to
interpret their results based on reliable stratigraphic, cultural, and chronological
attributions. Peer-reviewers with archaeological expertise should play the role of
authoritative devil's advocates to identify inconsistencies in or with the evidence
derived from other sites analyzed or discussed in the paper. It is legitimate to question,
however, whether this role of guarantor and critical analyst of background data has been
played satisfactorily here, at least for Denisova, the only site that has produced truly
exploitable palaeogenetic results and on which we will therefore focus our attention.
Admittedly, a chronological attribution of the perforated tooth is proposed in the article
in question based solely on palaeogenetic results, but the margin of error is far too wide
to consider that it frees us from the obligation to assess the chronological setting based
on stratigraphic analysis, cultural attribution and dating of the archaeological layers. Yet,
it is the interpretation of the stratigraphic sequence and the dating of these layers that
would have required a critical appraisal by archaeology co-authors.

The Denisova Cave stratigraphy and dating

According to the archaeologists responsible for the excavation, the perforated tooth was
found at the center of the South Chamber of Denisova, in the upper part of layer 11,
square E3. It is said that the piece was 3D plotted, but the resulting coordinates and the
find number are not provided. It is said that the thickness of layer 11 does not exceed
50 cm in the area in which the tooth was found but ranges in depth from -150 to -230
cm due to a substantial slope toward the southwestern rock wall of the chamber at an
angle ranging from 20–25° to 50°. Such a marked slope, however, would have not
affected the integrity of the layers. The age of the canine is assessed, according to the
authors, by two 14C ages obtained on charcoal samples discovered in the proximity of tooth in layer 11 (OxA-X-3089-11: 19,990 ± 61 BP; 24,200-23,830 calibrated years before present (cal BP) and (OxA-X-3089-12: 33,500 ± 220 BP; 39,180-37,560 cal BP) at 95.4% probability. It is however explained that the first dated charcoal was collected as much as 1.5 m away from the tooth and comes from the middle rather than from the upper part of layer 11, in which the tooth was found, and the second sample comes from the lower part of layer 11. This means that neither of the two 14C ages comes from the portion of layer 11 in which the tooth was found. Neither of them can be claimed to provide an estimate on the age of the tooth. This is more so considering that a 15,000-year difference between the two ages in a less than 50 cm thick sedimentary sequence suggests the layer is far from being chronologically homogenous. In sum, in absence of more 14C determinations and precise depths for the samples, not provided in the article, it is difficult to draw any reliable inference on the age of the tooth based on the reported 14C evidence apart from proposing that the tooth should be probably younger than the 14C age obtained on the charcoal sample from the middle portion of layer 11.

More 14C ages, many of them published in the same journal, are however available for the Denisova sequence and layer 11 that, surprisingly, have not been considered by Essel and colleagues when discussing the age of the tooth. Several striking inconsistencies emerge when putting the chronological attribution proposed by Essel and colleague in the context of previous radiometric determinations and ensuing narratives. Four other 14C ages are available for layer 11. Reich *et al.* (2010) reported three 14C ages on bones from the lower part of Layer 11, including two >50,000 BP (OxA-V-2359-17 on an Ovis/Capra bone and OxA-V-2359-18 on a Bison bone from layer 11.2), and one on a Hyena bone that produced an age of 48,650 ± 2,380 BP (KIA-25285). The fourth age was
on charcoal, comes from the interface between layers 11 and 10 and produced an age
of 29,200 ± 360 BP (AA-35321). In their 2010 article, Reich and colleagues interpreted
the discovery of a Denisovian molar in the upper portion of layer 11, the same one in
which the perforated tooth was found, and the four just mentioned associated 14C ages
as the proof that Denisovans inhabited the cave for more than 50 ka and that this
frequentation was followed, around 30 ka by another, probably by modern humans. The
presence of Denisovians at the site during the accumulation of layer 11 was later
supported by the palaeogenomic proximity between the Denisova 4 molar and the
Denisova 3 phalanx found in the same layer, in the East Chamber (Douka et al. 2019).

This summary leaves us with several important questions that we would have expected
Essel et al. to discuss. To what do the authors attribute a discrepancy of at least 15,000
years between the age they obtained for the lower part of Layer 11 and the 14C ages
previously obtained for this same part of the layer? How do they explain a 10,000-year
inversion between the age they obtained for the middle part of Layer 11 and the age
previously attributed to the interface between this layer and Layer 10? What
conclusions do they draw from the fact that, apparently, the upper part of layer 11 has
yielded both Denisovan remains and a pendant impregnated with modern human DNA?

From a cultural standpoint, two subunits are distinguished in layer 11. The first, found
in the lower and middle part of layer 11, comprises an Early Upper Palaeolithic stone
tool industry characterized by parallel knapping aimed at producing elongated flakes,
large blades and small bladelets. The second, found in the upper part of layer 11,
presents a high concentration of small bladelets and microblades, including backed
pieces. Interestingly, most of the non-utilitarian objects, i.e., beads, pendants, and
ornaments made of animal teeth, bone, mammoth tusk, soft stone, ostrich eggshells and freshwater shells, and bone tools, such as bone eyed-needles, points, polishers, awls, etc., were found in the lower part of layer 11 (Essel et al. 2023).

When describing the pendant, Essel and colleagues point out that pendants of this type are common in the Initial and Early Upper Palaeolithic at Denisova Cave as attested by some specimens found in the East Chamber, layers 11.2 and 11.1, and directly dated to 41,900–38,400 years BP (OxA-30005) and 32,660–31,100 years BP (OxA-30006). In the original article reporting these ages, the authors cautioned that the determinations should be considered minimum ages owing to the small size of the objects and the marginal collagen yields, ranging between 1.0 and 1.6 w% collagen, which prevented the application of robust chemical pre-treatment methods (Douka et al. 2019). If the ages reported by Essel and colleagues are not underestimated, the cultural relationship between the bearers of the pendants found in the East and the South Chambers becomes an open issue, and the homogeneity and/or stratigraphic integrity of the deposit must be interrogated.

In sum, the stratigraphy of Denisova Cave is extremely complex, and there can be no question that the sequence underwent significant post-depositional disturbance, as originally argued by Zilhão (2007) and since corroborated by the geological work carried out at the site (Morley et al. 2018, 2019; Kuzmin & Keates 2020). Any ground-breaking discoveries would benefit from a critical in-depth appraisal of the sequence to clarify rather than muddle its understanding. Considering the above, the agreement between the proposed dating based on genetic evidence and the new 14C ages published in the paper must be considered as largely artificial and difficult to accept without resolving
the major inconsistencies revealed by the site's dating record. It is the responsibility of
the archaeologists to carry out this critical analysis and to propose interpretations that
are acceptable to the scientific community. The application of new and increasingly
sophisticated DNA extraction techniques cannot solve these problems.

Combining archaeological and genetic results to build a unified narrative

The second question that arises from Essel *et al*. article is why the representatives of the
different disciplines did not set about developing, as far as possible, the dialogue and
cross-fertilisation process that they call for in the conclusion of their article. If the aim is
to combine the results of different disciplines to create a better informed narrative of
our past and not only to show that techniques common in forensic medicine can be
perfected and applied to the Palaeolithic, the integration of archaeological and
palaeogenetic data represents an epistemological imperative, and all the more so when
the results of the research are published in a high ranking multidisciplinary journal such
as *Nature*. A supplementary reason for the need to contrast results was, as will be shown
below, that the inclusion of pertinent archaeological data in the scientific discourse
would have had a direct impact on the interpretation of the genetic data and the
discussion of the potential of the technique for future applications.

The body fluid of the human who wore this pierced tooth left a genetic signature that
was identified by the researchers, but the analysis of this genetic material does not
inform us about the duration of the object's use. We have no way of knowing at present
whether it took a day, a week, or a much longer period of time for this ornamental object
to become impregnated with human body fluid nor whether this process erased
previous traces left by other individuals of the same or the other sex. This question is relevant because the anthropological implications of the information provided by palaeogenetics would not be the same. For instance, in the case of objects worn for a single day or by a single person, how might paleogenetic data be used, as the authors explicitly contemplate in their conclusions, to obtain information on task specialization by sex? One way of approaching the lifespan of an object of adornment, which also provides information on its social life, is to study the wear over its surface and, in particular, the wear of the perforation, the area that is often the most worn due to friction against the thread used to sew the pendant onto a garment or thread it into a necklace, bracelet or diadem. Archaeologists have investigated and experimentally reproduced this wear, and tools exist to document and quantify it (d'Errico 1993; d'Errico et al. 1993, 2022; Vanhaeren et al. 2013; Martisius et al. 2018; Falci et al. 2019).

No steps seem to have been taken in the study to identify these traces and to integrate the results of a functional analysis with those provided by paleogenetics. This is surprising because an analysis of the surface condition of the object, carried out with a confocal microscope, is presented in the article, but it was carried out with the sole aim of showing that the surface of the object had not been substantially damaged by the sampling intended to test for the presence of DNA. The same microscope could have been used, in combination with optical and scanning microscopy, to assess the degree of wear of the perforation and thus contribute with archaeological data to the generation of a unified interpretation of its paleoanthropological significance.

There is ample evidence that adornment objects were exchanged over great distances during the Paleolithic and we know for certain that in some cases deer canines travelled several hundred kilometers and were worn by individuals living in cold regions where
this species was not present (Vanhaeren & d'Errico 2003a, 2005; Rigaud et al. 2019). The
canine in question may have arrived in Denisova following exchanges with populations
living in eastern Siberia and been lost in Denisova before losing the genetic imprint of
the individual that originally made and carried it and before absorbing the genetic
material of the individual that may have carried it to Denisova (figure 1). This is especially
true considering that ornaments were often not worn in contact with the skin as
necklaces or bracelets but sewn onto clothing (Vanhaeren & d'Errico 2001; d'Errico et
al. 2021). The object may therefore have been impregnated with the body fluid of the
individual who handled it during its manufacture rather than the one who wore it as a
result of exchange. Taking this into account, it is problematic to draw conclusions from
the genetic analysis of this object in terms of population dynamics. Creating a dialogue
between disciplines is therefore essential.

One way that archaeology contributes to weighing up such hypotheses is to verify
whether wapiti remains are present in the archaeological layer that yielded the
ornamental object. If absent or rare, several hypotheses about how the canine was
acquired must be considered. For instance: (a) cultural exchange rather than acquisition
from an animal that the cave's inhabitants hunted, (b) scavenging from frozen carcasses
exposed by erosion of permafrosted deposits, (c) on-site acquisition of reworked wapiti
remains (provided the species is present in underlying deposits, as indeed it is; Vasiliev
et al. (2021)). Essel and collegues claim (2023, SI: 5-6) that remains of wapiti (Cervus
elaphus sibiricus) were found in layer 11 and cite Vasiliev et al. (2021) in support of this
statement. That paper, however, tells a different story, namely, that layer 11 is the only
one of the eleven archaeological layers analysed that yielded no wapiti remains.
Considering the large time span covered by the accumulation of this stratigraphically
complex unit this is quite remarkable and strongly supports scenarios where the wapiti
canine is an exotic good acquired by Denisova inhabitants through exchange or a fossil
whose age, regardless of it being established by genetic or radiometric techniques,
differs from the age of the archaeological event of interest (its modification and use as
an item of jewelry).

Essel and colleagues argue that their extraction technique makes it possible to
systematically combine genetic and cultural analyses in the study of Palaeolithic
artefacts. The study of ornaments used by traditional societies shows that they can
provide different types of information to archaeologists. Through the raw material from
which they are made, their morphology, their colour, their association and arrangement
on the body, ornaments make it possible to express, depending on the human group,
the individual's belonging to an ethnic, linguistic or social group or to an age class (Kinietz
1972; Wiessner 1984; Joyce 2005; Micheli 2012; Nowell & Cooke 2021). But Palaeolithic
ornamental objects are also the result of techniques used to transform natural materials
into cultural objects and attach them to an individual's body. It is known that the
materials chosen (e.g., shell, tooth, ivory, antler, bone, eggshell etc.), the animal species
preferred, and the techniques of shaping and attaching, differ among prehistoric human
groups (Vanhaeren & d'Errico 2006; Rigaud et al. 2015; d'Errico et al. 2021; Baker et al.
submitted).

It is therefore necessary to characterise these behaviours when publishing objects of
adornment. This naturally requires adequate photographic documentation, which for an
object as important as this wapiti tooth requires photos of all four sides of the object,
as well as microscopic photos to document and identify the techniques of perforating
the tooth on both sides. Documenting the location of the wear on the perforation is
important in order to know, for example, whether the tooth was part of a necklace or
was sewn onto a garment (Vanhaeren & d'Errico 2001; Vanhaeren et al. 2013; Mårgårit
et al. 2018; Falci et al. 2019; d'Errico et al. 2021). This information is essential if we are
to use DNA extracted from personal ornaments to find out whether individuals of
different sexes perhaps wore different ornaments or perhaps the same but attached or
displayed differently. In addition, when present, such as it is the case for the pendant
described by Essel and colleagues, mineral pigments and residues should be chemically
characterized as this information is crucial to document what raw materials were
involved in the manufacture and use of the object and where they came from (Cristiani
et al. 2014; Rigaud et al. 2014; Martí et al. 2017; Collins et al. 2020; d'Errico et al. 2021;
Velliky et al. 2021). All this documentation is missing in the Essel et al. paper.

But there is more to consider here. Genetic analysis of the tooth identifies it as belonging
to the Asian wapiti (Cervus canadensis). Like the European red deer (Cervus elaphus),
this species has two non-functional canines in its jaw that do not have opposing teeth
and therefore gradually wear away over the course of the individual's life against the
gums covering the mandible in the space between the incisors and the premolars (Greer
& Yeager 1967; d'Errico & Vanhaeren 2002). Archaeologists who have studied the use
of these teeth as personal ornaments have shown that they are characterized by both
strong sexual and inter-individual dimorphism (figure 2) and that it is possible, by relying
on qualitative and metric parameters (figure 3), to identify the sex and age of deer from
their canines, and to know whether two canines originally belonged to the same deer
(Greer & Yeager 1967; Vanhaeren & d'Errico, 2003b). These methods have been
repeatedly successfully applied to address issues such as exchange networks and social
inequality during prehistory (Vanhaeren & d'Errico 2005; Rigaud et al. 2010; Rigaud 2013). An analysis of the available photographs of the perforated deer canines discovered at Denisova (Figure 4) reveals the interest of such an approach. It highlights the use as ornaments of both stag and hind canines of different maturity, including very old hinds and, interestingly, the absence of pairs of canines originating from the same animal. Although this information needs to be confirmed by in-depth analysis, it supports an interpretation of the perforated canines as the result of exchanges.

Combined with results provided by a successful paleogenetic analysis of the ornaments, sex attribution of wapiti canines opens the possibility of looking into whether men exclusively or preferentially wore stags' and women hinds' or vice versa. It may even be possible to compare the genetic ancestry of two humans wearing canines originating from the same deer or establish whether a person was wearing two canines of a single deer. Judging from the photo of the tooth presented in the article, the individual who left her DNA on the pendant wore a hind canine. But how can we move forward in this pursuit if the sex attribution or at least the diagnostic characters and metric data allowing such an attribution are not provided, if nothing else in the accompanying supplementary information of the article?

Among the possible outcomes of such kinds of studies, one might be able, for instance, to show that the canines systematically worn by women were of hinds, thereby opening new horizons for the understanding of the symbolic systems of the populations living in this region of Eurasia during the Upper Paleolithic. Reaching conclusions on this matter requires that archaeologists critically evaluate the genetic results and ask their geneticist colleagues to do the same in light of the archaeological evidence. For instance, Essel et
al. state that the person who produced or wore the ornament they analysed was a woman. However, judging by their text and Extended Data Table 3, that is not exactly what their data seem to be saying: "Among the mtDNA fragments overlapping these positions, 86.6% (95% CI: 82.2-90.5%) match the state of DCP1, suggesting that the mtDNA fragments recovered in this fraction originate predominantly, but not exclusively, from a single ancient human individual, presumably the user or the maker of the pendant". So, since they convincingly show that almost all the DNA recovered is ancient, this means that the human DNA extracted from the pendant could in fact have come from more than one ancient individual, which, if consistent with the archaeological analyses that Essel et al. failed to carry out, might support its interpretation as evidence for long-distance cultural exchange. In short, the tooth's archaeological retrieval and analysis, subsequent to its having been lost or disposed of in the cave, is but the last episode in a long string of processes involving different actors, both anthropogenic and natural. Considering all these methodological aspects will in the future help in adequately address the multiple avenues of paleoanthropological interpretation raised by such aDNA evidence.

## Conclusion

The issues discussed here do not reveal any unbridgeable conflicts between archaeology and paleogenetics, but they do highlight that the integration of rich and reliable archaeological data is an essential prerequisite for the advances in DNA research to be able to make meaningful contributions to our understanding of human history. The potential of these advances is not in question. However, this potential cannot be realized
in the absence of reliable chronological and stratigraphic data and quality archaeological
observations. In a nutshell: geneticists are ill-advised if they believe that their discipline
will one day allow them to reconstruct human history without the support of quality
archaeological data; and archaeologists must likewise avoid the temptation to consider
that their work ends with the excavation of a site and the transference of samples to
their geneticist colleagues.

Acknowledgements

We thank Solange Rigaud for the critical reading of the manuscript. This research was
funded by the following agencies: Initiative d'Excellence IdEx, University of Bordeaux,
Talent program grant # 191022-001 (Fd'E, LD); French government in the framework of
the University of Bordeaux's IdEx "Investments for the Future" program / GPR "Human
Past" (Fd'E, LD); Research Council of Norway, Centres of Excellence (SFF), Centre for
Early Sapiens Behaviour, SapienCE grant # 262618 (Fd'E); European Research Council
(Synergy Grant for the project Evolution of Cognitive Tools for Quantification (QUANTA), No.
951388); French Ministry of Education and Research (JB).

Authors’ contributions

FD wrote the article with inputs from all other authors.

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Figures
<figure><img src="image_2.png" /><figcaption>Figure 1. Pictorial representation of an exchange of ornaments between two women living in Siberia during the Upper Palaeolithic; creation by NightCafe Studio free AI (https://creator.nightcafe.studio) after a text of Fd'D.</figcaption></figure>
<figure><img src="image_3.png" /></figure>

Figure 2. a: perforated and incised red deer canines from the late Upper Palaeolithic site of the Avène des Iboussière, Malaverne, France, one of the collections to which methods to detect sex, age and detect pairs from the same deer were applied; b: modern stag (top) and hind (bottom) canines from the Island of Rum, Scotland, showing from left to right increasing wear of the crown corresponding to aging; c: dramatic individual dimorphism in stag canines from the Ruhm island. Scale = 1 cm. (photos F. d'Errico, b-c modified after d'Errico & Vanhaeren 2003).
<figure><img src="image_4.png" /><figcaption>Figure 3. Metrical and morphological variables used to identify sex and age of red deers from their canines (modified after d'Errico & Vanhaeren 2002).</figcaption></figure>
<figure><img src="image_5.png" /><figcaption>Figure 4. Perforated canines of large cervids found at Denisova cave (modified after (Derevianko & Shunkov 2004; Shunkov et al. 2020; Essel et al. 2023): 4, 8, 9: EUP, Main Chamber; 1-3, 5, 7: IUP, East Chamber, IUP, 6) South Chamber. Canines extracted from male individuals (1-4), female individuals (5-9). From the top to the bottom of the figure, canines are ordered according to the maturity of the individuals, indicated by the degree of use-wear of the occlusal side of the crown (preliminary assessment following the method established by</figcaption></figure>
d'Errico & Vanhaeren, 2002). Younger individuals are at the top of the figure, older individuals
at the bottom.