# An Integrated Evolutionary Scenario for the Culturalization of the Human Body

Francesco D'errico

► To cite this version:

Francesco D'errico. An Integrated Evolutionary Scenario for the Culturalization of the Human Body. Franz Manni; Francesco d'Errico. The Oxford Handbook of the Archaeology and Anthropology of Body Modification, 1, Oxford University Press, 2024, Oxford Handbooks Online, 9780197573433. 10.1093/oxfordhb/9780197572528.013.1. hal-05410933

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# An Integrated Evolutionary Scenario for the Culturalization of the Human Body

Francesco d'Errico

## Introduction: Permanent Body Modification, and Beyond: An Integrated Approach

The culturalization of the human body refers to the processes by which individuals and societies imbue physical bodies with cultural meanings, values, and symbols (Lock 1993; Joyce 2005, 2007; Borić and Robb 2014; Nowell and Cooke 2024; Collins and Nowell 2024). These practices include both nonpermanent modifications like clothing, body painting, personal ornaments, hairstyles, and embroidery, and permanent alterations such as tattooing, scarification, body piercing, implants, genital and dental modifications, and bone shaping, as well as amputation (DeMello 2007).

In every society, the human body carries intrinsic meanings and symbolism long before any permanent or temporary modification takes place (Waskul and Vannini 2016; Joyce 2005; Allolio-Näcke 2019; Goodson 1991; Mascia-Lees 2011). These meanings generally reflect cultural beliefs, social roles, and individual identities, and they shape perceptions and behaviors within communities worldwide. Before any type of body modification occurs, living bodies are already imbued with societal expectations and norms associated with their gender identity, age group, and social affiliation (Frederick and Reynolds 2021). These frameworks influence perceptions of masculinity, femininity, youth, maturity, and expected cognitive and physical performances and shape how individuals navigate their roles, relationships, and interactions within and beyond their communities.

The simplest “treatment of the body” encompasses practices of not only deliberate exposure to natural elements like sunlight, water, and wind, but also of cleansing, anointing with oils, and applications of substances such as clay, charcoal, perfumes, makeup products, and others (Havlíček and Roberts 2012; Ward 2019; McCabe, Malefyt, and Fabri 2020). These practices not only maintain physical hygiene but can also carry symbolic significance, reflecting spiritual purification, healing, and connection to nature.

In all known human societies, however, the “naked body” is also a canvas on which technologically complex, time-consuming, and, in some cases, permanent and potentially harmful practices take place.

These practices are important for individuals and societies for different reasons. Some, such as clothing, fulfill practical functions of protection, insulation, hygiene, and modesty (Gilligan 2010; Gilligan et al., 2024; d'Errico et al. 2018). The culturalization of the human body is also a form of self-expression, allowing individuals to construct and assert identity within a cultural context (Tiggemann and Hopkins 2011). In ethnographically documented societies, practices of body culturalization are often deeply rooted in cultural and religious traditions (Rush 2005; Sanders and Vail 2008; Hambly 2009). They can serve as visible markers of cultural heritage and spiritual beliefs, helping to strengthen communal bonds and shared cultural narratives. They may also have therapeutic functions (Alter-Muri 2020; Lynn et al., this volume; Renaut, this volume) or, as in the case of talismans, amulets, and charms, possess protective or magical properties designed to bring good luck, ward off evil, or provide other specific benefits to the wearer (Varner 2008; Pennick 2021).

Temporary body modification offers greater flexibility by allowing experimentation with different identities or self-expressions (Hester and Hehman 2023), particularly in societies that allow social mobility, when social roles and status change. In contrast, permanent body modifications involve an irreversible change in the appearance of the body (DeMello 2007). Techniques for removing tattoos and other invasive body modifications exist in contemporary Western societies, but such practices were not available or practiced in most ancient societies. Permanent modifications generally reflect significant and durable cultural and symbolic meanings, serving as enduring symbols of social and ethnic identity.

Wherever they occur in historically attested or contemporary societies, permanent modifications certainly tend to and have a profound impact on how individuals are perceived within and outside their societies, within and outside the private sphere, and can evoke strong reactions (Wohlrab et al. 2009).
While in most ethnographically and historically documented societies body modifications were not a free
individual choice (but see LaTosky and Robert, this volume), in many contemporary societies rules have
often become more elusive, leaving room for practices designed to break those same rules in order to
signal affiliation to subcultures (King and Manni, this volume). This is also why body modifications
practices are now seen as a form of artistic expression in their own right (Lodder 2022, 2023).

From an evolutionary perspective, the emergence of both temporary and permanent body modifications
presents fascinating paradoxes. Hominins would have gradually lost their fur approximately two million
years ago and developed a sweaty, naked skin as a genetic adaptation to hot savanna environments to
increase hunting mobility (Jablonski 2006). According to others, the loss of human body hair would have
occurred more recently, 1.2 million years ago (Rogers, Iltis, and Wooding 2004). At the same time,
hominins living close to the equator would also develop a darker skin to be protected from the high
ultraviolet irradiation of these latitudes (Jablonski and Chaplin 2010). When, why, and how, after
reaching such adaptations, some members of our genus started to alter their appearance on purpose, that is
culturally, remains a largely open question. Similar open questions concern the evolutionary paths of the
culturalization of the human body and the reasons for this behavior that has reached a remarkable
complexity since several thousands of years as this volume shows (Gilligan 2010; Vanhaeren and
d'Errico 2011; Dubin 2009; DeMello 2007; Allolio-Näcke 2019).

It has been suggested that early humans might have used clothing in ways similar to how animals use mimicry or camouflage. For example, they might have used it to sneak up on prey or hide from predators (Buckner 2021). However, if this idea is true, it would only explain why humans started wearing clothes, not their subsequent evolution.

The emergence of practices devoted to permanently modifying the appearance of the human body
represents, in evolutionary terms, a specific corundum. Although permanent modifications may reduce
fitness because of the risks of severe infections, their large diffusion in all continents suggests they
offered significant advantages at the societal and individual levels (Valentova, Mafra, and Varella 2022).
One possible benefit of permanent modification is in signaling group membership in a durable and reliable
way (the flag function). By manifestly displaying the permanent adherence to cultural norms and values,
permanent body modification may have facilitated cooperation, strengthening group cohesion and
ultimately promoting the survival and success of the community as a whole. Today, contemporary body
modification practices everywhere show that permanently changing the appearance of the body can buster
self-expression and contribute to psychological well-being, social integration, and mating success, factors
that are crucial for individual and collective fitness.

More generally, the culturalization of the human body may have also facilitated the emergence and
reinforcement of cumulative cultures—that is, cultures in which knowledge, skills, and practices are built
up over time through social learning and passed down from generation to generation within a group
(Dean et al. 2014). By promoting transmission by imitation rather than emulation (Whiten et al. 2009),
culturalization may have contributed to the perpetuation and evolution of complex cultural traditions,
including those not related to permanent or temporary body modification.

It is clear that by understanding the evolutionary implications of these practices, we can better apprehend
who we are and what makes human societies different in many ways from those of our close relatives.
Current research on the evolution of our lineage reveals, on the one hand, a large and ever-increasing
number of fossil species and, on the other hand, the observation, confirmed in the last decade by
paleogenetic data, that several of them, such as Neanderthals and Denisovans, appear more and more to
be the phenotypic expressions of populations that, over long periods, constantly exchanged their genes
with modern populations, which were themselves the result of a long evolutionary process within the
African continent. How do we integrate the culturalization of the body into this changing evolutionary
scenario, which supports both ancient and complex origin of our species and its genetic and cultural
proximity to other fossil populations? Did the culturalization of the human body experience recognizable
tipping points in the way humans culturally transformed their bodies, possibly corresponding to
speciation events, or are we facing a gradual process with different past populations described in the past as different species sharing comparable practices?

These are challenging questions considering that the vast majority of these practices leave no trace or only ambiguous or circumstantial traces of themselves in the archaeological record before recent periods, making it difficult to determine when they developed, how they changed over time, and how they eventually coevolved.

A telling example of what may be outside the grasp of archaeological inquiry is seen in the grass-in-the-
ear behavior documented in the Chimfunshi Wildlife Orphanage in Zambia (Van Leeuwen, Cronin, and
Haun 2014). After a chimpanzee named Julie inserted a grass stem into her ear, the practice was adopted
by 8 out of 12 group members and continued to spread even after Julie died, indicating that practices to
alter the individual appearance can be transmitted through social learning in animals even in situations in
which the behavior appears to lack adaptive significance. Although the new behavior appeared in an
orphanage, and not in a chimpanzee population living in the wild, Julie does not seem to have learned it
from humans. This example indicates that practices leading to the culturalization of the body may have
very ancient roots, possibly going back to our last common ancestors with chimpanzees, and that the
available archaeological evidence probably illustrates situations in which these practices were already
consolidated. Their first adoption may have long preceded the first robustly attested cases.

In addition, each type of body culturalization practice has a different likelihood of leaving direct or
circumstantial traces in the fossil and archaeological record. Because they are applied to skin, and only
preserved in exceptional cases, permanent modifications such as tattooing, scarification, and genital
modifications are much less likely to leave direct archaeological traces than, for example, amputations or
bone and tooth modifications. It is therefore not surprising to find (Webb 1995; Willman and Lacy 2020;
Kaifu et al. 2011; Ni et al. 2020) that bone modifications such as teeth ablation, attested at Mungo,
Australia circa 45 kilo annum (ka); at Ohalo II, Israel circa 22.5 ka; and at Minatogawa, Japan circa 12-
11 ka, and cranial modifications described at Songhuajiang, China and dated to circa 11 ka, feature
among the earliest known permanent modifications. On the contrary, the oldest attested tattoos (Spindler
2013; Deter-Wolf et al. 2024), scarifications (Roehrig 2015), and genital modifications (Cox and Morris
2012) date back no further than 5 to 4 ka, and this thanks to the exceptional discovery of the tattooed
body of Ötzi, the so-called Iceman, found in 1991 in the Italian Alps.

Because they are often applied to the skin (which is only preserved by mummification), the oldest attested
tattoos (Spindler 2013; Deter-Wolf et al. 2024), scarifications (Roehrig 2015), and genital modifications
(Cox and Morris 2012) date back no further than 5 to 4 ka). Differently, because they leave traces on
osseous tissues, teeth ablations are attested as early as circa 45 ka at Mungo, Australia; circa 22.5 ka at
Ohalo II, Israel; and circa 12-11 ka at Minatogawa, Japan: they are among the earliest known permanent
body modifications (Webb 1995; Willman and Lacy 2020; Kaifu et al. 2011; Ni et al. 2020). Cranial
modification described at Songhuajiang, China, dates to circa 11 ka and is among the oldest examples.

Circumstantial evidence for the origins of tattooing and scarification remains exceedingly equivocal. Body piercing, because specific nonperishable ornaments, being made of shell, bone, or stone, often resist the deteriorations of time, probably dates from as early as 40 ka (Langley, O'Connor, and Aplin 2016). Numerous engraved, painted, or carved human representations dated to the Upper Paleolithic, starting from the 40 ka Aurignacian carvings found at sites of the Swabian Jura, are available (Bourrillon 2009; Dutkiewicz 2021; see also Sernicola and Manzo, this volume). It is difficult to establish whether such designs represent scarifications, tattooing, body painting and clothing, or other symbolic markings. The same ambiguity applies to archaeological stone drills, bone awls, pins, and needles that may have been used for tattooing and body piercing, and to the sharp blades that may have been used for scarification or tattooing, because these tools were also used in a variety of other activities.

The differential persistence of permanent and temporary body modifications in the archaeological
record—that is, their different taphonomy—underscores the need for an integrated approach to identify
long-term trends and their wider implications. In this chapter, I summarize what we know about the origin
and evolution of three key technologies for modifying the appearance of the human body: clothing, mineral pigments for body modification, and personal ornaments. The analysis of these three cultural proxies can shed light on the emergence and evolution of other fundamental practices of the culturalization of the human body. To argue that temporary and permanent body modifications are "communication technologies," facilitating the process of niche expansion undertaken by our genus at least 500 ka, I will build on syntheses (Gilligan 2010, 2023; d'Errico et al. 2018; Doyon et al. 2023; Gilligan et al., 2024; Dapschauskas et al. 2022; Rosso 2022; d'Errico et al. 2023) and a recently published experimental work (Salagnon et al. 2024). The available evidence supports the view that the gradual complexification of these practices reflects cultural exaptations—that is, the repurposing of existing cultural features for new functions due to changing circumstances or social dynamics (Colagé and d'Errico 2020; d'Errico and Colagè 2018). The timing and pace of the emergence of more complex practices suggest that the exaptations marking new steps in the culturalization of the human body were primarily cultural in origin, and largely independent from the biological changes associated with the emergence of new species.

## Clothing and Needles

Clothing has been an essential support onto which other practices devoted to the culturalization of the human body were grafted (Gilligan 2010; Gilligan et al., 2024; Abadia and Nowell 2015; d'Errico et al. 2018). Bone-eyed needles have traditionally been considered the most reliable archaeological artifact for inferring the manufacture of clothing, but this assumption is now being challenged on a number of grounds. The earliest known eyed needles appeared around 40,000 years ago in Siberia and around 30,000 years ago in East Asia (Figure 1). They did not appear in Europe until about 26,000 years ago (d'Errico et al. 2018). Though small in size, eyed needles have been consistently found in recent and older excavations in both East Asia and Europe after those dates, indicating that when the needles become an integral aspect of the technical system of a society, they are found in significant numbers. Their delayed appearance in Europe, and their absence from older sites, is therefore unlikely to be due to the loss of earlier specimens. To summarize, the absence of needles at Middle Paleolithic and initial European Upper Paleolithic sites in Europe suggests that needles represent a cultural innovation appearing around 40,000 years ago in eastern Eurasia, and somewhat later in Europe, and that eyed needles had not been previously used in Eurasia or Africa. This observation prompts questions about how the so-called archaic hominins who lived in Africa and Eurasia between two million years ago (Ma) and circa 50 ka survived in cold environments without specialized sewing technologies. The same question applies to early Homo sapiens living in Africa after 300 ka BP and arriving in Eurasia after 60 ka BP. It is hard to imagine that no clothing existed in those cold environments; this is why the reliability of eyed needles as a proxy for the emergence of tailored clothing has been questioned. Although physiological and cultural adaptations, along with a high caloric intake, may have contributed to cold-weather survival, the absence of eyed needles suggests that early human populations may have favored technological strategies for alternatives to clothing to cope with low temperatures and wind chill.
<figure><img src="image_2.png" /><figcaption>**Figure 1.** Bone eyed needles found at Xiaogushan (a) and Zhoukoudian Upper Cave (b), China, compared with Magdalenian needles from France (c) and Moravia (d). Scale = 1 cm. Source: Adapted from d'Errico et al. (2018).</figcaption></figure>

An indirect way to date the appearance of clothing has been attempted by comparing the genetic diversity of body lice and hair lice. Body lice (Pediculus humanus humanus), which are well-adapted to clothing, had diverged from head lice (Pediculus humanus capitis) by about 80 ka, possibly even earlier (Kittler, Kayser, and Stoneking 2003; Toups et al. 2011). The most likely estimate for the time when head lice and clothing lice diverged is about 83 ka, as shown by the mode of the posterior probability distribution. The median value of this estimate is 170 ka, with a 95% highest probability density range from 29 ka to 691 ka.

The enduring relationship between body lice and clothing suggests that clothing has been an integral part of human culture for millennia, well before the advent of eyed bone needles. Another reason to question eyed needles as “the specific marker” providing evidence for tailoring skills is their relative fragility. Eyed needles are particularly suited to sewing soft leather skins or embroidering ornaments, but when it comes to sewing thick skins, then the perforations to be connected through sewing must have been practiced beforehand with other types of tools.

Although archaeological evidence for early clothing made before the invention of eyed bone needles remains eminently circumstantial, data on human physiology, ethnography, and findings dating back to the Early and Middle Pleistocene indicate that tools for hide preparation and thermal insulation must have been already in use in those times (Gilligan 2010, 2023; Collard et al. 2016; Gilligan et al., 2024). Evidence of animal skinning and hide scraping has been uncovered at various sites in Eurasia since 780 ka (Carbonell et al. 1999) suggesting early clothing technologies (Gilligan 2010; d'Errico et al. 2018; Gilligan et al., 2024). Bone smoothers, presumably used for working skins, have been discovered at several sites, including Schöningen (Germany), Grotte des Contrebandiers (Morocco), Pech-de-l'Azé I, Combe-Grenal, and Abri Peyrony (France). Delicate bone awls suitable for piercing hides have been found at Blombos Cave and Sibudu in Southern Africa (Henshilwood et al. 2001; d'Errico and Henshilwood 2007) dating to approximately 80 ka to 61 ka, as well as Carpenter's Gap 1 in Australia and Grotte du Renne in France, dating to 47 ka and 45-42 ka, respectively (Figures 2 and 3). That skin, or possibly some other material, was probably used to cover the body as early as the Middle Paleolithic is also suggested by the discovery of human footprints indicating the use of shoes. The earliest evidence of the use of footwear comes from the Theopatra cave in Greece, where footprints of children wearing shoes, probably Neanderthals, have been dated to 110-135 ka (Kyparissi-Apostolika and Manolis 2021). More recent are the footprints discovered in the Cussac cave, Dordogne, France, dated to 28-31 ka (Ledoux et al. 2021). Circumstantial evidence for the use of footwear comes from the marked reduction, in the same period, in the robusticity of the lesser toes (Trinkaus 2005).
<figure><img src="image_3.png" /><figcaption>**Figure 2.** Bone awls from Middle Stone Age layers of the Sibudu rock shelter (a) and Blombos Cave (b), South Africa, dated to between 80 ka and 60 ka. Source: Adapted from d'Errico, Backwell, and Wadley (2012) and Henshilwood et al. (2001).</figcaption></figure>

<figure><img src="image_4.png" /><figcaption>**Figure 3.** Bone awls produced by late Neanderthals found in the Châtelperronian layers of the Grotte du Renne, France, dated to c. 45–42 ka. Source: Adapted from Julien, Vanhaeren, and d'Errico (2019).</figcaption></figure>

Based on thermal insulation properties, clothing can be categorized into two forms: simple and complex (Gilligan 2010). Simple clothing is loose in structure, while complex clothing is fitted and may consist of multiple layers. Neanderthals may have opted for the former, possibly combining their clothing with the application of fat on the skin for insulation purposes. We have, however, direct evidence that Neanderthals produced complex, twisted cordage (Hardy et al. 2020), supporting the hypothesis that they could make fitted clothing. Thus, the absence of specialized bone tools cannot be used as evidence for the absence of clothing during the European Middle Paleolithic, and available paleoenvironmental reconstructions indicate that Neanderthals must have worn clothing to cope with the climatic conditions of the late Middle and Upper Pleistocene (Gilligan 2010; Collard et al. 2016).
Further, a recent discovery from Canyars, a site in Catalonia, Spain, reveals the use of prehistoric sewing techniques before the advent of eyed needles in Europe (Doyon et al. 2023). Microscopic analysis of the punctures on a mammal bone fragment dated to 39,600 cal BP and discovered in an Aurignacian layer suggests they were made with robust flint burins, probably to perforate leather pieces for clothing, indicating early tailoring skills. This finding suggests that initial Upper Paleolithic hunter-gatherers could craft tailored leather clothing for enduring harsh climates 14,000 years before eyed needles were introduced to Europe. Experimental reproduction of use wear on Noailles burins from the Gravettian culture (33–24 ka) indicates they may have been used to pierce skins (Calvo et al. 2019), further challenging the assumed link between eyed needles and tailoring.

A comprehensive examination of earliest bone needles from Eurasia and North America indicates regional variability in dimensions, shapes, and potential evolutionary patterns in morphology over time (d'Errico et al. 2018). The earliest examples of eyed needles unearthed in China are notably large and robust, implying their utilization in crafting thick, custom-made attire and durable leather items. The noticeable disparity in size between early needle specimens from Siberia and those from China hints at the likelihood of independent innovations in these regions, which correspond with distinct lithic technologies associated with the earliest needles. In Siberia, there is minimal fluctuation in needle size throughout the Upper Paleolithic period; whereas in China, smaller, flatter needles began to emerge between 26 ky and 23 ky. Further diversification occurred in China after 23 ka, resulting in the noticeable variations in size observed at the site of SDG12, dating back 11,200 years. This shift likely correlates with the diversification of tasks necessitating needle use, with smaller subcircular specimens possibly intended for embroidery, appliqué, and sewing delicate furs for undergarments. Flat, more robust needles might have been utilized for assembling winter clothing. The emergence of bone needles at the Gravettian and Gorotsovian sites on the East European Plain around 26 ka may signify either an independent innovation or a geographical extension of early Siberian sewing practices. Interestingly, despite ample evidence of clothing and garment use during this period, no eyed needles have been discovered at numerous Gravettian sites in Central and Western Europe (ca. 33 ka–25 ka) or at primary burial sites, such as Sungir in Russia, where the use of elaborated, possibly multilayered clothes is inferred from the spatial distribution of the ornaments on the skeletal remains (Bader and Lavrushin 1998; Trinkaus and Buzhilova 2018). Eyed needles made their initial appearance in Europe during the Solutrean culture, approximately 26 ka, coinciding with the lower winter minimum temperatures characteristic of the Last Glacial Maximum. The narrow size range of needles utilized by Solutrean hunter-gatherers likely suggests a limited degree of specialization, with needles of similar dimensions serving various purposes. However, in the subsequent Magdalenian period, a distinct size differentiation emerges, with small, delicate needles on one end of the spectrum and large, sturdy ones on the other. This suggests that the sewing toolkit of these Late Glacial hunter-gatherers contained needles of varying sizes that were specifically crafted for different tasks. Differences in morphometric variation among needles discovered in Western Europe and in Central Europe indicate a significant regionalization in manufacturing traditions during the subsequent Magdalenian culture.

Paleoindian eye needles from North America, the earliest of which have been dated to 14 ka, are characterized by a narrow size range that reflects an unprecedented level of mastery in the production of this bone tool. It raises the question of whether this specialized expertise was inherent in the cultural practices of the human groups that first colonized North America, or whether it was developed after their arrival on the continent.

Whatever the case, it is clear that the production of these thin needles played a crucial role in humans' adapting to the environmental challenges of the mid- to high latitudes of North America. Bone eyed needles are only attested at recent prehistoric sites in Africa and are not recorded in Australia and Tasmania during the last glacial maximum.
Building on the foregoing evidence, Gilligan and colleagues (2024) propose a long-term evolutionary scenario for the emergence of clothing and dress (Figure 4). They posit that following the initial migration of early humans out of Africa, possibly two million years ago, the primary use of animal skins was to protect against the elements. These skins were likely draped over the body in a basic manner, offering rudimentary insulation and shielding. The emergence of notched stone tools and borers suggests that humans residing in the middle latitudes might have started to develop basic sewing techniques and methods for wrapping animal skins around the body between 500 ka and 300 ka years ago, if not earlier. The invention of bone awls around 80 ka in Southern Africa enabled humans to create tailored garments by perforating skins with greater precision.

This technological leap led to clothing that was better fitted to the contours of the body, offering enhanced protection and comfort. Bone awls, however, are not recorded at most African and Eurasian sites before 45 ka or later. It is very possible that bone awls remained an innovation that for a long time was restricted to certain regions or that their function was fulfilled by stone borers or awls made of perishable material.
<figure><img src="image_5.png" /></figure>

**Figure 4.** Tentative evolutionary scenario of the emergence and diversification of clothing in different regions and on a global scale, contrasted with the evolution of the climate for the las 5 Ma from variation in the δ¹⁸O record (climatic data from Lisieck and Raymo 2005).

Around 40 ky BP in both eastern Eurasia and Europe, the development of eyed needles and adapted punching techniques, respectively, allowed for a more efficient and durable sewing of clothing. This innovation paved the way for the creation of intricate stitched garments with improved functionality and aesthetic appeal. The use of differently sized bone eyed needles, documented after 30 ky, facilitated the crafting of even more elaborate tailored clothes and undergarments. As humans mastered clothing construction techniques, they began to embellish their garments with ornaments and decorations. This may have happened as early as 70 ka when we see, at sites such as Blombos and Sibudu, the association of both thin bone awls and shell beads, in particular, after 50 ka, in the attaching of ostrich egg shell beads on clothes, and during the Upper Paleolithic of Eurasia, when bone needles were used not only for practical sewing purposes but also for attaching decorative elements like beads and shells, adding cultural and aesthetic significance to clothing.

When this evolutionary scenario is contrasted with the climatic evolution of the planet over the last five
million years (Figure 4), it becomes clear that explaining the emergence of clothing solely as an
adaptation to the gradual climatic deterioration observed over this long period is challenging. Climate degradation was already well established before 500 ka, the time at which early clothing technologies are thought to have emerged. An interpretation more consistent with the climatic data suggests that this innovation and subsequent exaptations arose in response to shifts in the frequency and intensity of glacial-interglacial cycles. These shifts led to more pronounced glaciations toward the end of the Middle-Upper Pleistocene transition. Viewing the first items of clothing and their subsequent refinements in this context suggests that they were a symptom of the development of cumulative cultures, allowing cultural adaptations to successfully address climatic changes that human populations could not cope with through natural selection or epigenetics alone (Boyd and Richerson 2005).

This evolutionary narrative underscores the multifaceted role clothing has played in the gradual process of niche construction undergone by members of our genus. Clothing allowed our ancestors to inhabit a wider range of environments, granting them access to various resources and ecological niches (Gilligan 2010). By increasing individuals' chances of survival, longevity, and overall health, clothing facilitated the accumulation of cultural knowledge, leading to heightened cultural complexity and social identities. The technologies that enabled prehistoric populations to manufacture clothing may have had an influence on or co-evolved with technologies for permanent body modifications. Bone awls, the earliest of which are 80 ky old; bone needles; and sharp lithic tools could have been adapted for use in body modifications, including permanent modification practices. Needles may have been used for tattooing. The pigments used to dye clothing might also have been used for tattooing, and sharp objects used for cutting and sewing clothing could have been used for scarification.

## Ochre

Red ochre holds profound significance for many human cultures, serving as a means of symbolic communication to express individual, social, collective, and religious identities (Rosso 2022).

Archaeological evidence suggests that the use of pigments did not suddenly emerge 40 ka, as was thought a quarter of century ago, but evolved gradually, from ancient origins spanning different fossil human species. The use of a variety of iron-oxide-rich rocks, known as ochre, have been traced back at least 400,000 years in Africa (Rosso 2022 Dapschauskas et al. 2022). Outside Africa, the use of ochre is attested in Europe and the Near East starting 380 ka, for example, at the site of Terra Amata in France and Maastricht Belvedere in the Netherland. Evidence from more recent Middle Paleolithic sites has shown that European Neanderthals used red pigments but also black ones, mostly manganese oxides, and stained with red and yellow ochre shells, fossils and large stalagmites located in deep caves. In Asia, apart from microscopic red residues found on an engraved bone at the Chinese site of Lingjing, dated to circa 105-125 ka; a fragment of ochre found at a site in India possibly dated to more than 70 ka; and ochre pieces found at another site in Australia, all other instances of pigment use are more recent than 55 ka and compatible with the hypothesis that a systematic use of ochre was introduced in this vast region by expanding modern human populations (d'Errico 2021).

The concomitant use of ochre by traditional societies for utilitarian purposes (protection from the sun or insects, in the tanning of hides etc.) has led scholars to regard mineral pigments as an ambiguous element of the archaeological record. Their presence alone, without further compelling evidence, is considered by many to be insufficient to conclusively prove symbolic use, let alone symbolic use for the culturalization of the body.

Against this view, it has been argued that the symbolic and utilitarian functions of ochre are inherently intertwined in traditional cultures, and that, as a consequence, it is unlikely that the systematic use of pigments would have persisted over time without quickly acquiring a symbolic dimension. Evidence for the early involvement of colored powders in symbolic practices includes the gradual expansion of their use since 500 ka.
A recent synthesis by Dapschauskas et al. (2022) supports this view. These scholars delineate three phases of ochre utilization during the African Middle Stone Age: an initial period, from 500 ka to 330 ka; an “emergent” phase spanning 330 ka to 160 ka; and a “habitual” phase, from 160 ka to 40 ka. They interpret the latter phase, characterized by the presence of ochre at a third of African archaeological sites, as indicative of intensified ritual practices among early Homo sapiens populations. Other evidence supporting a symbolic use of mineral pigments include the preference for specific colors, the procurement of specific high-quality mineral pigments from distant sources, the deliberate heating of iron-rich rocks to change their color, the application of ochre to modify the color of personal ornaments, the creation of abstract engravings on ochre pieces or drawings with ochre pencils, and the demonstrated production, at some sites, of small quantities of powder that would have been more suited to symbolic activities than utilitarian ones.

Although not everyone subscribes to these viewpoints, several discoveries demonstrate that ochre was used in increasingly complex practices between 400 ka and 40 ka, and that some of these practices were symbolic in nature. Geochemical analyses show, for example, that Middle Stone Age populations from Southern Africa preferred to travel long distances to collect ochre of a particular color and purity. It has also been demonstrated that since at least 110 ka, yellow ochre has been heated to produce red ochre, as seen at the Skhul site in the Near East. One hundred ka ago, at Blombos cave in Southern Africa, two types of ochre were knapped, pounded, ground, and probably mixed with animal fat to produce a semiliquid compound that was stored in containers made of large marine shells. At the same site, abstract designs are engraved on ochre pieces between 100 ka and 70 ka, and ochre pencils were used to draw abstract motifs 70 ka. From 140 ka onward, ochre was deliberately used to color personal ornaments in North Africa and Southern Africa, a behavior with no apparent functional purpose. This suggests that, by this time at least, the use of pigments had become an integral part of the culturalization of the human body.

Ochre is ubiquitous in early Upper Paleolithic sites across Europe and Eurasia, including those attributed to Neanderthals, such as the Grotte du Renne in northern France. At this site, more than 18 kilograms of red and yellow ochre, as well as black pigments consisting of manganese oxides, were found in Châtelperronian layers in association with numerous Neanderthal remains (Salomon 2009). Residues of ochre are also abundant on bone tools used for working and piercing skins, suggesting that the clothing produced was also dyed red (Julien, Vanhaeren, and d'Errico 2019). The use of ochre in connection with the body became even more prevalent from the Gravettian period (33–24 ka) onward. The many buried human skeletons of this period and their associated ornaments are systematically ochre-stained, indicating that individuals were extensively covered in ochre during their lives and that their bodies may have been wrapped in ochre-stained skin shrouds (Pettitt 2010).

The synthesis just outlined shows that the use of ochre by human populations has a long history beginning at about the same time as the possible invention of the first rudimentary clothing. The use of ochre became more complex in some regions over time, but remained absent or latent in others until its use became systematic from about 160 ka, first in Africa and then in Europe. The use of ochre continued to increase throughout the Upper Paleolithic in Eurasia, covering virtually all aspects of the individual, social, and spiritual lives of these populations. It is likely that in the early stages of this long process ochre was used primarily for utilitarian purposes—for example, to counteract the negative effects of ultraviolet radiation—and that later these functions were exapted and the use of ochre was incorporated into a growing body of symbolic practices. Some of these exaptations may have involved the use of pigments in the production of permanent body modifications. Without providing proof, the search for ne ochre with high tinting power and the production of ne bone awls indicate that, technically, permanent and temporary modifications involving pigments, such as ochre and charcoal in tattooing, body painting, and the coloring of clothing and personal ornaments, may have been practiced from 80 ka ago and possibly earlier, and leave no direct trace in the archeological record.
Personal Ornaments

Personal ornaments have played a central role in the evolutionary history of human societies, serving as symbols of social cohesion, means of mate selection, and markers of social identity and strati cation, thereby influencing patterns of human behavior and social organization through the ages (Kuhn and Stiner 2007; Vanhaeren and d'Errico 2011; Kuhn 2014; Abadia and Nowell 2015; Mattson 2021). For a number of good reasons, personal ornaments are often regarded as an important indicator of the development of a culturally modified body. Unlike mineral pigments and needles, which serve as indirect evidence for body painting, tattooing, and clothing, ornaments offer direct testimony to the process of body culturalization. Their production and use involve the selection of specific materials, development of technologies for their manufacture and display on the body, and the leaving behind of discernible wear patterns that attest to their decorative use. In periods in which they take the form of many different types (marine and freshwater shells, animal teeth, shaped beads and pendants made of stone, ivory, antler, amber etc.), the study of the geographical distribution of these types makes it possible to propose hypotheses about the cultural geography of the populations involved and its evolution.

Nearly two decades ago, a landmark discovery was made at the Southern African Middle Stone Age site of Blombos Cave, where marine shells used as beads were found in layers dating approximately to 73 ka (Henshilwood et al. 2004; d'Errico et al. 2005; Vanhaeren et al. 2013). This finding challenged the prevailing notion that ornamental practices began with human populations in Europe and appeared no earlier than 42 ka. In the following years, additional marine shell beads were identified at various sites across North Africa, South Africa, East Africa, and the Mediterranean Levant dating back to approximately 142 ka to 40 ka (see Steele, Álvarez Fernández, and Hallett 2019 and d'Errico et al. 2023 for a synthesis). These discoveries clearly indicated that, contrary to previous beliefs, personal ornaments did not suddenly emerge with the level of complexity observed in numerous human cultures today. Rather, they appear to have evolved gradually, progressing through stages whose significance and implications we are only now beginning to understand.

It has been recently proposed that we can currently identify at least ten developmental stages (Figures 5 and 6) in the progression of these practices (d'Errico et al. 2023). The sketched evolutionary scenario is not seen as a strict one-way path. Independent invention and cultural transfer may have bypassed, or even reversed, some of these stages in some regions and historical contexts.

<figure><img src="image_6.png" /></figure>

**Figure 5.** A tentative multistep evolutionary scenario for the culturalization of the human body based on archaeological evidence of pigment and ornaments. The scenario includes evidence from both Africa and Eurasia. Thicker lines indicate continuous and widespread use of the relevant body culturalization technology in large regions. The tapering from thick to thin lines indicates a gradual reduction in the practice. Thinner lines indicate continuous, less intensive use. Dashed lines indicate periods during which we may reasonably assume that the practice was implemented, although the record has not been investigated in detail (after d'Errico et al. 2023).
<figure><img src="image_7.png" /><figcaption>**Figure 6.** Early personal ornaments: (1) *Glycimeris* shell with traces of red pigment and a natural perforation, Qafezh, Israël, c. 95 ka; (2) *Glycimeris insubrica* shell with a natural perforation, Cueva de los Aviones, Spain, c. 115 ka; (3) *Tritia gibbosula* shell with a natural perforation, Skhul, Israël, c. 115 ka; (4-5) Grotte des Pigeons, Taforalt, Morocco, c. 80 ka; (6) *Tritia gibbosula* shell with a natural perforation, Rhafas, Morocco, c. 80 ka; (7) *Tritia gibbosula* shell with a natural perforation, Ifri N’Ammar, Marocco, c. 120 ka. Note that the dark coloring of the shell most likely due to heating; (8) *Tritia gibbosula* shell with a natural perforation, Oued Djebbana, Algeria; (9-12) *Nassarius kraussianus* shells, Blombos Cave, South Africa, c. 73 ka; Note the difference in perforation size and type between shell beads 9-10 and 11-12. The dark coloring of the latter is due to heating; (13) naturally perforated *Conus apexes* used as beads, Panga Ya Saidi, Kenya, c. 60-40 ka; (14) *Conus ebraeus* shell beads associated with the burial of an infant, Border Cave, KwaZulu-Natal, South Africa, c. 74 ka; (15) *Conus ebraeus* shell bead, Border Cave, KwaZulu-Natal, South Africa, c. 74 ka; (16) ostrich egg shell beads, Panga Ya Saidi, Kenya, c. 33 ka; (17-23) personal ornaments made by Châtelperronian Neanderthals at the Grotte du Renne, Arcy-sur-Cure, Yonne, France, c. 45-42 ka.</figcaption></figure>
The first stage would have involved the use of mineral pigments for body decoration. The remarkable increase in the use of ochre during the Middle Stone Age in Southern Africa (see section "Ochre" above) suggests that, though ochre may have served practical purposes at the onset of the Middle Stone Age, around 500-300 ka, its symbolic functions, including its use in body decoration, likely became prominent after 160 ka. A subsequent stage, observed so far in Southern Africa at Blombos and Pinnacle Point (Jerardino and Marean 2010) and at the Middle Eastern site of Misliya Cave (Bar-Yosef Mayer et al. 2020), involved the collection and curation of marine shells, and possibly other natural objects, without modifications. The uniqueness of these specimens, coupled with the absence in some of natural or anthropogenic perforations, suggests that they may have initially served as amulets or similar items. This phase may have coincided with, or been followed by, the use of shells or other objects with natural perforations or features to facilitate attachment. This practice is evident at such sites as Blombos Cave and Pinnacle Point in Southern Africa, Qafzeh in western Asia, and Cueva de los Aviones in Europe between 150 ka and 80 ka. Already at this stage, pendants were coated with ochre, either deliberately or accidentally through contact with ochre-stained skin.

The fourth stage involved the assembly of several homologous objects with natural or minimally modified perforations, such as shells of the same or morphologically similar species, into ornamental compositions. This practice appears in northwest Africa from about 140 ka onward, and it continued in this region until 80 ka.

These beads are associated with a more systematic use of ochre and heating techniques to alter shell colors. The predominance of natural perforations limited the variability of shell arrangements.

The next stage, attested at the Sibudu rock shelter from 80 ka and at Blombos Cave between 73 ka and 70 ka, involved the development of perforation techniques that allowed greater variability in ornamental arrangement and easier production of different ornaments from the same shells. The fifth step involved the incorporation of larger shells of different species into ornaments composed predominantly of smaller shells, or the presentation of the two types of objects separately. This practice is evident at Blombos Cave around 72 ka and at Sibudu around the same time.

A subsequent stage saw the use of subcircular, naturally perforated shell fragments as ornaments. This stage is currently known only from Panga Ya Saidi, Kenya, where naturally perforated Conus apexes were used as beads, possibly between 60 ka and 40 ka (d'Errico et al. 2020). This practice paved the way for the seventh evolutionary stage: the production of fully worked circular ostrich eggshell beads from about 50 ka, probably initially in East Africa. Recent research on these objects shows regional variation, reflected in significant differences in bead size between East Africa and South Africa from 33 ka (Miller and Wang 2022).

The subsequent eighth evolutionary stage, around 45 ka, witnessed a significant diversification, particularly in large regions of Eurasia, characterized by an explosion of ornament types encompassing different raw materials, shapes, production methods, and undoubtedly different modes of attachment to the body. The geographical distribution of these highly diversified Paleolithic ornaments suggests their use as ethnic markers (Vanhaeren and d'Errico 2006; d'Errico and Vanhaeren 2015; Baker et al. 2024). The inclusion of exotic objects obtained via long-distance trade and their variable presence in Paleolithic burials suggests the use of ornaments as markers of social inequality in these societies (Vanhaeren and d'Errico 2005). The use of ornaments alongside other practices of body adornment for this purpose was accentuated by the ninth stage: the production of ornaments by specialized artisans for members of different social groups following the emergence of production economies and the accumulation of wealth. This process resulted, on one hand, in the development of increasingly intricate visual codes and the diversification of ornamental functions, and
on the other hand, in the transformation of ornaments into valuable commodities to be traded,
exchanged, or inherited. The final, tenth stage corresponds to the industrial production of ornaments,
which allowed a wide range of members of a society to have access to various ornaments, including to
beads that were given special significance in the previous steps, either because of their rare raw
materials or the special skills required for their production. This stage is achieved through the
industrial production of beads made from synthetic gemstones, pearls, amber, and similar materials.

The proposed evolutionary scenario is limited to considering technologies associated with the cultural
transformation of the body that leave tangible traces in the archaeological record. Although this
approach allows us to rely on solid, dated evidence, it is evident that these practices were likely
complemented, possibly from very early on, by a range of other expressions, such as gestures, dance,
mutilation, scarification, tattooing, and clothing.

This stepwise progression is mirrored by the increasing artificialization of ornament shapes: from
unaltered natural items, culture progressed to natural objects transformed in various ways through
perforation, coloring, and arrangement. This was followed in subsequent periods by a transition to
entirely artificial geometric shapes and colors, accompanied by a proliferation and standardization of
those shapes and colors, without necessarily an abandonment of the natural objects used in previous
stages.

Evolutionary and Cognitive Implications

Our attempt to outline an evolutionary scenario for the culturalization of the body that incorporates
both permanent and temporary modifications still has grey areas, affecting in particular, as one would
logically expect, the practices that are less likely to leave a trace of themselves in the archaeological
record. Despite uncertainties, however, this scenario highlights a process that lasted for at least
500,000 years, and probably longer. A key factor that may have triggered this process was, as
mentioned in section “Clothing and Needles” in summarizing the available data on the origin of
clothing, the Middle-Upper Pleistocene transition. This crucial moment in the evolution of our planet’s
climatic system led to climatic changes of higher amplitude, combined with ice ages of unprecedented
severity. The amplitude and pace of these events meant that attempting to cope with them by organic
evolution through natural selection or epigenetics alone was a perilous and ineffective undertaking.
Some human populations handled the new challenges by developing and accumulating cultural
innovations, including clothing, which enabled them to deal with the harsh climates and expand their
niche rather than tracking their ecocultural footprint, that is following the displacement of the
ecological systems they were more adapted to, and abandoning previously colonized regions. In this
context, it is not surprising to discover that this niche-building process appears to have begun with
essentially utilitarian practices, designed, for example, to protect the body against ultraviolet radiation,
low temperatures, wind, humidity, parasites, and infections.

It is hard to say how we moved from this situation to one in which these practices also became a way
to assert individual identity and a source of information about social and group affiliation. Information
from the archaeological proxies explored in this chapter, however, indicates that this process
developed over a long period, and that some populations, including those living in the African
continent, independently invented or adopted practices hundreds of thousands of years before others.
This process cannot therefore be explained as the consequence of an abrupt change corresponding to a
cognitive revolution that was linked directly to the emergence of our species in Africa.

The gradual increase in the quantity of ochre used at African Middle Stone Age sites and the gradual
diversification of ochre types and complexification of the technologies applied to ochre modification
and storage, on the one hand, and the absence of ochre in vast areas of Africa and Eurasia or the discontinuity in its use, on the other hand, suggest that this process was gradual and probably did not involve all human cultures for some time. What is clear, as evidenced by the appearance of ornaments covered in ochre or whose color has been modified with heating techniques, as well as by the divergence between head and body lice, adapted to clothing, is that from 160 to 140 ka, and perhaps earlier, the culturalization of the body must have reached a tipping point in certain cultures, leading to the definitive integration of clothing into cultural systems and to a symbolic transfiguration of the body through the use of ornaments. This transition marked a decisive move toward the creation of a niche that was primarily defined by both its cultural and environmental dimensions. The numerous subsequent steps can be envisioned as cultural exaptations that enabled new and increasingly more complex messages to be communicated to oneself and others. The underlying logic that guided this stepwise progression from the outset appears to be a gradual increase in the complexity of body culturalization technologies, alongside the investment of increasing time and effort in the acquisition and transformation of different types of raw materials to signal more and more complex social statuses and cultural affiliations. Such exaptations certainly entailed more refined and demanding cultural transmission strategies to facilitate the transfer to new generations of the knowledge necessary to implement these practices (Colagè and d'Errico 2020).

The use of the body itself as a raw material that can be modified by shaping it, removing parts of it, or permanently adding pigment and objects to it using dedicated techniques has probably followed the same logic of increasing technological complexification. It is plausible that the process we have identified, though focused on personal ornaments and ochre use, also sheds light on the evolution of other body culturalization practices. It is foreseeable that these practices, too, advanced toward increased complexity in terms of techniques, symbolic codes, and the involvement of specialized individuals to ensure their continuity and the potential for further development. Contrary to what a taphonomically biased archaeological record may lead us to believe, we may expect that practices linked to skin cutting and scarification were the earliest to emerge. Humans had already mastered this technology by using it to process the carcasses of prey and by using stone tools for other functions. In addition, the need to treat injuries must have already given humans means to reduce the chance of infections. Evidence indicating that permanently or semipermanently disabled individuals were taken care of by the community goes back to at least 400 ka (Gracia et al. 2009; Bonmatí et al. 2010) if not much earlier (Lordkipanidze et al. 2005), and a growing body of evidence suggests that Neanderthals were mastering the use of medicinal plants (Hardy 2022). This evidence entails that human societies of those times possessed practices with which to face the eventual health problems consequent to the production of permanent body modifications. Since effective cultural-transmission strategies to pass this body of knowledge through generations were presumably already in place, the exaptation leading to a change of function and the transfer of this knowledge to the human body must have been, technically, a quite trivial affair. Considering the similarity in color of ochre, blood released by prey, and human blood and the fact that scarification often does not reduce individual fitness, there is no reason to think that this type of permanent body modification may have not been a feature of cultural systems characterized by a habitual use of ochre, after 160 ka and probably earlier. One may even posit that the gradual increase in ochre use coevolved with practices involving permanently leaving scars on the skin produced by the releasing of blood. It is logical to think that the practice of inserting pigment into the body by rubbing it into open wounds or tattooing, perhaps following the practice of rubbing medicinal products on wounds to prevent infection, developed after cutting and scarification.

Although it is difficult to predict when this happened, it is interesting to notice that scarification and, even more so, tattooing are techniques that allow the permanent inscription of a visually perceptible
design into the human skin. Apart from isolated instances, engraved or painted designs do not appear
in the archaeological record before 120 ka. Unlike with paintings, it is difficult to attribute the virtual
absence of earlier engravings at sites older than 120 ka to taphonomic reasons, which may suggest
scarification and tattooing were not implemented or were only sporadically implemented before that
date.

The shaping of osseous raw materials such as bone, antler, tooth, and ivory using techniques specifically designed for that purpose, such as scraping, grinding, and gouging, does not appear in the archaeological record before 90–80 ka (Bouzouggar et al. 2018; d’Errico et al. 2022). It is reasonable to think that these techniques must have been applied to bone tool shaping before being exapted to, for example, teeth shaping, and that this transfer may have been preceded by technically simpler, but no less traumatic, dental evulsion carried out by knocking teeth out with rocks or similar techniques.

There is reason to believe that practices involving bone shaping, such as arti cial skull deformation and
foot binding, may have been among the most recent to develop. Technically, these practices may be
perceived as relatively simple and less risky, because they generally only involve the use of devices to
apply pressure to parts of the body, but they require a continuous and long-term commitment by
relatives or recognized experts. This includes care during the healing process, the maintenance of the
modification equipment used, and strong adherence to cultural practices. Unlike most other permanent
changes, which are inflicted at one or more specific points in an individual’s life, bone shaping are
done over long period of time and begin in early childhood. They are generally intended to mark the
individual’s membership in particular social groups in complex societies with enduring ranking
systems.

The Perception of the Culturalized Body and Its Possible Evolution

The integrated scenario I have attempted to outline raises the question of how human cognitive
capacities have been able to cope with learning and decoding the increasingly complex messages
conveyed by the accumulation of practices linked to permanent and temporary modifications of the
body. It is reasonable to think that this process necessitated the development of dedicated cognitive
functions and neural circuits that would enable the deciphering of the increasingly complex messages
these practices conveyed. In a recent study (Salagnon et al. 2024), participants were asked to assign
social roles or statuses to faces adorned with paintings or beads, or both, while their brain activity was
monitored using functional magnetic resonance imaging (fMRI). Results showed the activation of a
complex network, including the occipito-temporal regions, the salience network, the prefrontal cortex
and the intraparietal sulcus. The fusiform gyrus helps us recognize faces as social indicators, while the
orbitofrontal cortex evaluates social status. The salience network integrates relevant information,
aiding in social categorization. The hippocampus and parahippocampus aid memory and associative
abilities, helping us recall past experiences to make social judgments. Interestingly, the inferior frontal
gyrus, which is linked to language, is active when a subject is interpreting facial ornaments
symbolically, suggesting a connection between language comprehension and symbolic interpretation.

Analyzing resting-state functional connectivity revealed complex interactions among these brain
regions, highlighting their collective role in interpreting facial ornamentation for social status
attribution.

Was this complex network fully functional when the first archaeological evidence of culturalization of
the human face was recorded or did it result from a gradual process of integration and
complexification? The gradual complexification and patchy emergence of body culturalization
practices over the last 500,000 years suggest a scenario of increasing but asynchronous integration of
the brain areas involved in social status recognition. This growing integration allowed the decoding of
increasingly complex symbolic codes, supported by more demanding technologies for face and body
adornment. The interplay between cultural and biological mechanisms likely drove this process, with
individuals who are gifted in acquiring, decoding, and creating these symbolic adornments. The
messages had selective advantages that favored the permanent inscription of a more integrated
connectivity in the brain. It has been suggested that in a comparable way a progressive co- option of
brain regions would have enabled the development of increasingly complex tools (Stout et al. 2015;
Hecht et al. 2015; Putt, Wijeakumar, and Spencer 2019).

The period between 140 ka and 70 ka may have been crucial for the fusion of cultural practices,
because it marks the time when the use of red pigments became widespread at Middle Stone Age sites
in Africa, and when marine shell beads first appeared in North Africa, the Middle East, and Southern
Africa. The diversity of colors, shapes, and technologies suggests an evolution of practices that
enabled individuals to convey information about their social roles through more complex, commonly
understood symbolic systems. It's therefore plausible that by 120 ka to 70 ka the human brain had
developed the capacity to process and make sense of these signals.

## Acknowledgments

The author's research is funded by the European Research Council for the project Evolution of Cognitive Tools for Quantification (QUANTA), No. 951388; Initiative d'Excellence IdEx, University of Bordeaux, Talent program grant #191022-001; French government in the framework of the University of Bordeaux's IdEx "Investments for the Future" program/GPR "Human Past" and the Research Council of Norway, Centre of Excellence (SFF), Centre for Early Sapiens Behaviour, SapienCE grant # 262618.

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