PHILOSOPHICAL TRANSACTIONS
OF THE ROYAL SOCIETY B

BIOLOGICAL SCIENCES

Scaffolding Minds: Human Collective Intelligence through
Space, Body and Material Symbols

<table><tr><td>Journal:</td><td>Philosophical Transactions B</td></tr><tr><td>Manuscript ID</td><td>RSTB-2024-0448.R1</td></tr><tr><td>Article Type:</td><td>Research</td></tr><tr><td>Date Submitted by the Author:</td><td>n/a</td></tr><tr><td>Complete List of Authors:</td><td>d'Errico, Francesco; Bordeaux 1 University, Archeological Science; University of Bergen Faculty of Humanities, Centre for Early Sapiens Behaviour (SapienCE), Department of Archaeology, History, Cultural Studies and Religion<br>Meneganzin, Andra; KU Leuven, Institute of Philosophy<br>Colagè, Ivan; Pontifical University of the Holy Cross, Faculty of Philosophy and DISF Research Centre</td></tr><tr><td>Issue Code (this should have already been entered and appear below the blue box, but please contact the Editorial Office if it is not present):</td><td>COLLECTINTELL</td></tr><tr><td>Subject:</td><td>Behaviour < BIOLOGY, Cognition < BIOLOGY, Evolution < BIOLOGY, Palaeontology < BIOLOGY</td></tr><tr><td>Keywords:</td><td>Epistemic niche, Niche construction, Exosomatic storage, Epistemic artefacts, Externalized cognition, Cognitive archaeology</td></tr></table>

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# Scaffolding Minds: Human Collective Intelligence through Space, Body and Material Symbols

Francesco d'Errico ¹⁻², Andra Meneganzin³, Ivan Colagè⁴

¹ Université de Bordeaux, UMR 5199 PACEA, Batiment B2, Allée Geoffroy Saint Hilaire, CS 50023 F - 33615 PESSAC CEDEX. francesco.derrico@u-bordeaux.fr, https://orcid.org/0000-0002-2422-3079

² Centre for Early Sapiens Behaviour (SapiENCE), Department of Archaeology, History, Cultural Studies and Religion, University of Bergen, Norway.

³ Institute of Philosophy, KU Leuven, Andreas Vesaliusstraat 2, 3000 Leuven (Belgium) andra.meneganzin@kuleuven.be, https://orcid.org/0000-0003-3641-3803

⁴ Pontifical University of the Holy Cross, Faculty of Philosophy and DISF Research Centre, P.zza di Sant'Apollinare, 49 - I-00186, Rome, Italy. i.colage@pusc.it, https://orcid.org/0009-0008-6824-4956

## Keywords

Epistemic niche, Niche construction, Exosomatic storage, Epistemic artefacts, Externalized cognition, Cognitive archaeology

## Summary

Human collective intelligence—the capacity of groups to solve problems, make decisions, and acquire knowledge beyond individual capabilities—is here understood as an emergent phenomenon that evolved in our lineage from a distinct trajectory of epistemic niche construction (ENC), and progressively sustained the latter. Humans systematically alter their informational landscapes in materially visible ways by creating enduring spatial and artefactual scaffolds for improved cognitive performance and social coordination. In this paper, we propose a set of criteria to define ENC and track its emergence in the archaeological record. These criteria highlight the importance of persistent, publicly accessible, and evolutionarily incremental modifications that sustained behavioural coordination among individuals in space and time. We apply this framework to three major domains of material culture: the structuring of space for collective action, the culturalization of the human body, and the emergence of exosomatic artefacts to store coded information. We argue that these practices did not merely externalise knowledge but progressively transformed material culture and environments into targeted epistemic infrastructures able to scaffold and amplify group-level performances characteristic of collective intelligence, thus shedding light on the evolution of human cognition and social organization.
# 1. Introduction

Collective intelligence generally refers to the cognitive performance of a group of agents that exceeds what would be achievable by any individual alone [1–3]. The emergence and evolution of collective intelligence (CI) and cognate phenomena, such as collective knowledge and action, have garnered increasing attention in both human and animal studies over the past decades. Core research questions span from the mechanisms allowing groups to achieve shared levels of information, the maintenance of collective processes over time, the excogitation of novel behaviours, and the expansion of shared informational pools [4]. Our focus in this paper will be on the collective structuring of *information* supporting group-level decision-making and information retrieval as tracked via the archaeological record.

A few notes of caution are due. First, ‘information’ is a notoriously vexed notion across a plethora of fields, not least in the evolutionary sciences. While commitments to formal definitions of information and their application across disciplinary boundaries typically lead to trouble [5], we do not wish to leave our usage entirely unspecified either. In this paper, we are interested in the *material* vehicles of information in the environment (broadly understood) that are relevant to a given organism and that, once perceived, can influence its present or future behaviour. Thus, information is always information-for-an-agent (in a relative sense). Further, our use is compatible with broader conceptions of information as anything that ‘reduces uncertainty’ [6,7], thereby guiding and constraining appropriate behaviour within specific circumstances. Accordingly, in what follows, “information” and, more specifically, “informational landscape” refer to material features of an environment that function as cues or signals for specific organisms—that is, elements that can be detected given their perceptual and cognitive capacities and that can consequently scaffold their behaviours. Second, the structuring of information can be collective in two ways. In one sense, it is the consumers of information who form a collective; in another, the structuring action is performed by a collective. In our account, both dimensions will become relevant, although we note that even in what appear to be individual acts affecting information availability (e.g., the production of an artefact), these often draw upon collective skill and competence.

Here, we propose a chronologically structured scenario for the emergence and intensification of resources scaffolding collective intelligence in human evolution. To do so, this paper will leverage an *epistemic niche construction* (ENC) approach—which we will explicitly characterize—to three categories of archaeologically detectable behaviours and artefacts as gradually acquiring collective informational roles: (i) the alteration or construction of spatial settings to support synergetic thinking and action (e.g. structured exchange corridors, domestic spaces and underground environments); (ii) the implementation of technologies mediating the culturalization of the human body (e.g. clothing, pigments and personal ornaments), (iii) the development of exosomatic artefacts to store coded information (e.g. artificial memory systems, writing, number symbols). These three categories of artefacts capture three trajectories of the evolution of materially scaffolded human collective intelligence, each likely taking off at different temporal horizons. In keeping with a niche-
construction approach, we frame the evolution of such artefacts within a co-constructive and coevolutionary scenario that, coupled with other eco-cultural evolutionary signals, supports inferences to a stronger epistemic function of those materials in handling coordination problems within informationally demanding societies.

In light of calls for bridging work on collective intelligence and culture [4], it is important to clarify how our framework relates to existing approaches within cultural evolutionary studies [8–12]. Following traditional definitions of ‘culture’ as an individual-level phenomenon [13], cultural evolutionists are typically interested in modelling how ideas and practices spread through chains of individual transmission, thereby explaining cumulative change and population-level distributions of cultural traits. By contrast, our focus on collective intelligence highlights phenomena that cannot be reduced to individual components alone, nor merely treated as their aggregates. We argue that ENC creates epistemically structured environments—through spaces, bodies, and artefacts—that enable distributed, synchronised, and scalable forms of reasoning and memory. These infrastructures clearly support and bias cultural transmission but also transform it into genuine collective intelligence, understood here as a higher-level phenomenon emerging from group-level interactions with epistemic environments. Importantly, our focus shifts the emphasis away from the pivotal role attributed to high-fidelity transmission mechanisms and toward the multiple, graded ways in which an epistemically structured environment can scaffold intelligent action. These latter forms likely have deep evolutionary origins, with high-fidelity transmission building upon a pre-existing epistemic platform. To see this, in the next section, we analyse and expand upon the ENC framework before applying it to discuss archaeological traces.

## 2. Constructing the informational niche

Humans and non-human animals actively shape the environment by transforming habitats, modifying resource availability and influencing the distribution of other species within their ecosystems. Examples range from beaver dams to the sophisticated agricultural practices of leafcutter ant colonies and the habitat alterations made by burrow-dwelling animals [14–16]. When organism-mediated environmental modifications lead to persistent effects on selective pressures, evolution by niche construction occurs [17–20]. Niche construction is typically defined by the following criteria [20]:

1. An organism must significantly modify environmental conditions;

2. The organism-mediated environmental modifications must influence selection pressures on a recipient of niche construction;

3. There must be a detectable evolutionary response in a recipient of niche construction that is caused by the environmental modification of the niche constructor.

It has long been noted that much *human* niche construction is *epistemic*, meaning it is directed at altering the informational character of environments, often in ways that make important
features salient [21–25]. This, in Sterelny’s words, involves “making cognitive tools and assembling other informational resources that support and scaffold intelligent action” [23]. While humans are epistemic engineers *par excellence*, animals too engage in activities that can be interpreted as epistemic actions [22,26] with actual or potential niche-constructing effects. These include trail-marking behaviour in ants [27], spider webs building informationally overloaded spider worlds [28], or nest concealment in passerine birds—a form of epistemic sabotage, complicating information retrieval for predators [29,30]. Crucially, some of these activities employ features of the environment or tools as external ‘props’ to informational and cognitive processing [26,31].

Expanding on the three criteria for general niche construction mentioned above, we propose five explicit criteria for *epistemic* niche construction (ENC).

1. An organism must significantly alter the informational conditions of its environment. ENC implies that organisms must modify how information is made available, structured or encoded in the environment. These modifications can range from incidental by-products of ecological behaviours (e.g., termite mounds signalling stable microclimates) to purposeful epistemic actions enhancing environmental predictability or guiding behaviour (e.g., trail marking, tool-mediated informational storage, or symbolic encoding). Such informational alterations can vary in resource investment, ranging from low-effort cues (e.g., scent marking) to resource-demanding constructs (e.g., concealed nests).

2. Organism-mediated informational modifications must persist enough to scaffold behaviour or cognition. For ENC to be effective, environmental changes must produce durable modifications allowing organisms to access, interpret, or build upon the altered information. Persistence may result from structural stability (e.g., nests, webs) or active maintenance (e.g., refreshed scent trails, territory marking). Potentially informative but fast-fading cues are unlikely to make a significant contribution to ENC.

3. Informational modifications must be usable and retrievable by multiple agents. For ENC to extend beyond individual benefit, the modified information-bearing features of the environment must be publicly accessible, interpretable, and reusable by others, within or across generations. The salience of these features is dependent upon the perceptual and cognitive capacities of the organisms. This allows shared epistemic ecologies where multiple agents rely on, maintain, and potentially expand common informational resources.

4. Informational modifications must generate evolutionary feedback in at least one recipient population. ENC implies affecting evolutionary trajectories beyond immediate utilitarian advantages. ENC drives evolutionary change when a modified informational environment leads to heritable changes in cognition, perception or behaviour.
Importantly, this does not necessarily presuppose genetic heritability: ENC can affect
individual and collective traits through different channels of inheritance: epigenetic,
ecological, social, and cultural.

5. *Informational modifications must support behavioural coordination among individuals (either synchronically or diachronically).* ENC facilitates social coordination in space and time by enabling organisms to align with past or future actions, or to synchronise with others' behaviour. These modifications can structure joint attention, store group-level knowledge, or regulate task sequences. In the context of 'swarm intelligence' research, this is sometimes conceptualised as "marker-based stigmergy" [32,33].

Let us unpack these. Criteria 1-3 clarify the presence of epistemic niche construction, criterion 4 captures its evolutionary consequences, and criterion 5 its relation to collective intelligence (as also discussed in the Introduction). Criteria 1 and 2 allow for a graded reading. This is deliberate and critical. Modifications of environmental features sustaining ENC are not always produced intentionally (criterion 1). A simple example is the formation of paths through woods, hills, or clearings, where the repeated passage of individuals unintentionally induces others to follow the same route (this is also an example of a maintained albeit perishable scaffold, criterion 2). Other modifications, however, are purposefully created, as captured by the distinction between 'cues' and 'signals' [7]. Symbol systems – understood here as relying on degrees of conventionality and shared informational background among users – are perhaps the clearest examples of intentionally produced cognitive or epistemic scaffolds supporting the cognitive performance of multiple individuals who learn and use them.

Criterion 3 points to epistemic activities involving more than one individual. This has important implications. As with criterion 1, resources scaffolding cognition in a group can range from various forms of affordances [34] that are already part of the furniture of the environment to highly targeted systems that maximize the fit to an organism's cognition and knowledge [35–37]. Crucially, however, as Sterelny [22] emphasised, epistemic tools for joint use are typically suboptimal for any individual user. Theorists of extended and scaffolded cognition have long discussed practices that reduce memory and perceptual load [38,39], improve efficient workspace engineering [40], and bring about alterations of learning environments to support future generations' learning [22]. However, retrieving information or correctly interacting with spaces and devices meant to "speak to the group" requires adequate cognitive and sociocultural support: the benefits of cognitive offloading, scaffolding, and perceptual facilitation are only possible when an adequate informational and cognitive base is already in place. As we will see below in discussing the structuring of space, the culturalization of the body and the production of exosomatic artefact to store information, such intentional environmental modifications with epistemic and cognitive import may also impact the very perceptual and cognitive capacities of their creators and learners, as well as those of subsequent generations (criterion 4) [41,42].
Criterion 5 is crucial for understanding how ENC underpins the emergence and development of human CI. Epistemic modifications enable cognitive labour to be distributed, synchronised, and scaled. This socio-temporal scaffolding transforms individual efforts into group-level cognition, facilitating complex collaboration, coordination, division of labour, and accumulation of collective memory. In this sense, ENC is not merely a background condition for cognition; it also *actively* engineers the temporal and social architecture through which “group minds” emerge and operate. The more robust and integrated these externalised supports, the greater the potential for sustained, large-scale collective reasoning, planning, decision-making and cultural innovation. Put this way, ENC is also likely to be impacted by CI: with improved higher-level cognitive performance, it is plausible that better skills emerged for modifying the very conditions that enabled such performance. In our view, this already carries an important implication: alterations to the informational profile of spaces, environments, or objects should have been advantageous to hominin groups *well before* the proliferation of symbolically-soaked worlds. The use of specially designed devices to code and store information is a non-trivial activity that appeared relatively late in the hominin evolution. As we will argue in the context of ancient spatial settings, it is probable that epistemic functions were initially superimposed on other artefactual and spatial functions — artefacts were epistemically co-opted, or *'culturally exapted'* [41,42] — before highly designed informational devices could emerge.

The long-term coevolution of hominin cognition, sociality, and material engagement unfolded within changing ecological and climatic conditions, shaping the scope and persistence of epistemic modifications. Figure 1 synthesises the interplay between three critical macro-evolutionary variables over the last 3.3 million years: global climatic fluctuations, temporal distribution of major hominin taxa, and cumulative terrestrial area colonised by hominins. This visualisation highlights two core features relevant to epistemic niche construction. First, the shift from 41,000-year to 100,000-year glacial cycles (after ca. 1 Ma) implies increasing ecological instability and unpredictability, which likely intensified the pressures for innovations in social learning, spatial planning, and informational offloading. Second, the progressive and punctuated expansion of the hominin geographic niche — initially within Africa, later across Eurasia and finally into remote island environments — suggests that material and symbolic scaffolds for decision-making and collective action became increasingly critical for adaptation. These offer a deep-time backdrop against which the three archaeologically visible behaviours this paper focuses on (spatial setting, body culturalization and exosomatic artefacts) can be understood as adaptive responses to expanding informational demands.

### 3. The epistemic structuring of space

Human societies construct culturally meaningful spatial divisions, often marked by material cues or the symbolic appropriation of ecological features (Figure 2, top panel). These spatial
demarcations distinguish inside from outside, private from public, domestic from ritual, and
ordinary from extraordinary. Spaces in human contexts are not only inhabited but encoded
with normative expectations—what one can do in a space is contingent on its cultural framing.
This transformation of space into an epistemically charged medium is fundamental to the
evolution of collective cognition, as spatial organisation imposes salient scaffolds for group-
level decision-making, behavioural coordination, and shared memory.

The origins of such practices are likely ancient. The earliest evidence for this transformation
can be found in the emergence of culturally mediated “inside” spaces [43]. We will briefly
revisit core shifts in the character of domestic spaces, highlighting how they trace ENC and
impact CI.

Between 2.6 and 1 Ma, archaeological sites appear primarily organised around activities
related to food procurement and processing — stone knapping, carcass processing, and food
sharing — with limited evidence for spatial differentiation (e.g., clusters of artefacts and
debris). Around 1 Ma, the intermittent use of fire, documented at sites such as Gesher Benot-
Ya'akov, signals the growing value placed on modifying and controlling the immediate
environment for social and epistemic purposes. By 400 ka, the regular control of fire and the
appearance of structured hearths mark a significant shift in the structuring of domestic space.
Between 400 ka and 250 ka, more sustained occupation patterns emerge with the
organisation of hearths into stacked features. It is at this stage that significant opportunities
for social and epistemic gains come online — from storytelling to the cooperative effort
required by fire management and early cumulative cultural processes. Possible ritual or
symbolic spatial practices also appear in this timeframe, including the deposition of bodies at
Sima de los Huesos (ca. 450 ka). From 250 to 125 ka, pest-repellent bedding (e.g., at Border
Cave, [44]), dietary diversification, and subterranean seemingly symbolic spaces (such as
Bruniquel Cave at 176 ka) suggest increasing engineering of complex socio-domestic ecologies
(possibly with aesthetic character too [45]). Thus, from 250 ka onwards, the scaffolds
supporting novel social and epistemic gains show not only stability but also signal
complexification. Between 125 and 40 ka, spatial organisation becomes more complex, with
constructed hearths, differentiated areas for distinct activity (e.g., sleeping, working, waste
disposal), and signs of ritual deposition (inhumations). Spatially segregated zones for symbolic
expression emerge, including cupules or geometric motifs at South African and European sites
(Klipbak 1, Nchwaneng, Rhino Cave, Roche-Cotard, Gorham's Cave, Ardales). During the Upper
Palaeolithic (40–10 ka), these trends culminate in highly structured domestic spaces, the
regular use of artificial structures with distinct hearths, decorated interiors, and tent rings for
attaching skins, suggesting purpose-built camps. Symbolic spaces are now consistently
separated from domestic areas and often located in deep cave chambers, likely facilitating
shared cognitive and emotional experiences. The proliferation of cave art across Eurasia
exemplifies the diversification of aesthetic and symbolic spatial practices and their integration
into broader informational ecologies.
Culturalization progressively affected “outside” spaces as well. While hearths, ritual deposits, and domestic organisation scaffold collective cognition within inhabited spaces, we suggest that the procurement of distant resources epistemically structures external space by establishing, recognising, and negotiating territorial boundaries and their permeability. These practices thus extend the informational landscape of groups beyond the domestic sphere and integrate external territories into shared epistemic frameworks. In traditional societies, border-crossing between groups is often managed through cultural strategies such as message sticks (e.g., among Australian Aborigines) or the negotiation of neutral meeting zones. Such ethnographic examples are, of course, not to be read as establishing direct continuity with Palaeolithic practices but serve to illustrate mechanisms of boundary permeability through shared conventions. These strategies cannot plausibly depend on individual initiative alone but presuppose collective processes, social conventions, and perhaps even intergenerational transmission of norms.

Early direct archaeological evidence of spatial reconfigurations remains sparse [46–48]. In the Schelelean, raw material transport over distances up to 20 km is attested but may reflect movements within a single group’s territory. With the onset of the Middle Stone Age in Africa and the Mousterian in Eurasia (ca. 350 ka), we observe more compelling signs of intergroup interactions, including exchanges of lithic and ochre materials over distances of 25–50 km (rarely up to 300 km). The presence of non-utilitarian manuports, including quartz crystals and fossil shells, further supports the hypothesis of normative spatial practices from this period onward [49]. The procurement of ochre across distances of 25–80 km is documented as early as 110 ka (e.g., at Skhul and Diepkloof), while the transport of marine shells over 90 km at Border Cave (South Africa) or 60 km at Grotte des Pigeons (Morocco) attests to the cultural valuation of distant resources. Evidence for occasional and opportunistic open-water movement to close islands appears around 200 ka [50]. By around 60 ka, systematic seafaring is attested by the colonisation of Sahul and offshore island chains, suggesting the emergence of planned, repeated long-distance navigation strategies. From 40 ka onward, long-distance transfer becomes a defining feature of Upper Palaeolithic societies, with marine shell beads travelling up to 400 km [51], red deer canines covering distances of 250 km [52], and extensive use of non-local ochres.

In sum, the progressive structuring of both external and internal space played a pivotal role in hominin ENC. These practices not only shaped how knowledge was organized and transmitted but also enabled the emergence of increasingly complex, temporally extended, and collectively coordinated forms of cognition.

These practices meet criteria 1 to 3 for ENC, as they show informational modifications of the environment that endure in time and support collective use — from increasingly function-transparent interior areas to established corridors for material exchange. Moreover, the sustained process of spatial epistemic structuring of internal and external spaces likely exerted novel evolutionary pressures on occupants’ cognitive and behavioural performance, promoting novel forms of cooperation and ways of sharing informational resources (Criterion
4)—storytelling being a prominent candidate example. Criterion 5 appears to be at work as well, to the extent that those practices structured group-level coordination and progressively promoted distinction of roles and division of labour.

## 4. The culturalization of the human body

The culturalization of the human body refers to the processes by which individuals and societies imbue physical bodies with cultural meanings, values, and symbols (Figure 2, middle panel). This phenomenon encompasses both *transient* modifications—such as clothing, body painting, personal ornamentation, and hairstyles—and *permanent* alterations, including tattooing, scarification, dental and cranial modification, piercing, and even intentional, non-medical amputation [53]. It is important to note, however, that not all of these modifications need to function as fully symbolic expressions in the richer sense in use among semioticians. In many cases, they facilitate social signalling or identity marking as ‘badges’ or ‘insignia’, without involving strictly conventional, abstract or referential meaning.

Human body modification is a cumulatively evolving, socially transmitted, and often value-charged practice. In many cases, it extends beyond adaptive or aesthetic functions by purposefully constructing visual epistemic environments through which individual and group identities are displayed and negotiated. It has long been suggested that body modifications can act as preconstructed channels of communication, reducing the cognitive load in synergetic interactions between individuals [54,55]. Such visual clues (clothing, hairstyles, body adornments) allow for quick assessment of others’ status and fast adjustment of behaviour, and facilitate more complex interactions (such as problem-solving or decision-making) as well as inferences about shared values, beliefs, customs, and norms. From an evolutionary perspective, the emergence of more and more complex societies brought about increasingly challenging coordination problems. It is easy to see how practices of body culturalization aid in these problems — by reducing uncertainty about roles, status, or affiliations, and streamlining inference into simpler perceptual tasks — thus reducing negotiation costs and freeing cognitive resources to navigate interactions in larger or more stratified groups. The logic, unsurprisingly, is one of mutual reinforcement: more complex social worlds made investment in bodily cues increasingly profitable, which, in turn, facilitated the development of a platform for more expanded, multi-layered interactions.

Body-culturalization practices are often transmitted across generations, thus fostering a sense of shared history and identity within a community. They serve to reinforce social bonds and cooperation within communities. This sense of belonging and social cohesion enhances collective intelligence by promoting collaboration, empathy, and a shared sense of purpose.

Recent interdisciplinary research has begun to explore the neural basis of the perception of culturalized bodies. In a recent functional magnetic resonance imaging study, participants
were asked to assign social status to human faces culturalized via painting, beads or a combination of both [56]. Participants' brains exhibited activity in an extended neural network, including visual, limbic, insular, and prefrontal regions. This suggests that the perception and interpretation of symbolic bodily cues engage integrated circuits for executive control, affective processing, and social cognition. These findings align with the hypothesis that the increasing complexity of body culturalization exerted selective pressures on cognitive functions, reinforcing neurodevelopmental pathways for bodily cue learning, memory, and interpretation. This is supported by extensive psychological and neuroimaging research on stereotyping and ethnic cognition, which demonstrates that bodily and visual cues trigger social categorisation and stereotype activation [57,58]. Such evidence reinforces the idea that culturalized bodies scaffold multi-layered social cognition, extending beyond individual recognition to group-level categorisation and norm enforcement. The coevolution of body culturalization and cognitive architecture thus represents a key feedback loop in the broader framework of ENC, through which bodily surfaces gradually became canvases for encoding socially meaningful information.

The culturalization of the human body unfolds along two interrelated dimensions in hominin evolution:

1. *Behavioural, social and symbolic* – the emergence of identity markers and socially structured bodily display;

2. *Technological* – the invention and use of tools and substances (e.g., needles and pigments) to alter, adorn, and signify the body.

In the following subsections, we describe four main, non-strictly sequential steps within this process by highlighting archaeological evidence along the above dimensions. A final subsection will discuss the relationships between body culturalization and ENC.

### (a) Clothes

Although early clothing leaves few direct archaeological traces, a growing body of indirect evidence strongly suggests its emergence during the Middle Pleistocene. From as early as 780 ka to 500 ka, we begin to observe cut marks indicative of skin removal, traces of use-wear on lithic tools associated with hide scraping, and the use of pointed stone tools interpreted as perforators. Particularly significant in Europe is the appearance — beginning at Schöningen (ca. 380 ka) and attested at several other more recent sites — of bone tools manufactured from ribs and long bone fragments, often interpreted as smoothers or “lissoirs” to soften skins [59]. Their standardised morphology and intensive use-wear suggest that such practices were increasingly formalised components of hominin adaptive strategies. Further indirect support for the widespread use of clothing comes from data on the genetic divergence of body and hair lice (at around 170 ka, [60]), implying the presence of clothing as a stable environment for lice. From approximately 100 ka onward, additional evidence emerges from sites such as Sibudu and Blombos Cave in South Africa, where bone awls appear and may have served in
hide piercing or sewing. Eyed bone needles — long viewed as a hallmark of Upper Palaeolithic innovations — appear first in Siberia and China around 40 ka (Figure 3) and only later in Europe, by approximately 26 ka [61]. Their emergence coincides with the appearance of regionally distinct traditions in personal ornamentation and clothing manufacture, suggesting an intensification of bodily culturalization. In the period preceding the earliest eyed needles in Europe, additional techniques for hide working and skin preparation appear to have been used. Recent research at the site of Canyars has demonstrated the presence of bone punching boards used to perforate hides, suggesting sophisticated sewing or fastening systems already in place by the Mid-Upper Palaeolithic [62]. Importantly, the emergence of clothing and hide-processing technologies does not strictly correspond to long-term reduction of temperature during the last 3 My (Figure 1). Rather, their development appears more closely tied to the need for adaptive flexibility in the face of rapid climatic oscillations and increasingly variable environmental conditions during the Middle and Late Pleistocene. In this view, clothing likely emerges not as a passive response to cold, but as a culturally driven strategy within the broader framework of niche expansion and epistemic niche construction, enabling hominins to externalise both ecological and social identity onto bodily surface.

### (b) Pigments and chromatic body culturalization

Pigments — particularly red ochre — represent another powerful means through which human groups began to inscribe cultural meaning onto the body. A comprehensive database of ochre use records 86 African Middle Stone Age (MSA) sites yielding ochre between 500,000 and 40,000 years ago [63]. From 160,000 years ago, the increasing frequency and complexity of ochre use suggest a shift from primarily utilitarian applications (e.g., protection from sunlight, antiseptic or antiparasitic use) to communicative practices, including body painting. At Blombos Cave (South Africa), ochre originating from different sources was knapped, pounded, ground, mixed with animal fat and stored in shell containers. Abstract engravings on ochre and the creation of ochre “crayons” to mark surfaces attest to the emergence of externalised graphic communication. Outside Africa, the earliest use of ochre is recorded around 380,000 years ago in sites such as Terra Amata in France and Maastricht-Belvédère in the Netherlands [64]. Later European Neanderthal sites demonstrate the use of both red and black pigments to mark shells, fossils, and cave walls. In the Near East and Africa, evidence for heat-induced colour transformation of yellow to red ochre further supports intentional crafting of visual effect. These developments signal the integration of pigments into emerging systems of communicative and eventually symbolic practices related to self-identity and inter- or intra-group identification — a form of epistemic marking through body modification.

### (c) Ornaments and the visual grammar of identity

Personal ornaments represent one of the most direct and durable traces of bodily culturalization. Perforated shells are documented in African and Near Eastern MSA contexts between 140 and 70 ka [65]. These early ornaments (Figure 3) generally involved a narrow selection of shell species, were often curated and subjected to surface modifications, including
heating and ochre-coating, to enhance chromatic properties and visual appeal. The evolution of ornament traditions reveals a long and multi-step process of gradual complexification.

The first stage may be represented by the habitual use of ochre, attested as early as 500 ka (as discussed in Sub-sec. 4.b). This was followed by the collection and curation of unmodified natural objects such as marine shells, selected for their colour, shape, or texture, and potentially worn or displayed without further alteration. As adornment practices developed, hominins began assembling series of homologous shells — often naturally perforated — into composite items (necklaces or pendants). Subsequently, perforation techniques were refined to enable greater control over bead production allowing for standardised arrangements and modular designs. These developments paved the way for the integration of heterogeneous ornaments into single decorative items, combining shell species of different sizes or visual properties. In some contexts, naturally modified or heavily eroded fragments were selected and repurposed as symbolic objects, further expanding the visual grammar of ornamentation.

By around 50 ka, fully manufactured ostrich eggshell ornaments appear in East and Southern Africa, marking a significant step in the intentional production of symbolic materials. The Eurasian Upper Palaeolithic, from circa 45 ka, saw an explosive diversification of ornament types and raw materials, with regionally distinct styles emerging across Europe and Southeast Asia. These regional differences appear to map onto cultural boundaries, functioning as ethnic markers within expanding social networks. As ornamentation systems became more elaborate, we begin to see evidence of increasing social stratification reflected in the differential distribution of rare or exotic ornaments. The use of imported materials or demanding production techniques signalled status or affiliation, suggesting the emergence of symbolic resources as vectors of social inequality. This development culminated in the appearance, with the emergence of production economies, of professionalised artisans producing ornaments for members of selected social classes, as well as the eventual mass production of imitation beads, which democratised access to visual symbols formerly restricted to elites.

Although this sequence is not strictly linear and likely involved regional variation, cultural transfers, and reversals, it nonetheless reflects a broader evolutionary trend: the gradual intensification of technological investment, symbolic abstraction, and transformation of ornaments into durable carriers of social, political, and economic conventional information. As such, the history of personal ornamentation offers a powerful lens to examine the interplay of materiality, identity, and ENC in the human past.

### (d) Permanent modifications

While practices such as painting and adornment have left substantial archaeological evidence, the earliest permanent bodily modifications are more difficult to trace. Practices such as tattooing, scarification, piercing and dental ablation typically leave ambiguous or indirect hints in the archaeological record. Nevertheless, isolated examples of dental ablation and intentional limb amputation — some dated to 40 ka — suggest that permanent body modification was part of a broader repertoire of symbolic expression and group differentiation
from an early stage [66]. Ethnographic parallels show that these practices could materialize a
wide spectrum of meanings, from initiation and status marking to ritualized responses to grief
— for instance, amputation as a form of mourning [67]. In this sense, they serve as durable
and publicly visible markers of cultural identity, signalling not only group membership but also
allegiance to shared traditions, beliefs, and values. They materially inscribe identity onto the
body, becoming a persistent epistemic medium for social information. These embodied signals
help reduce uncertainty in social interactions, facilitating recognition, norm enforcement, and
role differentiation. They contribute directly to the formation of epistemic niches that support
coordination, memory, and collective intelligence within human groups. Beyond their
communicative and symbolic roles, permanent modifications often serve as costly signals. The
endurance of pain, risk of infection, and the irreversible nature of such practices convey
commitment and loyalty. Importantly, the social, often ritualized, acceptance of practices that
modify the body in potentially harmful or dangerous ways creates powerful mechanisms of
intra-group cohesion. Participation in such acts by initiates, practitioners, or witnesses,
reinforces emotional bonds, aligns experiences, and contributes to shared narratives and
identities. In this way, permanent body modifications do not merely reflect social integration;
they actively produce and reproduce it. These dynamics enhance trust, solidarity, and
synergetic cognition, all of which are foundational to collective intelligence. The capacity to
think and act together, particularly in contexts requiring high levels of cooperation and long-
term planning, is strengthened when individuals bear visible, enduring signs of mutual
commitment inscribed on their bodies.

(e) Cultural body modification and ENC

Cultural body modification, considered in the four forms discussed above, appears to meet
our five criteria for ENC. The first step, hinged on clothing, opened the possibility of arbitrarily
modifying bodily appearance, independently of the reasons that triggered the behaviour.
Once pigments and ornaments were widely in use by 160-140 ka (if not earlier), human
environments were deeply characterised by social conventional information supported by
visual infrastructure (Criterion 1). The archaeological record attests to a considerable
investment in such practices and their cultural transmission [68]; hence, such informational
modifications of the environment were durable enough to shape and scaffold the behaviour
and cognition of those exposed to them (Criterion 2). The signalling conveyed through cultural
bodily modification addressed, and was naturally exposed to, a multitude of agents in the
shared visual environment (Criterion 3). In keeping with our Criterion 4, the cultural evolution
of pigments and ornaments use displays a cumulative pattern along which single practices and
artefacts were progressively combined, in turn prompting further evolutionary processes, at
both the behavioural-cultural [65] and the neural-cognitive levels [56]. Finally, Criterion 5 is
matched in that cultural body modifications specifically serve functions related to inter-
individual interaction, synergy, and coordination, as well as to within-group social
structuration and division of labour. This is particularly evident in the last considered step
(costly permanent body modifications).
5. From symbolic artefacts to Artificial Memory Systems: Externalising knowledge and scaling collective intelligence

All artefacts created by both human and non-human species embody a degree of exosomatic information; their materiality inherently suggests how they should be crafted or utilised, and the range of possible actions they afford. Artefacts infused with symbolic significance fall into a distinct category. They encapsulate information by forging an intangible connection between a natural or artificial characteristic and its cultural meaning. These include (Figure 2, bottom panel) both mobile functional objects (decorated tools, weapons, containers) and non-functional artefacts such as manuports (amulets, charms, figurines, etc.). While these objects are *not designed* to store structured or coded information, they often carry indexical or symbolic meanings that inform social cognition, regulate behaviour, and scaffold shared understanding. By embedding culturally salient meanings into material culture, such artefacts contribute to the externalization of knowledge. They function as engineered cognitive scaffolds — perceptually accessible structures that guide attention, evoke memory, and support communication across individuals and generations. Hence, they crucially contribute to ENC.

Such symbolic artefacts form the cognitive and cultural substrate upon which proper Artificial Memory Systems (AMSs) later emerge [69]. AMSs are devices *specifically conceived* to record, store, transmit and retrieve *coded* information outside the physical body. At present, these systems condition the lives of virtually all human beings and actively contribute to shaping our cognition, revolutionising the way we accumulate and transmit knowledge. Writing systems, numerical symbols, and digital devices are among the most widely recognised forms of AMS. Digital technologies, particularly through their integration into intangible information networks, now play an increasingly crucial role in how humans store, access, use and transmit knowledge across time and space.

Because AMS can maintain and recall information with high fidelity, they serve as repositories of collective intellectual effort, enabling successive generations to access, preserve, and expand upon accumulated knowledge. In doing so, they dramatically extend the temporal and social scale of information transmission, paving the way to knowledge processes that would be impossible through biological memory or oral tradition alone. AMSs extend cognitive operations into material substrates and play a foundational role in ENC, scaling up and temporally stabilizing collective intelligence. The most widely recognised AMSs — writing systems, numerical notations, and digital technologies — emerged in complex past societies.

Writing first appears in Mesopotamia around 3400 BCE, and then in Egypt (ca. 3300 BCE), Greece (ca. 1800 BCE), and China (ca. 1400 BCE) [70,71]. Other early AMSs include numerical clay tokens (ca. 9000 BP) and proto-cuneiform tablets (ca. 5500 BP), which exemplify the
gradual transition toward abstract, manipulable signs embedded in syntactic systems. AMSs are not exclusive to state-level societies. Across diverse small-scale societies — including hunter-gatherers, pastoralists, horticulturalists, and fishing communities — devices such as tally sticks, notched bones, knotted cords, carved staffs, and incised wooden objects have been used to track time, navigate space, record quantities, aid memory, and transmit ritual or mythological content [72–74]. This suggests that external symbolic storage systems predate writing, and that the human capacity for exosomatic information management has deep temporal roots. A growing body of archaeological evidence suggests that rudimentary AMSs were already in use by the late Middle Stone Age and early Upper Palaeolithic. Between 60,000 and 10,000 years ago, both in Europe and in Early Later Stone Age contexts in Africa, numerous artifacts bearing sequences of incised or engraved markings have been interpreted as early AMS instances [69,75,76].

Recent archaeological research further suggests that the emergence of numerical notations and number symbols arose not from a linear evolutionary process but through a series of cultural exaptations — instances in which pre-existing cultural practices were repurposed for novel cognitive or communicative functions. As proposed by d'Errico et al. [69], this process can be modelled as a five-stage scenario reflecting shifts in how hominins used material culture to externalize, manipulate, and transmit information. The foundation of this trajectory lies in the earliest (2.6 My) production of cutmarks during butchery, which contributed to the development of perceptual and motor skills necessary to create and visually process repeated incisions. The first cultural exaptation occurred when hominins began to intentionally produce parallel or equidistant marks that were no longer utilitarian but carried iconic, indexical, or symbolic meaning, as seen at Trinil and Bilzingsleben (Figure 4). A second exaptation is represented by artefacts such as the Les Pradelles hyena femur, where identical marks were created during a single session, and meaning was attributed to each individual unit rather than to the overall pattern — arguably the earliest evidence of numerical notation. A third stage is exemplified by the Border Cave fibula (44–42 ka), where similar marks were added over time, introducing the ability to incrementally record quantities. The fourth exaptation involved a more complex system in which multiple variables — morphology, spatial distribution, accumulation, and number of marks — were individually or jointly encoded, as in objects dated to 40–38 ka. This trajectory was accompanied by a clear tendency toward miniaturization, likely reflecting both the demand to record larger volumes of information and the consolidation of more abstract symbolic grammars. A fifth exaptation eventually led to the invention of number symbols proper — autonomous, manipulable units embedded in syntactic systems. This development, though cognitively transformative, occurred without requiring any biological change. Rather, it capitalized on the teaching strategies and neural plasticity already present in members of our genus.

This tendency echoes patterns observed in the evolution of writing systems. As d'Errico and Colagè [42] have argued, the emergence of such systems may result from a process of cultural exaptation coupled with cultural neural reuse, whereby neural circuits originally evolved for
other purposes are re-employed in culturally novel tasks, producing cascading effects on cognition and behaviour. The progressive externalisation, densification, and reorganisation of symbolic information in material media reflects not only cultural innovation, but also the recursive shaping of cognitive capacities through interaction with epistemically structured environments [77]. In this light, number symbols, writing and other AMSs should be viewed not as isolated breakthroughs, but as the outcome of a long-term process of ENC, through which increasingly complex information was stabilized, transmitted, and reconfigured. These systems became crucial scaffolds for the expansion of collective memory, planning, and inferential reasoning — key components of scalable human collective intelligence.

## 6. Discussion

Each of the three considered domains — the culturalization of space, the modification of human body, and the development of AMSs—played key roles in the evolution of human epistemic niche construction (ENC) and human collective intelligence. The chronological perspective of those three domains highlights the intensification of ENC across evolutionary time. We now pull the threads of the discussion of the archaeological evidence presented in Secs. 3-5 in the light of the five criteria proposed in Sect. 2.

**Criterion 1: Significant alteration of the informational conditions.** In all three domains, hominins introduced novel structures that reshaped information accessibility, interpretation and encoding. Spatial modifications involved meaning-charged partitioning of the environment, giving rise to cognitively and socially meaningful zones. Through clothing, ornamentation, and permanent modifications, the human body became a canvas for conveying and retrieving information about group identity, social status, or ritual role. AMSs provided a highly fine-tuned external and material medium for recording abstract information, decoupled from either immediate perception or biological substrate. In all cases, these transformations went beyond strictly functional modifications, reshaping or adding informational value to the environment.

**Criterion 2: Persistence of informational modifications, scaffolding cognition or behaviour.** The spatial modifications identified in the archaeological record are marked by long-term use and sometimes by repeated maintenance, indicating their durability and intended preservation as epistemic scaffolds. Similarly, body culturalization created stable, visually persistent markers that guided social interaction. AMSs, by definition, are designed for long-term information retention. The progressive miniaturisation and increasing density of markings in Upper Palaeolithic AMSs further support their use as cumulative, stable memory devices. Thus, in all three domains, material modifications persisted — or were purposefully crafted to persist — long enough to influence not just immediate actions but longer-term behavioural and cultural trajectories.
*Criterion 3: Informational modifications usable and retrievable by multiple agents.* Spatial arrangements reflect shared conventions on domestic or external zones for subsistence activities or ritual practices. Bodily signals are public and group-readable, serving as interfaces for collective knowledge, interaction and coordination. AMSs often exhibit standardisation and repeated use, implying they were most plausibly meant to be interpreted by others. In all three domains, there is a visible trend toward increasingly public, interpretable informational structures.

*Criterion 4: Informational modifications contributing to evolutionary processes.* All three domains, in virtue of the preservation or active maintenance of epistemic alterations, affected cognitive and behavioural evolution. The culturalization of the body possibly influenced mate choice, group cohesion, and social learning. Spatial structuring promoted new forms of social organisation and possibly the emergence of formalised roles. AMSs, by enabling external memory and cumulative elaboration, provided new conditions for cultural transmission and planning. Together, these practices reshaped developmental and social ecologies in ways that likely had lasting evolutionary consequences. Such consequences, as discussed, might have involved multiple channels of inheritance, with preliminary evidence on the effects of cultural practices on cognitive faculties and neural substrates [42,56].

*Criterion 5: Informational modifications supporting coordination among individuals.* Structured spaces facilitate shared routines and group-level task distribution. Body culturalization enables fast social categorisation, expectation formation, and the regulation of social interactions. AMSs facilitate the temporal decoupling of knowledge from individual memory, allowing for delayed retrieval, teaching, ritual performance, and institutionalisation of norms. All three domains show a progressive increase in the capacity to scaffold joint attention, collaborative action, and intergenerational knowledge transmission [68].

## 7. Conclusion

The five proposed criteria for ENC emphasize how human collective intelligence has developed through, and has likely reinforced, a niche-constructing process where aspects of the environment were gradually brought under control for their informational import. Crucially, the epistemic value of niche construction is not a mere by-product of functional modifications of the environment, but the result of long-term, collective and increasingly targeted investment. This aspect is mainly emphasized by the first three criteria, which are all met by the three cultural-evolutionary processes discussed in this article (culturalization of space and the body, and production of AMSs). The fourth criterion emphasizes the evolutionary role of such epistemic environmental modifications. It is worth noting that the trajectories of the three considered cultural-evolutionary processes suggest a cumulative evolutionary pattern with effects on the behavioural, cognitive and possibly neural constitution of the members of the groups that developed such practices. The fifth criterion directly links human ENC and its evolutionary consequences with collective intelligence. The
three cultural-evolutionary processes discussed above indeed have momentous social implications. Collective intelligence is not just social cognition, however. The key feature of the three considered processes is that they manipulate the informational dimension of the environment in a *generalised* manner. Space arrangements, bodily modifications and AMSs are often not instantiated in view of a specific, designated interlocutor; on the contrary, they embody information made available in the environment to a generalised audience. This, in our view, makes these processes true *loci* of genuine human collective intelligence.

Hence, viewed through the lens of our five criteria, the archaeological record discussed in this paper offers strong support for interpreting the culturalization of space and body, and the development of AMSs as converging trajectories of ENC. Together, these processes illustrate a broader evolutionary trend: the progressive dependence of cognitive functions on stable, interpretable, and socially shared material structures. This transformation not only enhanced individual cognition but enabled the emergence of a typically human collective intelligence — a higher-level phenomenon emerging from epistemically engineered environments recursively shaped by cultural practices. The archaeological record thus provides a unique window into the deep-time coevolution of materiality, meaning, and mind. Our framework demonstrates a progressive convergence of environmental, bodily, and artefactual epistemic resources. It supports the interpretation of hominin material culture as a medium of cumulative, collective cognitive scaffolding — a recursive dynamic that characterises human ENC and underpins the emergence of large-scale collective intelligence. Notably, while the expansion of the hominin geographic range accelerates markedly from ca. 2 Ma onward (Figure 1), this process is not consistently accompanied by robust archaeological signatures of ENC (Figure 2). Many newly colonized regions appear archaeologically “thin” with regard to structured spaces, symbolic artefacts, or evidence for shared informational scaffolding. This disparity suggests that geographic expansion alone does not necessitate epistemic restructuring; rather, it may precede or even occur independently of the widespread externalisation of information into material forms.

Understanding when, where, and why ENC intensified requires attention to contextual thresholds — ecological constraints, population density, intergroup interaction, and cognitive-cultural tipping points— that modulate the visibility and function of informational artefacts within the archaeological record.

## Acknowledgements

We thank Cathal O'Madagain and Andrew Whiten for inviting us to present a paper at the memorable Collective Intelligence workshop held at the School of Collective Intelligence of the University Mohammed VI Polytechnic in Rabat on 15-16 May 2024, and for making the event a unique opportunity for interdisciplinary exchange. Moreover, we are grateful to the audience of BSPS 2025 in Glasgow for productive feedback on aspects of this work. We also thank the Reviewers for their constructive engagement with the manuscript. Their comments
helped us refine the manuscript to better serve an interdisciplinary readership. Finally, we
thank Lila Geis for help in formatting the reference list. Research of FD and IC is supported by
the Grand Programme de Recherche ‘Human Past’ of the Initiative d’Excellence (IdEx) of the
University of Bordeaux. FD research is supported by the European Research Council (Synergy
Grant for the project Evolution of Cognitive Tools for Quantification [Quanta], no. 951388);
the Research Council of Norway through its Centres of Excellence funding scheme, SFF Centre
for Early Sapiens Behaviour (SapiENCE), project number 262618; the Talents Programme of
the University of Bordeaux (grant number: 191022_001) and the Région Nouvelle Aquitaine
Threads project. AM gratefully acknowledges funding from the Research Foundation –
Flanders (FWO), Grant No. G070122N. IC acknowledges an internal research grant by the
Pontifical University of the Holy Cross on “A cross disciplinary inquiry into human creativity”
(grant PRI-HCR).

Authors' Contributions

Conception and design: FD, AM, IC

Drafting the article: FD, AM, IC

Revising the article: FD, AM, IC

Competing Interests

All the authors declare no competing interests.

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## Figure Captions

Figure 1. Global climate evolution, hominin taxa distribution, and geographic expansion over the last 3.3 million years.

Top panel: Global climate fluctuations inferred from benthic $δ^{18}$O values, showing dominant 41,000-year and 100,000-year glacial-interglacial cycles [78]. Middle panel: Temporal ranges of major hominin taxa, illustrating periods of overlap and succession. Bottom panel: Cumulative expansion of the terrestrial area colonized by hominins (in million km$^2$), showing progressive ecological niche expansion from the East African Rift Valley to the colonization of arctic regions and remote islands.

Figure 2. Chronology of key behaviours shaping the human epistemic niche across three domains.

Top panel: *Structuring of space* (cyan = inside space, blue = outside space) shows the progressive organization of activity areas, including early hearths, bedding, mortuary and symbolic spaces, long-distance transport of symbolic materials, and eventually the construction of artificial structures. Middle panel: *Body culturalization* highlights changes to the body as a cultural medium—symbolic practices like ochre use, bead-making, body modification, and specialized clothing and ornament production.

Bottom panel: *Information storage* traces the emergence of symbolic marking and external memory—from cut marks and abstract engravings to additive marking systems, portable information-encoding artifacts, and formal writing systems, culminating in printing.

Figure 3. Early personal ornaments.

(1) *Glycimeris* shell with traces of red pigment and a natural perforation, Qafzeh, Israel, 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, Israel, c. 115 ka 2, (4-5) Grotte de 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, Morocco, c. 120 ka. Note the dark colouring 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 colouring 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 bead 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; (17-21) personal ornaments made by Châtelperronian Neanderthals at the Grotte du Renne, Arcy-sur-Cure, Yonne, France, c. 45-42 ka; bone eyed needles found at Xiaogushan (22) and Zhoukoudian Upper Cave (23), China, c.39-33 ka. (Photos F. d’Errico apart from 1; modified after [53].)

Figure 4. Archaeological instances of marking practices and symbolic information storage.

(a) Freshwater bivalve from Trinil, Indonesia (c. 540 ka) bearing a deliberately engraved zigzag pattern, the earliest known instances of abstract marking; (b) ochre fragment with a complex geometric motif from Blombos Cave, South Africa (c. 73 ka); (c) photo and drawings of a baboon fibula from Border Cave, South Africa (c. 44 ka) incised with 29 notches using with different cutting edges, interpreted as one of the earliest known artificial memory system. Scales = 1 cm; close-up from Trinil scale = 1 mm. (photos and drawings F. d’Errico).
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