Abstract
Art making, understood as making music, dancing, and producing visual artworks, has a deep evolutionary history. Traces of musical instruments, figurative arts, and rock art, like engravings and cupules, are found all around Europe, Asia, and Africa since the Pleistocene. The debate about the evolution of art making has produced many theories of how arts evolved in humans’ deep past. In this article, I add my contribution to one of the debate’s most influential positions, Ellen Dissanayake’s account of art making, focusing in particular on her explanation of the antecedents of arts, i.e., mother–infant interaction. I first integrate Dissanayake’s account of ape infant behavior with new evidence from comparative studies and develop a new approach to build an account of the hominin suite of behaviors in the context of parental care using evidence from cognitive paleoanthropology and comparative neurobiology. I then propose a novel method to identify, in the fossil record, the evolution of arts as described by Dissanayake. These two lines of research are importantly intertwined: I suggest that changes in hominin parental care may help identify new cognitive skills relevant for tracing the origin and early developments of arts. I conclude that the capacity of art making may have evolved multiple times since the Middle Pliocene (≈4 mya) in different hominin lineages.
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Prelude
Ellen Dissanayake’s evolutionary account of arts and their early foundations in Pleistocene mother–infant interaction is a milestone in studies of the evolution of arts and aesthetics. Few accounts, even when diverging from her views, fail to cite Dissanayake. Her work also attracts transdisciplinary interest, with references in psychology (Chittar et al. 2023; Varella 2023; Bamford et al. 2024; Nadal and Skov 2024), biology (Leongómez et al. 2021; Savage et al. 2021), neuroaesthetics (Chatterjee and Vartanian 2016; Brown 2021), anthropology (Clark et al. 2024; Cross and Morley 2008), archaeology (Bednarik 2003; Davidson 2013; Dapschauskas et al. 2022), and philosophy (Portera 2020; Davies 2021; Cappelletto 2022; Killin 2024).
Important insights can be drawn from Dissanayake’s ideas, which I motivate in this article, presenting her view of art making, the foundational role of Pleistocene mother–infant interaction, and the evidence to trace the evolution of the artistic capacity. Then, building on Pietro Allegretti’s (2026) backdating of mother–infant interaction and on data from paleoarchaeology, developmental paleoanthropology, and comparative studies, I propose a complementary approach to Dissanayake’s research to account for hominin mother–infant interaction and to trace the evolution of arts in the fossil record.
The conclusion is that paleoarts should be reconceptualized. While Dissanayake locates the artification capacity in some Homo species (Pleistocene hominins from ≈2 mya), this capacity should be extended to include Australopithecus (and potentially other Middle-Pliocene hominins from ≈4.2–3.5 mya).
Introduction: Explanation of Artification, Its Evolution, and New Implications
I explain Dissanayake’s account of artification before motivating its reformulation.
Artification is “an evolved behavioural predisposition in members of the genus Homo to intentionally make the ordinary extraordinary (i.e., to “make special”), by means of artistic/aesthetic operations (e.g., formalization, repetition, exaggeration, and elaboration), particularly in circumstances about which one cares (considers important)” (Dissanayake 2014, p. 55). These operations are repetition of elements, like drum pulse; formalization, like movements from ordinary life in pantomime; exaggeration of behaviors or marks, as in choreographed dancing; elaboration through colors and other ornamental additions; and manipulation of the perceiver’s expectations, as a strongly accented sound against a sequence of softer ones in a trumpet solo. These operations can be applied to “face and body (masks and costumes), voice (song), movement (dance), story (poetic language and performance), and surroundings (decorated paraphernalia, shaped and embellished surroundings, and built structures)” (p. 51)Footnote 1. Artification highlights what people do while doing art, and wherever we find this doing, Dissanayake expects to find at least one of those aesthetic operations.
Allegretti (2026) explains how, for Dissanayake (2011), these operations constitute the process through which artists produce perceptual effects described by aestheticians, philosophers of art, and evolutionary psychologists, e.g., the universal, gestalt-like visual art principles like symmetry suggested by Vilayanur Ramachandran and William Hirstein (1999). This process is deployed to produce different emotionally-valenced effects (Brown and Dissanayake 2009): reactions to the outcome of goal-directed actions (e.g., joy/sadness, Brown 2021), to the appeal of objects (liking/disliking, associated to aesthetic emotions), to the actions of agents (approval/disapproval), and to social interactions (comfort/discomfort, associated with affiliation/prosociality).
On Dissanayake’s (2021) view, arts evolved as a subgroup of the artification capacity, expressed as the human universal feature of ritual ceremonies (Allegretti 2026), gatherings socially oriented to pragmatic ends, e.g., channeling individual aggression or facilitating courtship (Rappaport 1999; Legare and Nielsen 2020). These ceremonies would be activities undertaken by the entire community, comprising components, like coordinated dances, music, and visual artworksFootnote 2, that for Dissanayake (2018a) can be distinguished from each other in separated arts only artificially as Western scholars often do.
For Dissanayake (2003), artification in hominins may have produced idiosyncratic cases, e.g., making marks, but these were not adaptively transmitted across generations, unlike ceremonies, which served the functions of group socio-emotional coordination and individual stress relief. A unique awareness, the uncertainty of the future, developed in larger-brained hominins because of new cognitive advancements, e.g., better planning and imaginative abilities. However, it also meant that when survival conditions were unfavorable, individuals would have understood that obtaining desired outcomes was uncertain. Perceived uncertainty produces fear and anxiety (Keltner et al. 1993), releasing physiologically deleterious stress hormones such as cortisol (Lupien et al. 2009), whose harmful effects diminish when individuals feel in control of uncertainty (Sapolsky 1992). Ceremonies may have provided such a feeling: gathering during stressful periods—a typical primate behavior (Churchland 2011)—would be more reassuring than inaction or solitary actions (Taylor 2004). By gathering together, and potentially (but not necessarily) asking extraordinary entities such as spirits or forebears for help, hominins perceived that they were addressing uncertainty (Dissanayake 2017). Also, individuals felt they addressed challenges as a group, facilitating social bonds and more competent collective action, which provides a selective edge.
Dissanayake (2017) proposes evidence for the artification capacity in the fossil record and a method for identifying artification-based ritual ceremonies. Before artifying, hominins required the capacity to perceive extraordinariness, i.e., noticing items that attract attention but lack any clear immediate adaptive function (Dissanayake 2018a). This capacity is identified in the 3.0–2.5 mya perceptually striking Makapansgat pebble, so valued to be possibly carried for kilometers (Dart 1974), or in stored exotic quartz crystals ≈900 kya (Oakley 1997). Early examples of artification are remains of mark-making (Dissanayake 2018a), e.g., ≈900 kya ochre (Beaumont 1990), ≈500–200 kya engravings (Van Peer et al. 2003; Joordens et al. 2015), and ≈400–200 kya adornments (Bednarik 2003). These examples may indicate ceremonies, but this remains uncertain: Allegretti (under review) suggests that for Dissanayake (2003, 2017; Dissanayake in press), artification-based ceremonial practices are identifiable by evidence that they are cross-generationally culturally transmitted. Such evidence is still lacking for the Indonesian engraving-making (Joordens et al. 2015), so this practice would only count as an (idiosyncratically) artified item, but there is evidence for ochre-based ceremonies beginning ≈500 kya (Dapschauskas et al. 2022).
However, Dissanayake (2017) provides a method for identifying the timeframe of archaeologically invisible ceremonies. Using models of cognitive evolution (e.g., Donald 1991; Pellegrini and Bjorklund 2004), it is possible to trace hominin cognitive advancements, such as increased mimetic ability, “conscious, self-initiated, representational acts that are intentional yet not symbolic” (Dissanayake 2017, p. 91), including altered actions and vocal tones, through paleontological evidence of advanced social coordination, e.g., hunting cooperatively, and tool-manufacturing skills. For Dissanayake (2017), some cognitive advancements would be proxies of capacities underlying ceremonies, like human-like mental time travel, that is, “the ability to recall the past in order to imagine the future (Suddendorf and Corballis 1997)” (Dissanayake 2017, p. 90); metarepresentation, that is, “the ability to pretend and to understand pretense in others (Leslie 1987; Cosmides and Tooby 2000) and to appreciate fiction as distinct from reality (Tooby and Cosmides 2001)” (Dissanayake 2017, p. 90); and/or working memory (Kavanagh et al. 2005). She (2017) suggests these abilities predate fully H. sapiens-like symbolic-linguistic capacities (capacities that she dates to ≈50 kya), as the presence of these abilities in the paleontological record can be inferred from evidence of techno-social advancements such as spear-throwing and thousands of hunted animals remains at Schöningen, Germany, from ≈400 kya (Thieme 1997). Dissanayake (2017) also suggests earlier, though more controversial, examples of these abilities, namely, the earliest traces of mimetic abilities at 1.8 mya (Donald 1991), with African technological and hunting evidence reported by Allegretti (2026) further supporting this backdating.
The reason ceremonies could address their adaptive functions lies in hominin ancestral mother–infant early interaction (hereafter, AMIEI; an acronym borrowed from Dissanayake in press). This is an adaptive suite of universal, affective, concurrent, multimodal, and communicative operations to modify signals exchanged by mothers and infants between birth and about six months of age, which include peculiar vocalizations, like soft undulant sounds; facial expressions, head and body movements such as smiling, head nod, etc.; touches and pats; and turn-taking, coregulated, and dyadic interactions (for a review, see Dissanayake 2017). Dissanayake (2014) labels this multimodal suite proto-aesthetic operations: they are implicated in hominin artification, i.e., expressions of repetitive, exaggerated, etc., behaviors. Infants use them unintentionally, while in contrast artification operations are intentional.
This suite exapted great ape signals for friendliness (Dissanayake 2018b), e.g., open-eyes or eyebrow flash (Eibl-Eibesfeldt 2017). AMIEI is considered an ethological ritualization (Dissanayake 2014), the evolutionary process of stereotyping of a behavioral pattern and its cooption from the behavior’s original function for intraspecific communication (Tinbergen 1952; Huxley 1966; Tonna et al. 2020). For example, in some male birds, feeding-related ground pecking was exapted over evolutionary time into a mating display (Huxley 1966). In my view, Dissanayake (in press) suggests that infants have an innate capacity to engage in proto-aesthetic operations (Dissanayake 2014, 2017), and, in her argument, she explicitly connects AMIEI to examples of innate ritualizations, such as Nikolaas Tinbergen’s (1952) case of a gull’s pecking. But also, that AMIEI is partially learned, implying coregulated and contingent interactions based on mutually anticipated dynamic variation and manipulation of signals, especially from 4-month-old infants (Dissanayake 2007, 2011, 2014).
For Dissanayake, AMIEI evolved as an adaptive solution to the obstetrical dilemma (Washburn 1960; Grunstra et al. 2023), the health problems during parturition caused by the anatomical reorganization following bipedalism and the gradual evolution of larger-brained infants (Hublin et al. 2015). Giving birth to premature infants and delaying brain development until after birth, a condition called altriciality, were some evolutionary solutions to the obstetrical problems. Consequently, infants were born increasingly more helpless, requiring greater parental assistance to survive and develop socio-emotional skills, and AMIEI was naturally selected for providing such assistance (Dissanayake 2018b). While Dissanayake (2017) suggests this process started with Homo erectus ≈1.8 mya, here I (Allegretti 2026) assume the Lucy australopithecine mother–infant interaction (LAMII) hypothesis, which emphasizes the role of bipedalism as the origin of this dilemma (Webb et al. 2024) and backdates AMIEI to the Pliocene ≈4.2 mya.
Ceremonies would coopt LAMII’s operations and associated dopaminergic reward systems, e.g., oxytocin, that diminish physiologically deleterious cortisol levels (Lupien et al. 2009) and produce prosocial feelings of bondedness (Panksepp 2004; Brown and Dissanayake 2009). To this day, these systems are recruited for rewarding multimodal mother–infant interactions (Dunbar 2010; Kim et al. 2014; Peltola et al. 2018; Scatliffe et al. 2019; Hiraoka et al. 2020; Filippa et al. 2021; Dissanayake 2021; Hirschel et al. 2023) and group-based vocalizations and dancing (Phillips-Silver and Keller 2012; Savage et al. 2021).
The exaptation of AMIEI for ceremonies may have occurred multiple times and for different functions, e.g., sexual selection, social bonding, coalition signaling, parental attention, and aposematic display (Dissanayake 2014; Dissanayake in press). Moreover, Allegretti (2026) suggests that ceremonies at different evolutionary times would rely on different proximate mechanisms, coopting various cognitive, anatomical, material, and social components in these displays, e.g., refined vocal control or lithic sound production (Killin 2017). Indeed, Allegretti proposes ceremonies are more accurately characterized as having an evolutionary mosaic structure (Parravicini and Pievani 2019), a set of interrelated traits emerging following different evolutionary trajectories at different times for different selective pressures that coevolve together.
In summary, I explained Dissanayake’s artification and Allegretti’s reformulation, focusing on the timeline of AMIEI (reformulated as Allegretti’s LAMII), artification capacity, and ceremonies. The rest of the article follows a twofold aim: using evidence from comparative studies and cognitive paleoanthropology (Currie et al. 2024)Footnote 3, it provides support to the LAMII hypothesis with a new account of its proximate mechanisms and suggests a method to help identify the possible timeline of artification capacity, similar to that described above for ceremonies. This account and method will be used to suggest the conjectural hypothesis that artification evolved on small cognitive upgrades compared to nonhominin apes (hereafter, apes) and is shared with non-Homo species.
A Phylogenetic Perspective on LAMII: Proximate Mechanisms Account
I develop an account of australopithecine mother–infant interaction based on comparative studies of ape ritualization mechanisms. Using comparative evidence is justified, based on the relatively uncontroversial notion that australopithecines still led similar lifestyles to apes and were just developing new behaviors, e.g., longer foraging expeditions and inclusion of new food in their diet (Sterelny 2021). However, even the apes most closely related to humans, bonobos and chimpanzees, are inadequate as examples for australopithecines, since each species is divided by millions of years of independent evolution (Almécija et al. 2021).
To avoid overgeneralizing from a single species, it is important to first map a suite of behaviors shared across apes. From a phylogenetic perspective, traits shared among many closely related species are more likely to have evolved once in the common ancestor or multiple times through similar developmental patterns, so this suite would be more plausibly common to all Hominidae species. This perspective protects the LAMII hypothesis from the reverse engineering criticism (Gould and Lewontin 1979; Parravicini and Pievani 2018), that is, assuming the phenotype of a trait in the past based on its current characteristics without testing or justifying this assumption. After developing the ape suite, I suggest that proto-aesthetic operations are this suite’s specific upgrades in LAMII, i.e., increased intentional communication. This account of LAMII is a contribution to Dissanayake’s mother–infant interaction hypothesis.
Ritualization in Hominidae Mother–Infant Interaction
This section maps the Hominidae behavioral suite.
Human newborns interact intimately with their caregivers (Beebe and Lachmann 2013; Hrdy 2024), a characteristic shared with apes and other primates (Whitham et al. 2007; Luef and Liebal 2012; Rossano 2013; Knox et al. 2019; Fröhlich et al. 2022; Amici et al. 2023). Also, human and ape infants adapt their communication strategies in mother–infant interaction, e.g., repeating and varying signals in all sensory modalities (Falk 2004a, b; Hobaiter and Byrne 2011; Luef and Liebal 2012; Halina et al. 2013; Dissanayake 2015; Tomasello and Call 2019; Fröhlich et al. 2022). So, hominins presumably shared these characteristics too.
I assume Hominidae mother–infant interaction is at least partially based on ritualization. As an example of ritualization, Marta Halina et al. (2013) suggest that when either a bonobo infant or mother signals to the other the intention to depart together, e.g., the infant grabbing the mother’s fur to pull her to himself, or the mother pushes the infant, and so on. After repeated interactions, the partners understand each other’s intention from the initial steps of the signaling sequence, using these steps as gestures, e.g., grabbing without pulling (Halina et al. 2013), a process described as learned intention movements (Tinbergen 1952). Ape infants also develop attention-getter gestures, learned through the understanding that specific idiosyncratic movements attract attention in multiple contexts, e.g., a ground slap to initiate play and nursing (Tomasello and Call 2019).
Whether apes ritualize remains debated (Hobaiter and Byrne 2011; Fröhlich et al. 2025). Interestingly, Dissanayake (2017) uses Barbara King’s (2004) account of ape communication, which does not assign a role to ritualization. However, while Halina et al. (2013) agree with King (2004) that gestures develop through continuous interindividual adjustments to each other’s behavior, they disagree that signals’ form and function change continuously:
[u]nder ontogenetic[learned] ritualization, behaviors are ritualized into gestures with relatively stable forms and communicative functions. Indeed, it is the stability of a gesture that allows a signaler to use it to satisfy a particular goal in a wide variety of contexts. If the immediate context affected the communicative message of a gesture in major ways, then an individual would not be able to use that gesture to attain her goal unless the context was right. This problem is especially acute for those situations in which an individual is facing a context that is not conducive to her goal—for example, in the case of an infant trying to get her mother to change her behavior from resting supine to walking (Halina et al. 2013, p. 664).
Also, Halina et al. (2013) object to Catherine Hobaiter and Richard Byrne (2011) on the correctness of their arguments against ape ritualization, e.g., that the reach gesture is coopted from the taking action, an implausible ritualization since “chimpanzees do not respond to the initial sequence of behaviors involved in taking by handing an object over to the taker. In general, it is difficult to imagine that successful taking will depend on the actions of another individual in chimpanzees” (Halina et al. 2013, p. 663). I refer to Michael Tomasello and Josep Call (2019) for a detailed analysis of the debate. Here, I assume that Hominidae ritualize.
A potential criticism of using Halina et al.’s (2013) examples of 10–24-month-old bonobo infants to reconstruct LAMII’s proximate mechanisms is that Dissanayake’s AMIEI hypothesis focuses on newborns or infants under 6 months. To the best of my knowledge, no studies examined younger apes. Nevertheless, pending new data, it is plausible that they learn ritualizations, given evidence of ape and human cognitive capacities and developmental pathways relevant to emotion and communication. Ape and human infants engage in face-to-face interactions, vocalizations, and body signals with their caregivers a few hours after birth (Rijt-Plooij and Plooij 1987; King 2004; Myowa-Yamakoshi et al. 2005; Falk 2009; Amici et al. 2023), and week-old apes and humans have similar cognitive capacities for sustained attention to signals or cues, orientation, arousal, state regulation, and motor-attentional adjustment during caregiver-infant interaction (Amici et al. 2023; Bard et al. 1992, 2005, 2011; Knox et al. 2019; Meltzoff and Moore 1977; Myowa-Yamakoshi et al. 2004, 2005; Redshaw 1989), as well as for learning gestures, e.g., reaching, pointing, etc. (Gillespie-Lynch et al. 2013). The main difference is that humans have less advanced locomotory performance (Bard et al. 2011). So, early mother–infant interactions may play a comparable role in emotional and communicative development in apes and humans from birth on (Nadler 1974; Bard 2005; Bard et al. 2005, 2011; Fröhlich et al. 2025), supporting the hypothesis that if 4–6-month-old humans learn to ritualize (see “Introduction”), younger than 10-month-old apes and hominins may possess similar capacities. Also, since apes and humans are born altricial (Webb et al. 2024), and, as suggested, exhibit neonatal similarities in ritualization-related cognitive, emotional, and communicative capacities, these similarities suggest that if H. sapiens developed a neonatal predisposition for ritualization, it plausibly evolved in apes and other hominins too (cf. Dissanayake 2014).
Ape and human infants share partial control over intentional communication (Kret et al. 2020), a feature not emphasized in the AMIEI hypothesis (cf. Dissanayake 2011). Andrea Knox et al. (2019) suggest this control can be inferred in apes from proxies including:
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the orientation of the signaler’s body and gaze toward the recipient (Tomasello and Call 2007);
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the wait for a response and gestural repetition (Knox et al. 2019);
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the persistence toward a goal, e.g., adding new gestures (Tomasello and Call 2019).
The previous example of the bonobo infants exhibits these proxies: they orient themselves towards the recipient, wait, and persist for a response. Since ape infants ritualize (Tomasello and Call 2019) and, as suggested above, this process may begin in early infancy, this intentional communicative control may gradually develop from infancy onward. Additionally, this conclusion implies that the interindividual coordination in ape mother–infant interaction, developing ontogenetically (King 2004; Rossano 2013; Halina et al. 2013; Fröhlich et al. 2016; Knox et al. 2019; Tomasello and Call 2019), may be partially intentional.
In conclusion, partial intentional control over ritualization and interpersonal coordination in mother–infant interaction is a phylogenetically common ability in Hominidae. I now suggest a possible australopithecine novelty, i.e., increased intentional control.
Before proceeding, it should be noted that these cognitive components are not shared uniformly across apes and are restricted compared to humans. Apes show partial intentional control over gestures (Genty et al. 2009; Hobaiter and Byrne 2011; Gruber 2014; Tomasello and Call 2019; Liebal and Oña 2020; Kret et al. 2020; Lameira et al. 2025), not as proficiently as humans, e.g., in the expression of facial emotions (Kret et al. 2020). Whether apes control vocalizations for socio-emotional coordination is controversial (Hobaiter et al. 2023; Rodrigues and Fröhlich 2024; Lahiff et al. 2025; Fournier et al. 2025; Lameira et al. 2025), e.g., for alarm calls (Liebal and Oña 2020). Orangutan and bonobo infants are more responsive to social signals (Fröhlich et al. 2016); in orangutans, this is probably due to prolonged parental care, which rehearses the ability to understand their social partners’ communicative attempts (Knox et al. 2019), considerations useful for the next section.
LAMII, a Fosterer of Intentional Ritualization
Australopithecine parental changes were marked by increased intentional control. In support of this claim, I summarize Anton Killin’s (2017) account of Pliocene hominin cognitive advancements, to then suggest LAMII’s role.
Killin (2017) accounts for the stepwise, multispecies gradual coevolution of human system 2 cognitionFootnote 4, including “mental time travel, hypothetical reasoning, complex theory of mind, complex forward planning” (p. 226)Footnote 5. This system is linked to subjective experiences of agency and concentration (Kahneman 2011), and examples include maintaining a different-from-natural walking pace (Kahneman 2011), with Killin (2017) citing the 3.66 mya Laetoli trackways, comprising nested footprints (Crompton et al. 2012; Shaw-Williams 2014), as early evidence. System 2 upgrades are also connected to an advanced theory of mind compared to apes, possibly required for social learning of the 3.3-million-year-old Lomekwian industry’s manufacturing process (Harmand et al. 2015; Lombard et al. 2019), and for group scavenging, suggested––albeit controversially––by the 3.4 mya cut-marked bones in Ethiopia (McPherron et al. 2010; Domínguez-Rodrigo et al. 2012). I propose that system 2 evolution may have started with LAMII ≈4.2 mya, first presenting the Pliocene hominin ecological scenario, then integrating new paleontological and comparative evidence with Tomasello’s (2023) model of ape metacognition.
North African environments ≈4.2 mya were predominantly woodlands, just gradually starting to transition to more open landscapes with increasingly dispersed vegetation (Wrangham 2009; Su and Haile-Selassie 2022; Doman and Early 2022; Drummond-Clarke 2023; Orr et al. 2023; Trájer 2023; Drummond-Clarke et al. 2024; Amarathunga et al. 2024; Faith et al. 2024; Tierney et al. 2025). Bipedal hominins were likely mobile, wide-ranging foragers (Sterelny 2021), with a mostly plant-based omnivorous diet (Wynn et al. 2020; Lüdecke et al. 2025), living in habitats plausibly showing decreasing density of predators (Faith et al. 2024), and reduced herbivore competition, e.g., foraging spaces left vacant by continuous habitat shifts (Quinn et al. 2021; Trájer 2023). Concomitantly, slightly increasing dispersed vegetation would have favored ape-like anti-predator strategies, based on early detection of pack-hunting predators for arboreal fleeing (Tutin et al. 1981; Hiraiwa-Hasegawa et al. 1986; Pruetz et al. 2008; Nishida 2011; King 2024), favored by upper-body adaptations for climbing (Alemseged 2023), or mobbing solitary predators (Hiraiwa-Hasegawa et al. 1986; Boesch 2009; Nakamura et al. 2019; King 2024). These ecological conditions plausibly favored increasing Panini-like (chimpanzees and bonobos) scavenging behavior (Thompson et al. 2019), e.g., locating carcasses before advanced decomposition (Watts 2008).
Compared to Killin’s (2017) later Pliocene (≈4.0–2.6 mya) scenario, characterized by expanding open and mosaic landscapes at the expense of woodlands, increasing risks of predation and herbivore competition, my scenario describes more ape-like and less ecologically dramatic changes at this timeline’s upper bound. In Killin’s scenario, wide-ranging foragers, such as Australopithecus afarensis, in more open habitats, manufacturing stone tools (Harmand et al. 2015) and—controversially—engaging in group scavenging (McPherron et al. 2010), would have increased opportunities for rehearsing social vocal communication, potentially bringing it under partial voluntary control (Killin 2017). Here, I emphasize the process itself, the increased opportunities as fosterers of intentional control, suggesting that a similar process may have begun already ≈4.2 mya for LAMII multimodal control of ritualization. Albeit archaeological evidence from this time is lacking, the conditions for this process may have likely been in place: there were already hominins potentially able to produce Lomekwian technology (Plummer et al. 2025), e.g. afarensis living ≈4.2–2.9 mya (Alemseged 2023); as suggested later, this process of intentional control is present in neurobehaviorally similar apes; and in this ecological scenario new hominin neuroanatomical constraints may have advanced this process.Footnote 6
This process is similar to the one suggested in the previous section for the development of intentional behaviors in ape mother–infant interactions (Knox et al. 2019), where continuous engagement fosters repetition and innovation of ritualized signals under increasing intentional control. In my view, both Killin’s and my proposed processes align with Tomasello’s (2023) account of ape metacognition, suggesting apes and early hominins develop greater intentional control and focus through the repeated monitoring of their own and others’ intentions and actions. Foraging contexts can foster this control (Tomasello 2023), supporting Killin’s (2017) hypothesis, since Pliocene hominins were plausibly still cognitively ape-like (Sterelny 2021); but I suggest mother–infant interaction can foster this control too. In apes, such interactions depend on Tomasello’s monitoring (Knox et al. 2019), with orangutans showing particularly advanced responsiveness to signals, due to prolonged maternal dependance (Knox et al. 2019), suggestive of repeated interactions as fosterer of monitoring-related capacities, e.g., understanding intentionality, behavioral inhibition, motor control (Tomasello 2023), and coordinated dyadic interactions (Knox et al. 2019), a fundamental component of Dissanayake’s account (see the “Introduction”). In australopithecines, altriciality may have required continuous interactions (Allegretti 2026), fostering ape-like ontogenetic learning for intentional control of affiliative/prosocial communication, such as contact calls or ritualized attention-getting gestures exchanged in dyadic coregulated interactions.
This process may have been fostered by ape-like declines in caregivers’ responsiveness to signals produced by infants once they reach ≈4 months of age, e.g., ignoring food requests (King 2004; Amici and Liebal 2025). Such declines may have favored continued communicative efforts in more altricial australopithecine infants (plausibly crucial for survival; see Allegretti 2026), fostering monitoring intentions and intentional control. In contrast, ape infants abandon communicative attempts more readily due to greater locomotor independence, thereby reducing their reliance on caregivers (King 2004; Halina et al. 2013). Also, australopithecines may have developed neuroanatomical constraints favoring advanced intentional control. Paleoanthropological models of australopithecine skulls and obstetrical constraints suggest that the (controversial) early encephalization process from ape-like endocranial volume of ≈400 cc to >400–500 cc, that was associated with larger neocortices (Gamble et al. 2014), supporting advancements in intentionality (Powell et al. 2010), motor control (Baker et al. 2025), and prosociality (Gamble et al. 2014; Piantadosi and Kidd 2016), potentially begun ≈4.2 mya (for a review, see Allegretti 2026). Furthermore, models of australopithecine hand morphology indicate relatively long thumbs (Alba et al. 2003; Rolian and Gordon 2013; Almécija and Alba 2014), associated with larger neocortices in primates (Baker et al. 2025). Finally, larger-brained and altricial australopithecines may have developed advanced social skills (Gamble et al. 2014; Allegretti 2026) that are linked to improved intentional control and monitoring of others’ behaviors and intentions (Burkart et al. 2009). Apes already show individual variability in parental motivation (Abello and Colell 2006), which may have been selected for in LAMII (Allegretti 2026). Increased parental care fosters the ontogenetic development of prosocial skills in various contexts, e.g., courtship and play (Crockford et al. 2020; Francesconi et al. 2024; Sabbi et al. 2024), fostering intentionality-related abilities.
In summary, australopithecines would have undergone similar processes to the orangutan dependence, extending ape-typical metacontrol (Tomasello 2023), which, coevolving with new constraints for increased prosociality (Allegretti 2026), may have fostered small advancements in intentional control for communication than apes, i.e., a greater number of learned, intentional, and ritualized signals exchanged flexibly in coregulated interactions.
A potential criticism of this hypothesis concerns the uncertainty surrounding parental care strategies in australopithecines. There is no clear paleontological evidence of biological reproduction and social systems (Halcrow et al. 2020), and paleoanatomical reconstructions are consistent with a range of primate care systems, spanning from gorilla-like harems to more advanced alloparenting than the type present in Panini (Manthi et al. 2012; Gamble et al. 2014; Suwa et al. 2021). An almost exclusively maternal care system, the most common in primates and apes (Burkart et al. 2009; Ziegler et al. 2022), may provide a phylogenetically plausible explanation for australopithecines, given also the paleoecological context outlined above, which suggests ape-like lifestyles (Sterelny 2021). However, the obstetrical dilemma and its backdating to the Pliocene is linked to a more human-like transition to advanced alloparental, cooperative care, e.g., prolonged time-consuming support from helpers (Hrdy 2006, 2024; Burkart et al. 2009; Hawkes and Coxworth 2013), suggesting a possible earlier origin of this type of care. This advanced alloparenting would support my hypothesis of advancements in intentionality (Burkart et al. 2009), albeit not through continuous mother–infant interactions, but multiple caregivers–infant interactions.
However, this alternative seems unlikely. The australopithecine obstetrical dilemma would not be connected to the advanced socio-cognitive upgrades linked in the traditional account to advanced alloparenting, e.g., increased food intake (Zollikofer and Ponce de León 2010; Thompson and Nelson 2016), important advancements in prosociality (Gamble et al. 2014), new developmental patterns (Bogin 2020; Meneganzin and Currie 2025), and cognitive, foraging, technological, and social innovations, e.g., advanced cooperative hunting (Hrdy 2006; Hiscock 2014; Cofran and DeSilva 2015; Killin 2017; Halcrow et al. 2020; Sterelny 2021). These upgrades are usually attributed to later larger-brained (>900 cc) and probably more altricial hominins like H. erectus (Antón and Snodgrass 2012; Mitteroecker and Fischer 2024), from a period where there is paleontological evidence that hominins were markedly more prosocial than apes (Sterelny 2021), e.g., cooperatively hunting adult bovids and hunting and scavenging hippopotamids (Bunn and Pickering 2010; Plummer et al. 2023), implying that advanced alloparenting coevolved with already increased individual prosocial motivations compared to apes.
The evolution of these motivations may indeed have started with australopithecine obstetrical constraints fostering LAMII, but it is not plausible that this led to advanced alloparenting in early hominins. Altriciality may have indeed increased the energetic costs for infant transportation (DeSilva 2011), and time-consuming communication (DeSilva 2011; Dunbar 2012, 2022; Halcrow et al. 2020), e.g., more frequent separations between infant and mothers foraging on the ground or in trees (Falk 2009). However, the ≈4.2 mya Pliocene conditions described above suggest that mothers could have addressed these challenges. Hominins with smaller, less energetically costly brains than H. erectus may have found favorable ape-like foraging conditions with less herbivore competition and predation risk, allowing longer time-consuming interactions for mothers and infants foraging together.
Apart from LAMII, evidence of other ≈4.2 mya Pliocene selective pressures favoring prosocial motivations potentially supporting advanced alloparenting is less clear. Pliocene favorable foraging conditions alone would unlikely have selected for the maintenance of novel survival strategies requiring prosocial upgrades compared to apes. Later hominins’ strategies, e.g., systematically hunting difficult prey or group mobbing to deter predators and/or competitors (Sterelny 2021), entailed considerable individual safety risks and were unlikely to be adopted when more accessible foraging options were more readily available. Moreover, archaeological evidence for such risky behaviors remains absent. Also, eventual novel foraging methods for ape-like resources, e.g., plants or small vertebrates, in Pliocene australopithecines, cognitively comparable to apes, would not plausibly explain prosocial advancements alone. Apes are strongly conflictual interindividually while foraging, e.g., caching deceptively or withholding food (Whiten and Byrne 1988; Burkart et al. 2009; Tomasello 2023), do not form more stable, numerous groups even in favorable foraging conditions (Gamble et al. 2014), and in primates better foraging skills alone are not associated with larger neocortices fostering prosocial advancements (Dunbar 1995). Finally, it seems that evidence for prosocial advancements compared to apes prior to LAMII in the earlier Miocene 8–5 mya is missing. Paleoecological research of this period is suggestive of ape-like foraging conditions like predominant woodlands (e.g., Orr et al. 2023), and paleontological research provides much less clear evidence of obligated bipedalism and its associated obstetrical and foraging constraints (DeSilva et al. 2022; Webb et al. 2024; Prang et al. 2025; Allegretti 2026), of larger neocortices (Gamble et al. 2014; Gómez-Robles et al. 2024), and of increased toolmaking-related manual dexterity (Marzke 2013; Kivell 2015; Prang et al. 2021), possibly associated with Pliocene advanced social learning (Lombard et al. 2019).
So, I conjecturally hypothesize that prosocial upgrades in australopithecines developed in maternal care as effects of obstetric constraints during favorable ape-like foraging conditions, later coevolving with group-increased cooperative behaviors, e.g., advanced lithic technologies (Plummer et al. 2025), or alloparenting. While australopithecine mothers may have accepted assistance more than is usual for apes (Falk 2009, 2025), the socioecological constraints and parental costs discussed above undermine the hypothesis that this reliance already greatly exceeded, as in humans (Burkart et al. 2009), the limited, controlled interactions that ape mothers typically permit with infants to helpers (Burkart et al. 2009).
In conclusion, LAMII could have exacerbated the ape-like process of developing a suite of intentional ritualizations (Halina et al. 2013; Knox et al. 2019; Tomasello and Call 2019), fostering a hominin’s greater multimodally learned repertoire of prosocial signals under increasingly flexible voluntary control. While the nature of australopithecine parental strategies remains uncertain (Halcrow et al. 2020), I proposed here that almost exclusive maternal care is a phylogenetically highly plausible option. This has implications for Dissanayake’s AMIEI hypothesis: like her, I suggested that hominins ritualize and engage in dyadic contingent interactions, but contrary to her account, these components are also found in ape infants, together with partially intentional control over ritualization, so that the difference with hominin infants’ proto-aesthetic operations is in the latter’s upgraded degree of voluntary control.
This control may indicate the origin of artification. To justify this claim, I will first suggest that the cognitive proxies of ceremonies, e.g., metarepresentation, provide the conditions for tracing the evolution of the artification capacity as well.
The Evolution of Arts: Paleontological Identification of Artification
The evolution of artification capacity can be traced more precisely than Dissanayake suggests. While she offers examples, like ceremonies and ornaments (Dissanayake 2014), she specifies the timeline and cognitive conditions underlying artification less clearly than for ceremonies, e.g., 1.8–0.4 million-year-old metarepresentation (Dissanayake 2017). Such a clarification would contribute to identifying controversial archaeological cases. This section suggests that the same ceremonies-related socio-cognitive components, like metarepresentation, can help to trace artification, and concludes that this capacity may have evolved in the Middle Pliocene already.
Allegretti (2026) emphasizes that apes already show, to a lesser degree, the cognitive capacities underlying ceremonies (Laland and Seed 2021), i.e., metarepresentation, mental time travel, and working memory, and thereby further expands Dissanayake’s (2017) gradualistic view of artification. Apes already have some intentional control over ritualization (Tomasello and Call 2019), required for artification; limited prelinguistic symbolic capacities (Matsuzawa 2004; Vauclair 2022); and in artificial settings produce human painting-like marks (Morris 1962; Lenain 1997; De Waal 2001; Davies 2012). The gradual advancements in ceremonies-related cognitive components are associated with various artified items in the archaeological record. As suggested in the “Introduction”, Dissanayake identifies traces of artification scattered throughout the Pliocene–Pleistocene, e.g., the Makapansgat cobble (≈3.0–2.5 mya), ≈900 kya ochre, and ≈200 kya ornaments, all different periods associated with different cognitive advancements. For example, ≈2.9–2.5 mya hominins were already producing the Oldowan technology (Plummer et al. 2023), whose new production methods and uses in the tool industry (e.g., cores to produce flakes and/or as pounding tools) may reflect novel specializations in group coordination (Killin 2017). Improved coordination is associated with advancements in forward planning, theory of mind, and pretense abilities (Leslie 1987; Gowlett et al. 2012), which are relevant to human metarepresentation, mental time travel, and working memory (Wynn and Coolidge 2011; Allman and Mareschal 2016; Currie and Killin 2019). At 1 mya, hominins colonized greater parts of Eurasia (Sánchez-Bandera et al. 2023; Guo et al. 2023), which is associated with socio-cooperative upgrades (Sterelny 2021), and ≈400 kya hominins produced unique inventions in the animal kingdom, e.g., hafted tools (Kuhn 2020), suggestive of advanced forward planning (Killin 2017).Footnote 7
In my view, these considerations suggest that small advancements in the cognitive components for a specific artified invention—ritual ceremonies—may underlie and be connected to the production of all the other artified items and, by extension, the entire artification capacity. Also, given the emphasized gradualistic view of these capacities relative to apes suggested above (Allegretti 2026), and since these cognitive abilities underlie various artified items such as ornaments, the cognitive difference required for artification capacity may be incremental upgrades from an ape-like baseline. Evidence for these upgrades could therefore be used as proxies to identify the timeframe of artification in the fossil record even in the absence of archaeological evidence, an approach Dissanayake (2017) developed for ceremonies (Donald 1991), as I explained in the “Introduction”. However, I have admittedly not developed a framework to define the specific cognitive advancements underlying the origin of artification. I only suggest artification’s possible timeframe, providing conjectural hypotheses of artification’s archaeological evidence and origin.
Regarding the evidence, some Acheulean handaxes, as ancient as ≈2.0–1.7 mya (Diez-Martín et al. 2015; Mussi et al. 2023), may be artified items (Allegretti under review). Acheulean toolmakers may have intentionally produced items with symmetric patterns (Currie 2011; Wynn and Berlant 2019; Meneganzin and Killin 2024) that, as suggested in the “Introduction”, are the perceptual counterparts of aesthetic operations (Dissanayake 2011), in order to produce an aesthetic effect (Meneganzin and Killin 2024), one of artifiers’ intended reactions (Brown and Dissanayake 2009). Also, some Oldowan tools may be artified: for Thomas Wynn (2021), these tools already show perceptual ergonomic features of the Acheulean industry (Gowlett 2007), features likely manipulated by Acheulean toolmakers to achieve an aesthetic effect. Plausibly, some Oldowan ergonomic features were produced intentionally (although controversial, see Wynn 2021), as this technology may have required advanced cognition (for a review, see Killin and Pain 2023). More conjecturally, since 3.3 million-year-old Lomekwian toolmakers may show some Oldowan-related manufacturing abilities, e.g., rotating the cores during flake removal (Killin 2017), perhaps also these hominins share similar cognitive abilities, indicating the use of aesthetic-like operations and artification in Lomekwian industry too, now counting >250 artifacts (Plummer et al. 2025). Future discoveries may backdate such examples to ≈ mya, potentially revealing tool-related advanced cognitive capacities compared to apes, as suggested in the LAMII section.
Based on this review of evidence of artification and the consideration that this capacity includes many phenomena like ritual ceremonies, ornaments (Dissanayake 2014), Acheulean items (Allegretti under review), etc., all evolving at different times, I suggest a mosaic structure may characterize artification capacity too (Parravicini and Pievani 2019). These practices may have evolved for different functions, at different times, and coopted distinct proximate mechanisms, constituting together artification’s own building blocks (borrowing the concept from Savage et al. 2021). Artification may have exhibited internal variability, not only across populations, with differing cognitive capacities and material cultures, but also within individuals, who could flexibly use the aesthetic operations, for example, producing soothing or aggressive vocalizations in different ceremonies. For these reasons, I suggest that Tecumseh Fitch’s (2015) concept of cognitive toolkit, a trait that is used for different functions and shows variable forms (Savage et al. 2021), seems suitable to describe the artification capacity. Once evolved, the artification toolkit would follow different coevolutionary processes in different hominin populations, as suggested by the mosaic structure of ceremonies (Allegretti 2026).
Regarding artification’s origin, changes in hominin prosociality suggest cognitive advancements relevant for artification, such as theory of mind and pretense (Gowlett et al. 2012), and Allegretti (2026) suggests LAMII was the first prosocial upgrade in hominin lineages as described by the social brain framework (Gamble et al. 2014). The previous section suggests that earlier cognitive upgrades are unlikely, so perhaps LAMII may be the earliest minimal cognitive upgrade for artification capacity to evolve; Australopithecus mothers played with their infants by intentionally enacting the aesthetic operations, and the Pliocene conditions described in the last section may account for the emergence of the first hominin artified scenario. Apart from LAMII, resolving dubious cases of artification such as Oldowan and Lomekwian technologies could help establish the minimal cognitive conditions for artification and identify artification’s likely multiple origins and developments, e.g., with upgrades as in Laetoli footprints or Lomekwian tools, associated respectively with improved pretense (Shaw-Williams 2014), and theory of mind (Lombard et al. 2019).
Artification’s origin may be conjecturally traced to apes (cf. Dissanayake 2014), since the cognitive advancements suggested for LAMII are minimal, and I suggested that during the Pliocene different ecological pressures (e.g., decreasing predation risks) may have favored the expression of ape-like latent capacities, such as prosocial attitudes. Apes reared in semi-wild or artificial settings show increased communicative creativity (Van Schaik et al. 2016; Matsuzawa 2020; Gibson et al. 2023, 2024), exploiting proficiently human-made tools’ perceptual features (Gibson et al. 2023, 2024), suggestive perhaps of an Hominidae capacity to notice extraordinariness. Also, apes wear adornments (Van Leeuwen et al. 2014) and perform components of ceremonies (Kühl et al. 2016; Tennie and Van Schaik 2020). So, using the conceptual toolkits of comparative research (Currie 2021; Halina 2025), attention-getter and intention-movement signals (Tomasello and Call 2019), aimed at producing artification-related reactions, like, for example, prosocial emotions (Brown and Dissanayake 2009), may be suitable behaviors to investigate ape artification and potentially provide insights into artified behaviors of early hominins, using the methodological toolkits of cognitive paleoanthropology (Currie et al. 2024). Also, convergent evolution of Pliocene hominin-like advances in forward planning, episodic memory, pretense, and intentional ritualization may be minimal conditions of artification to evolve in different phylogenetically distant species.
In conclusion, it is possible to use the same type of cognitive advancements typical of hominin ceremonies to investigate the mosaic evolution of the artification toolkit. I provided a conjectural account of one of artification’s origins ≈4.2 mya, the Lucy australopithecine mother–infant interaction hypothesis (LAMII), suggesting that this capacity was later selected in hominins at different times and under different selective pressures to foster different emotional reactions (Brown and Dissanayake 2009).
Conclusion
I provided a new phylogenetic argument of the behavioral suite of Pliocene hominins, suggesting their mother–infant communication may have been voluntarily controlled to a greater degree than in apes and that this control may constitute a proxy of the origin of artification. I justified this claim after providing a novel method to trace artification’s timeline, i.e., using the same cognitive components implied in ceremonies as proxies, although more methods to identify artification in the fossil record are warranted.
Anchoring artification components to the paleoarchaeological record is important for testing Dissanayake’s timeline using empirical and surrogative evidence, e.g., models (Currie 2018), and for providing plausible scenarios of the mosaic evolution of artification in future research, using new approaches for reconstructing complex art-related traits (Killin 2017, 2018, 2024). What the minimal socio-cognitive advancements underlying artification are is still conjectural, but presumably this capacity evolved following a multiple and multispecies origin, suggestive of the necessity of adopting Adrian Currie’s (2015) approach of identifying relevant evidence and models to account for highly specific paleontological cases, so as to reconstruct the minimal cognitive requirements of artification in the past.
Data Availability
Not applicable.
Notes
For a detailed explanation, see Allegretti (2026).
Cognitive paleoanthropology deals with “the evolution of hominin cognition—that is, the cognitive variety and faculties of the complex lineage leading from our last common ancestor with Pan—as it is explored through the archaeological and paleontological record” (Currie et al. 2024, p. 4).
This system is described by the psychologist Daniel Kahneman (2011), that distinguishes between two systems in the human brain, that is, system 1 and 2. System 1 “operates automatically and quickly, with little or no effort and no sense of voluntary control” (Kahneman 2011, p. 20). Examples of the automatic activities of system 1 include “Detect that one object is more distant than another. Orient to the source of a sudden sound. … Read words on large billboards” (Kahneman 2011, p. 21). Instead, system 2 “allocates attention to the effortful mental activities that demand it, including complex computations” (Kahneman 2011, p. 21). Examples of this system include “Focus on the voice of a particular person in a crowded and noisy room. Look for a woman with white hair. Search memory to identify a surprising sound” (Kahneman 2011, p. 22). Intentional behavior in humans today is a result of system 2 cognition. For a detailed explanation, see Kahneman (2011).
Theory of mind is the ability “to explain and predict the behaviours of another person by attributing to them states of mind or intentions” (Powell et al. 2010, p. 3554).
Evolutionary constraints are the physical-structural limits of traits derived from an individual’s phylogenetic inheritance (Melis et al. 2024).
See Killin (2017) for a review of Pleistocene relevant cognitive upgrades.
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Acknowledgments
I warmly thank Justine Kingsbury and Anton Killin for fundamental comments on this article. I also incredibly benefited from conversations with Ellen Dissanayake, Telmo Pievani, Barbara Fischer, Nicole Grunstra, and Andra Meneganzin on the topics of this research, whom I also thank. Many thanks to Te Whare Wānanga o Waikato—the University of Waikato’s Doctoral Scholarship, from which the author received funding while conducting the research that inspired this article, and to two anonymous reviewers.
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Allegretti, P. Ancestral Mother–Infant Interaction and Its Relevance for the Evolution of Arts. Biol Theory (2026). https://doi.org/10.1007/s13752-026-00552-9
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DOI: https://doi.org/10.1007/s13752-026-00552-9