Neanderthals

O V E R V I E W

Triangulating Neanderthal cognition: A tale of not seeing the forest for the trees

Michael Breyl

Germanistik, Komparatistik, Nordistik, Deutsch als Fremdsprache, Ludwig- Maximilians-University of Munich (LMU), Munich, Germany

Correspondence Michael Breyl, Germanistik, Komparatistik, Nordistik, Deutsch als Fremdsprache, Ludwig-Maximilians- Universität München, München, Germany. Email: [email protected] muenchen.de

Abstract

The inference of Neanderthal cognition, including their cultural and linguistic

capabilities, has persisted as a fiercely debated research topic for decades. This

lack of consensus is substantially based on inherent uncertainties in

reconstructing prehistory out of indirect evidence as well as other methodolog-

ical limitations. Further factors include systemic difficulties within interdisci-

plinary discourse, data artifacts, historic research biases, and the sheer scope

of the relevant research. Given the degrees of freedom in interpretation ensu-

ing from these complications, any attempt to find approximate answers to the

yet unsettled pertinent discourse may not rest on single studies, but instead a

careful and comprehensive interdisciplinary synthesis of findings. Triangulat-

ing Neanderthals' cognition by considering the plethora of data, diverse per-

spectives and aforementioned complexities present within the literature

constitutes the currently most reliable pathway to tentative conclusions. While

some uncertainties remain, such an approach paints the picture of an exten-

sive shared humanity between anatomically modern humans and

Neanderthals.

This article is categorized under:

Cognitive Biology > Evolutionary Roots of Cognition

Linguistics > Evolution of Language

K E Y W O R D S

cognition, evolution, language, Neandertal, Neanderthal

1 | INTRODUCTION

Considering the diversity of attitudes on Neanderthal cognition that can be found within the literature, it may be best to start off with an extensively well-founded consensus opinion: From indigenous lifestyles to stock market brokers and from today's population into the deep past of our species, there is no doubt that general biology, cognitive ability, and language use have been a constant throughout all modern human groups for at the very least many tens of thousands of years. As a result, ecological, demographic, epidemiological, cultural, and technological factors are sufficient to explore transitional events between modern human populations. To exemplify, based on long-standing research it

Received: 28 May 2020 Revised: 9 August 2020 Accepted: 13 August 2020

DOI: 10.1002/wcs.1545

This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided

the original work is properly cited.

© 2020 The Authors. WIREs Cognitive Science published by Wiley Periodicals LLC.

WIREs Cogn Sci. 2021;12:e1545. wires.wiley.com/cogsci 1 of 20

https://doi.org/10.1002/wcs.1545

seems nothing short of preposterous to suggest that the demise of countless indigenous communities, tribes, and states following European colonization was due to some inherent cognitive difference between those peoples. The same holds true for prehistoric population-level events, including hunter-gatherer displacements and replacements in Europe (Fu et al., 2016; Lazaridis et al., 2014) as well as the extensive dispersals following the Neolithic revolution, rapidly replacing Mesolithic populations' lifestyles and heavily influencing the ongoing genetic legacy of the regions affected by these dis- persals (Dolukhanov et al., 2005; Hofmanová et al., 2016; Skoglund et al., 2012). The case of the Neanderthal extinction, however, seems to be everything but obvious.

Working through the literature, two diametrically opposed approaches become apparent: One presupposes that Neander- thals were cognitively inferior and one posits a substantial similarity to anatomically modern humans. Both default positions feature a certain level of intuitive plausibility and are being vehemently defended by their proponents. The classic interpreta- tion argues that our species dispersed out of Africa once it reached a fully modern cognition and consequently clashed with Neanderthals, quickly replacing them and thereby demonstrating that those modern humans were qualitatively superior in some or many cognitive domains. Coming to this conclusion is admittedly easy as much of the Neanderthals' distinct physi- ology, like their protruding faces, their big brow ridges and their robust build, is typically classified as archaic when juxta- posed with the comparably gracile anatomy of modern humans. From the point of stating this kind of physiological distinctiveness, it is only a small step to reason that there might have been obvious differences in cognition, language and culture too. In contrast to this view, considering the vast evidence that such replacements happened within our species pre- sumably without the existence of a cognitive gap, it stands to reason that any interpretation of a transitional event relying on those factors needs to present independent evidence for the existence of such an inequality.

Taking either one of these approaches will inherently alter the perception of the available evidence, which at least partly explains the contemporary indecisiveness of the pertinent literature, containing both positions within a multi- tude of interdisciplinary perspectives. But problems extend further, as a clear picture is additionally obscured by meth- odological limitations, historic data artifacts and the wide range of disciplines involved, which inherently leads to more complications. This overview aims to present the relevant parts of this puzzle in a concise manner. A measured under- standing of research on Neanderthal cognition and their extinction will be reached first with the aid of discussing his- toric reasons for the current state of the debate as well as inherent problems and limitations of the research and second through a broad, triangulating synthesis of research findings. The discourse will culminate in a summary of what those findings mean for our understanding of Neanderthals' cognition, culture, and language.

2 | UNDERSTANDING THE PLAYING FIELD

In order to fully grasp the modern state of the research with its differing opinions on the subjects of Neanderthal cognitive ability, how anatomically modern humans' cognition compares to Neanderthals' and if cognition was a factor in the Nean- derthal extinction, it is necessary to incorporate a broader perspective than provided by the data of recent primary studies. It seems to be rather easy to recognize that in the history of research, the social consciousness of the community had, and still has, a profound impact on the interpretation of available data. As Hammond (1982, p. 4) puts it: “[L]ike any other human activity, scientific research is to some extent inseparable from the historical context in which the activity occurs.” Drell (2002, p. 2) concurs: “[I]deas and theories are never produced in a vacuum, but within certain given socio-historical situations.” Both of these authors present broad and deep accounts of the historic context, relevant tendencies and ideolo- gies since the discovery of the first Neanderthal specimens until the respective date of publication. Trinkaus and Ship- man (1993) dive even deeper into the subject of the history of Neanderthal research. This overview will be restricted to certain selected bullet points that are helpful to sketching out relevant biases that still influence modern opinions to a cer- tain degree. This brevity is motivated by giving room to two more sections that illuminate an expanded view on the sub- ject. One, starting off the following explanations, will deal with the structure, methodologies as well as limitations of the research and the other will address the interdisciplinary nature of the field.

2.1 | Inherent issues of interpreting prehistoric cognition and how to cope with these issues

The recognition that any direct verification or demonstration of the origins of human language and thought for a spe- cific location or point in time lie beyond the grasp of any of the sciences is not a new one or one that is hard to

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comprehend (e.g., Müller, 1861). Those events are rooted so deeply in human prehistory such that any reconstruction by definition will have to rely on inferences from indirect evidence. Feasible approaches include a wide range of possi- bilities. The exploration of early human physiology and genetics provides insights into a biological readiness to mod- ern-like cognition. The archeological record features technological and other cultural products as well as traces of behavior, for example, fossil leftovers from subsistence strategies that imply a certain level of cognition on the side of the producer. Additional insights into prehistoric behavior and therefore cognition can be attained from peripheral dis- ciplines like paleoclimatology and non-natural sciences that explore language and cognition as it is constituted today— including linguistics, psychology and sociology. Despite having such a vast collection of disciplines to draw inferences from, it has to be admitted that none of these fields of study provide grounds for unequivocal direct data into the origin of modern thought and language. Instead, all evidence is indirect and inferences drawn from this indirect evidence are basically the scientific equivalent of the children's game telephone: Senseful, yet error-prone interpretations. In other words, this indirect mode of reasoning creates a major issue as it multiplies the ways in which the reconstruction might fail.

This reliance on indirect inference, being less clear and less precise than direct evidence and enabling degrees of freedom in interpretation on the side of the researcher, therefore constitutes a first inherent shortcoming of the relevant research surrounding Neanderthal cognition—similar to the situation in other historical disciplines. Second, there exist major demarcation problems within this mode of indirect reconstruction. One of these lies in the identification and evaluation of proxies that potentially could enable an inference of prehistoric cognition. Without going into detail at this point, finding suitable proxies and assessing their reliability constitutes a divisive issue within the literature (Box 1 includes some examples of common methods for such inferences). Another such controversial demarcation problem arises by the recognition that modern humans' cognition is neither a single monolithic trait nor one that came into exis- tence through a clearly identifiable event. In this manner, even conceptualizing and defining methodological approaches is a contentious undertaking (e.g., Johansson, 2014, as well as Botha, 2008's discussion on “bridging” strate- gies). Third, and in a related sense, with modern cognition being a complex trait, the evolution of it was a temporally extended and complex process. That being said, intricate multifactorial structures or processes are incredibly hard to untangle even under substantially better research conditions, as can be seen in the deep divides within research on con- temporary humans' cognition and consciousness (e.g., Thagart & Stewart, 2014), and thus seem almost impossible to explore prehistorically. Fourth and exacerbating the last point further, human intuition shows a clear propensity toward narrative and explanatory clarity, often trapping interpretations and explanations in overly simplistic views even at expert levels. This results in false dichotomies and similar oversimplifications that might impede a measured and suitable evaluation of the evidence.

Summarizing these points, degrees of freedom within conceptualization, methodology, and interpretation open up a range of variability when judging the available evidence. Thus, attempting to reconstruct the origin of a basically

BOX 1 NONEXHAUSTIVE LIST OF COMMON METHODS IN INFERENCING PREHISTORIC COGNITION AND LANGUAGE USE

• Comparing the morphology and biomechanics of physiological features between (anatomically) modern humans and fossil specimens, including the cranium, the larynx, and the inner ear.

• Inferring if technological products of prehistoric populations are advanced enough to require modern cogni- tion and/or language ability, for example, habitual fire use for cooking and in hearths, sufficiently advanced tool production, use of clothing and string.

• Inferring if the behavior of those populations matches to that of modern hunter-gatherers in subsistence strategies (e.g., hunting a wide range of game and further exploiting a variety of vegetation and marine resources), resource acquisition/management (e.g., implementing long-distance routes and local, specialized workshops), social support, and so on.

• Documenting if artifacts or behavior reflect symbolic thinking, like in the use of color and ornaments, the production of cave paintings or performing intentional burials.

• Comparing the genetic landscapes of (anatomically) modern humans and archaic populations in relation to regions involved in for example, neurogenesis and language ability.

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modern cognition is problematic in multiple dimensions at the same time. Following this realization, research into pre- historic cognition seems inherently untrustworthy, but there are two themes that counteract these issues. One lies in the ever growing scope and performance of a number of relevant sciences, which increasingly provide pertinent data and perspectives, allowing for powerful insights into existing issues. The other lies in a careful and broad triangulation of the questions at hand by means of these collective interdisciplinary insights, building a strong case of abductive rea- soning. Any single piece of evidence can inherently be challenged when being considered on its own merit. This is espe- cially true if, as previously discussed for the topic of inferring prehistoric cognition, certain methodological limitations or shortcomings are in place. Yet if a large and varied body of evidence triangulates a common question and predomi- nantly aligns to one common point of conclusion, then the level of certainty within the research rises beyond the robustness of single findings. In regards to the approaches listed in Box 1, finding conclusions to the best of the avail- able evidence does not rest on a single mode of inference, but in a cumulative, consolidated synthesis of a vast range of data within all of those approaches.

2.2 | Data artifacts and bias in interpreting Neanderthal cognition and culture

Traditionally, modern cognition and language use were inferred through archeological means by a set of behavioral traits that seemingly sprung into place rather suddenly and exclusively within anatomically modern humans. This reconstruction as well as the related concept of a behavioral modernity that is tied to this cognitive revolution have received severe criticism in recent decades, subsequently increasingly losing support, and consequently cannot be seen as a currently prevailing expert opinion anymore (d'Errco & Stringer, 2011; d'Errico et al., 2003; Henshilwood & Mar- ean, 2003; McBrearty & Brooks, 2000; Shea, 2011; Villa & Roebroeks, 2014). Nevertheless, it is important to understand how this view came to dominate the relevant literature for decades, how it correlates with intuitive biases toward Nean- derthals and how it still influences the current debate despite being rejected by a growing body of literature. Though oversimplifying, the following sketch shall outline these points.

The first perspective to be elucidated lies in the data artifact that historically grounded the traditional view on Nean- derthal cognition. There are two points to consider: One, fossil and archeological discoveries were temporally and geo- graphically sparse for most of modern paleoanthropology, which left huge gaps in the reconstruction of the physiological and cultural evolution of prehistoric humans. Two, within this sparsity of findings, European discoveries dominated the totality of available evidence. Through this empirical framework, a seemingly clear picture emerged: Prehistoric Europe was inhabited by a technologically and physiologically apparently archaic population of Neander- thals for hundreds of thousands of years before it was rather rapidly replaced by anatomically modern humans, which went on to feature a more advanced technological and behavioral toolkit as well as the gracile physiology that is associ- ated with human modernity. On the basis of these observations, it seems natural to conclude that this must have been a case of a cognitively superior species outcompeting and replacing an archaic one. What has to be understood is that given this long-standing limited data sample, there was not sufficient impetus to seriously doubt this kind of interpreta- tion, especially considering the enforcement by inherent biases, which constitutes the second clarifying perspective to be presented here.

It cannot be denied that humanity's self-awareness includes an intuitive superiority complex toward anything differ- ent from ourselves. This seems blatantly obvious for intraspecies tribalism where groups of people may discriminate against each other for nationalist, ethnic or religious reasons, but also extends toward our place within nature. Throughout history, humans have imagined themselves as the pride of creation, separate and above all other life. In this respect, early interpretations on Neanderthal fossils, during the 19th century and well into the 20th century, stressed the deep divide between that species and our own, leading to a dismissive view of Neanderthals as brutes and evolutionary losers that were outcompeted by the superior modern humans. In the context of the outlined data artifact and these socio-historical biases, early paleoanthropology upheld a picture of Neanderthals as not quite human for decades and ultimately caused this dismissive mindset to be embedded within society from the layman to the expert. Despite the accumulation of evidence that challenges this interpretation and the later formulation of more nuanced opinions, this view persisted throughout the 20th century and still lingers in pockets of the contemporary literature.

In the expert realm, this narrow view eventually opened up by the middle of the 20th century, when a growing body of evidence and successively stronger, ever more careful methodologies allowed for a broader and more open discussion of the Neanderthal extinction. Howell (1952, p. 404) for example states “Whether the ‘classic Neandertals’ were already extinct by the time of this new peopling, or whether they were extinguished by, or hybridised with these invaders,

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remains a moot point at present,” stressing that the exact nature of the replacement was at the time impossible to answer on the basis of the available evidence. Following more findings Solecki (1971, 1975) granted Neanderthals the emotional and cognitive breadth of modern humans. This conclusion was based on his view that Neanderthals appar- ently shared a human level of empathy and possibly acted in symbolic ways. They seemed to care for their weak and sick as fossil finds indicated that some individuals lived to an old age despite suffering debilitating ailments that made social support necessary for survival. He also concluded from unusually high amounts of pollen that were found in the soil next to a Neanderthal specimen that this individual was buried together with flowers in a symbolically intended rit- ual, though it has to be added that this interpretation was later discredited as there are competing explanations for how the pollen could have gotten there (Sommer, 1999). Nevertheless, these and further findings tended to a slowly chang- ing landscape of expert opinions on Neanderthals' humanity that continues to be in motion even today.

An explanation for this slow-moving, still incomplete and consistently controversial development takes two more steps. First, scientific progress necessarily includes a certain degree of conservatism. At the forefront of the scientific horizon, hypotheses, models, and theories are fiercely tested and debated—and only after an arduous process of harsh scientific scrutiny, a scientific consensus may emerge. Thus, not every new finding should overturn already established consensus opinions, but instead needs to be put into a carefully constructed context within the ongoing research. In this manner, new findings that contradict such a consensus need to build up to a threshold where the cumulative nature of the previous research history is successively overridden in order to shift to a new consensus. This in itself constitutes a healthy part of the scientific endeavor and provides some inherent quality control. Nevertheless, and second, this aggre- gate nature of scientific progress has its downsides too. For once, this process commonly leads to a quite vast and opaque body of literature that can only be efficiently navigated by the experts of a field. Furthermore, it is possible for things to take on a pathological quality when the following two criteria are in place: First, a previously consensus but now outdated opinion has accumulated massive credence throughout a lengthy research history and has firmly embed- ded itself in the cultural memory. Second, the field of research suffers from vast degrees of freedom as discussed in the previous section and therefore lends plausible deniability to proponents of the outdated opinion. It stands to reason that the research on Neanderthal cognition resembles such a case, though this point cannot be further substantiated here.

2.3 | General problems within interdisciplinarity and a pertinent formation of theory

The following will be of particular brevity, suggesting the need for an interdisciplinary approach and then summarizing some of the relevant factors that impede any attempt of synthesizing interdisciplinary research beyond what has been presented so far. To begin with, it is necessary to stress that the research question of the evolution of modern cognition, including correlates like language ability, within human ancestry is one that is to be answered in an interdisciplinary framework. It is a given that multiple disciplines have perspectives and insights into the issue—from biology, biochem- istry, and neuroscience through paleoanthropology and archeology to psychology and linguistics. However, a simple multidisciplinary approach will not yield the answers sought, as each and every one of these fields of research consti- tutes a blind spot for each other respectively. Instead, and drawing a comparison with the parable of the blind men and an elephant, these disciplines will have to work toward an interdisciplinary approach that brings their diverse perspec- tives together. This, in turn, entails a number of challenges that go beyond what has already been discussed.

First, those previously mentioned demarcation problems and the struggle with a multifactorial historical process are enhanced when put into such an interdisciplinary context. Further, the human tendency to conceptual clarity leads to miscommunication in a similar manner. This extends to a mismatch of technical jargon, methodologies, perspectives, and axioms between the collaborating disciplines. In a way, every expert involved is each other's layman, mis- appropriating each other's work and tending to a frustrating communication style full of misunderstandings and ambi- guities. In fact, the relevant literature already features countless examples of carelessly constructed arguments when borrowing work or terminology from outside one's own area of expertise. This is especially true for certain relatively recent discoveries like the FOXP2 gene and mirror neurons, which quickly garnered a widely misguided mania of spec- ulation about possible implications for human thought and language. It also holds true when primatologists and orni- thologists relate their research on communicative capabilities of certain nonhuman species to linguistic terminology, erroneously speaking of syntax and phonology whenever they identify structure in vocalizations. Yet, in the face of these exemplifying misadventures and all those problems that have been laid out up to this point, there is no shortcut to viable answers. Any conclusive reconstruction of the origins of modern cognition, language, culture and more will necessarily have to go through the path of an excruciating interdisciplinary synthesis.

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3 | THE EVIDENCE SURROUNDING NEANDERTHAL COGNITION

It has been stated previously that the concept of a behavioral modernity that established itself in a rather quick and holistic manner has attracted considerable criticism during the last two decades (see again McBrearty & Brooks, 2000, and others). During the same time span, archeological, and genetic evidence strengthened the interpretation of a Nean- derthal population that was culturally and cognitively closer to their contemporary anatomically modern humans than previously thought, albeit not exactly the same (among others: Dediu & Levinson, 2013; Dediu & Levinson, 2018; Finlayson et al., 2012; Hoffecker, 2018; Roebroeks & Soressi, 2016; Villa & Roebroeks, 2014). The following two sections will elaborate on both of these points and paint the current state of the evidence according to the relevant expert opin- ions as well as some of the reasons for these currently changing opinions. After this, in a third section, the discourse will go one step beyond the summary of archeological and paleoanthropological findings and triangulate the issue of Neanderthal cognition through an extended interdisciplinary framework.

3.1 | The antiquity and cumulativeness of cognitive modernity/variability

As has already been sketched out, some scholars granted Neanderthals an essentially modern mental world by the 1970s and the cognitive revolution scenario has come under scrutiny by the turn of the millennia. Yet, both the notions that Neanderthals were cognitively and culturally inferior to anatomically modern humans and that modern cognition came into being only within anatomically modern humans by more or less abrupt means persist within a strong tradi- tion (Berwick & Chomsky, 2016; Binford, 1981; Noble & Davidson, 1996; Stringer & Gamble, 1993; Tattersall, 2015). These authors uphold the argument that symbolic behavior, and with it modern cognition, language as well as a suit of modern technologies and behaviors, came about roughly during the tens of thousands of years before anatomically modern humans started to spread into Europe to ultimately replace the Neanderthals. In his manner, despite being on the side of reappraising Neanderthals' capabilities, d'Errico et al. (2003, p. 1) state “No firm evidence of conscious sym- bolic storage and musical traditions are found before the Upper Paleolithic.” and especially through nonexperts con- cerning the fields of paleoanthropology and archeology, views of a sudden emergence of modernity survive up until today, as in Berwick & Chomsky (2016, pp. 38–39). The following paragraphs will provide evidence that reflects the cur- rent state of the research, which came to acknowledge that multiple aspects of supposedly modern behaviors feature a surprisingly deep and complicated history within the human evolutionary lineage.

In this sense, while control of fire has to be seen as a drawn-out and for most of its history incomplete process that possibly started up to 2 million years ago in Homo erectus (Chazan, 2017), modern-like behavior includes a regular and multi-varied use in hearth-like structures. Such behavior was classically seen as having been established in the cogni- tive revolution toward behavioral modernity. This was later contrasted by studies (e.g., Goren-Inbar et al., 2004) as well as reviews (e.g., Gowlett, 2016) that locate these features as becoming widespread roughly between 800,000 and 400,000 years ago, enabling regular cooking as well as new tool technologies like hafting that is facilitated with adhe- sives. Concerning such tools, the stone work production alone reached whole new levels of complexity and planning by about 300,000 years ago. The techniques and modes of production that roughly developed around that timeframe involved a temporally and hierarchically extended process, requiring extended planning. This implies that there might not be as sharp a distinction to later human populations as has been suggested by the traditional literature (see for this point for example, Eren & Lycett, 2012). Further, in some areas, stone tool production at local workshops reached a level of intensity that Foley & Lahr (2015, pp. 10–11) described as a “humanly induced environmental change of a larger geographic scale,” while the acquisition and transport of high-quality materials involved long distance routes, sometimes exceeding 100 km (Blegen, 2017; Brooks et al., 2018). Such long-distance transport and the implementation of large-scale local workshops also includes pigment use for color production from at least 250,000 years onward, which indicates early symbolic thought (Barham, 2002; Brooks et al., 2018). Needless to say, all of those features were previ- ously linked to the interpretation of an Upper Paleolithic revolution.

Stone blades are a good place to start in order to discuss a more concrete example of technology within cultural evo- lution. Shea (2011, p. 6) states on this matter that “[i]n Europe, evidence for the systematic production of prismatic blades coincides with the first appearance of H. sapiens in Upper Paleolithic contexts. For decades, Paeolithic archaeolo- gists have routinely inferred a causal connection from this association.” In other words, a cornerstone in the interpreta- tion of the cognitive revolution and Neanderthal inferiority was the emergence of blade technology within anatomically modern humans directly antecedent to or coincident with the replacement of the Neanderthals. This stands in contrast

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to findings discussed by Muller & Clarkson (2016, p. 2), who elaborate that “it is now clear that technological evolution is far from linear, but is instead multidirectional, branching and recursive.” In this sense, the mentioned “blade tech- nology is securely dated to well before the Upper Palaeolithic, is not confined to anatomically modern humans, and appears and disappears in many regions over time.” This pattern of technologies appearing and disappearing through- out hundreds of thousands of years has been described by other authors, including d'Errco & Stringer (2011) and Meignen (2012), who review a broad literature that demonstrates a mosaic of cultural innovation and loss that seems to point to a rich dynamic of population fluctuation and movement that underlies this patchwork of cultural evolution. These findings suggest that there was no late biologically based cognitive threshold to at least some supposedly modern behaviors and technologies. Instead, this cultural gain and loss of technologies and behaviors can be made sense of more fittingly by different means.

Minding this state of the research while trying to identify the mechanism that underlies the documented pattern, a tentative conclusion lies in the observation that demography and population dynamics may play a major role in explaining technological-cultural patterns within and beyond the temporal limits of our species. Indeed, according to some authors, this interpretation can be seen as a major complementary, or even competing, explanatory approach to the default hypothesis, which posits that technological and cultural evolution of this deep prehistory is causally linked to biological changes that directly lead to changes in cognition. Eren & Lycett (2012, p. 9) make this quite clear: “[B]ehavioral changes that eventually emerge […] may be more the product of demographic change and increased con- nectivity of social networks than they were, necessarily, of fundamental cognitive changes.” Powell et al. (2009, p. 1298) similarly make a point of explaining the emergence of “modern behavior without invoking increased cognitive capac- ity” and another angle on this comes from the Australian archeological record. Anatomically modern humans arrived in Australia by about 65,000 years ago (Groucutt et al., 2015) and without a doubt featured the full biological and cogni- tive range of modern humans, yet the cultural evolution following the colonization was patchy and incomplete in regards to fully modern symbolic behavior until about 6,000–7,000 years ago, leading Brumm & Moore (2005, pp. 167– 168) to the conclusion that “if modern symbolic behavior in early Australia produced a patchy archaeological record, there is no clear reason for rejecting the ‘modernity’ of the Middle Pleistocene record of the Old World solely on the basis of its patchy distribution.” More insights into such cultural processes, but concerning movement in the opposite direction, are given by Henrich's (2004) reconstruction of the Tasmanian archeological record, which demonstrates the disappearance of technologies and behaviors over successive millennia after rising sea levels isolated Tasmania from the mainland Australian population, heavily restricting social networks and cultural exchange.

Additional research will be necessary to reach a new consensus on this issue, but the current evidence is pointing into the direction of what has been described in this section. More on the role of demography in cultural dynamics and the supporting evidence will be discussed in the next two sections when exploring Neanderthals' behavior according to the current state of research. Here, it shall simply be summarized that a number of technologies and behaviors that pre- viously were seen as part of an Upper Paleolithic cognitive revolution feature an early onset, partly predating the emer- gence of our species. This opens up the possibility of modern cognition having a much deeper prehistory than traditionally assumed, with cultural evolution not necessarily being constrained exclusively by biology, but also, or pos- sibly primarily, by other factors like ecological and demographic pressures and developments. Indeed, it seems pretty obvious that modern cognition and language is necessary, yet not sufficient, for cultural and technological innovation to spiral into our modern globalized world, as some of today's indigenous people live an essentially hunter-gatherer life- style, featuring technology not clearly superior to what was already in use hundreds of thousands of years ago, simply because neither the ecology nor the demography introduces the momentum that leads down the line toward an even- tual industrialization. Instead, those immediate-return societies (Woodburn, 1982) feature a stable survival strategy “[w]here goods are free” (Lewis, 2015, p. 1.) and therefore, there is no shortage or inequality of resources to be over- come by technological or behavioral innovation.

3.2 | The comparability of anatomically modern human and Neanderthal behavior

In the introduction to this overview, the traditional and quite intuitive way of thinking on the issue of Neanderthal cog- nition has been sketched out already. Nevertheless, it makes sense to exemplify this position in the words of its propo- nents. An especially succinct choice of words comes from Berwick & Chomsky (2016, p. 38): “[S]omething must have set us apart from the Neandertals, to prompt the relentless spread of our species who had never crossed open water up and out of Africa and then on across the entire planet in just a few tens of thousands of years.” Whenever this style of

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argument is brought up, it almost certainly goes on to invoke cognition and/or language as the thing that has set us apart from Neanderthals, and so it does in Berwick & Chomsky (2016, pp. 38–39). Reasons for why this intuition is so strong have been brought up in earlier sections. An opposed tradition claims that this line of reasoning might be wrong and that an assumption of Neanderthal inferiority is not feasible. Instead, the declaration of a cognitive or linguistic inequality necessitates independent evidence in favor of this interpretation. The following shall explore this point fur- ther and then assess the available evidence, primarily through archeological and comparative means. The next section will expand on additional points, sketching out, among other things, the demography and population genetics sur- rounding the Neanderthal extinction.

Interdisciplinary efforts between archeology, paleogenetics, and related disciplines show that a number of prehistor- ical replacements played out equally fast or even faster than the Neanderthal replacement. Sometimes, a roughly simi- lar geographical pattern of progression is involved, and often, the genetic continuity of the region is heavily influenced by the dispersing population. Two telling examples within our species have been mentioned before: The Neolithic expansion into Europe (Baralesque et al., 2010; Hofmanová et al., 2016; Malmström et al., 2009) and Upper Paleolithic transitional events between European hunter-gatherer populations (Fu et al., 2016; Lazaridis et al., 2014). Naturally, these events do not involve any necessity of a biologically based cognitive or linguistic inequality. The same applies to similar demographic dispersals and replacements between Neanderthal subpopulations that have been genetically inferred (Fabre, Condemi, & Degioanni, 2009; Hajdinjak et al., 2019; Peyrégne et al., 2019). Instead, population dynam- ics as well as cumulative culture may inherently lead, in combination with certain ecological and demographic condi- tions, to “explosive demographic transitions” (Ghirlanda & Enquist, 2007, p. 591) or “abrupt cultural regime shifts” (Aoki, 2015, p. 6). All of this strengthens the idea that the Neanderthal extinction may not by itself be seen as grounds for concluding that it was driven by a cognitive inferiority. Instead, this proposition needs to be addressed indepen- dently, as proposed by the younger of the two default approaches to this research question.

In respect of Neanderthals and their contemporary anatomically modern humans, archeological studies and reviews that state a comparability in tool production have accumulated a noteworthy history by now. Schwarcz et al. (1989, p. 658) for example, state on the question of the temporal relationship between Neanderthals and anatomically modern humans in the Levant that the situation “is obscured by the fact that the artifacts used by these two populations are indistinguishable.” In a similar manner, many others state a comparability of contemporary anatomically modern humans' and Neanderthals' technological products (Dediu & Levinson, 2013; Dediu & Levinson, 2018; Eren & Lycett, 2012; Hoffecker, 2018; Muller & Clarkson, 2016; Roebroeks & Soressi, 2016; Villa & Roebroeks, 2014). These include not only advanced stone work, but specialized soft tools made out of bone or wood as well as composite tools using tar, which are enabled by a free and variable use of fire, and throwing spears open up the spectrum of hunting strategies employed by modern hunter-gatherers (Aranguren et al., 2018; Degano et al., 2019; Goren-Inbar et al., 2004; Gowlett, 2016; Milks, Parker, & Pope, 2019; Niekus et al., 2019; Roebroeks & Villa, 2011; Schoch, Bigga, Böhner, Rich- ter, & Terberger, 2015; Soressi et al., 2013; Thieme, 1997). It has to be noted that most these technologies and behaviors were, according to the dating in those studies, in place by about 350,000 years ago. Further, the interpretation of those spears having been used as throwing rather than thrusting weapons is primarily based on their inherent weight distri- bution, but is consistent with abnormal skeletal and joint wear in Neanderthals that mirrors those documented in pro- fessional throwing athletes (Faivre et al., 2019). At the same time, other wear and kinematic analyses conclude that Neanderthals were adept precision workers on top of their effective power grip capabilities (Feix, Kivell, Pouydebat, & Dollar, 2015; Karakostis, Hotz, Tourloukis, & Harvati, 2018; Niewoehner, Bergstrom, Eichele, Zuroff, & Clark, 2003).

Documentation of the types of animals hunted by Neanderthals further substantiates the inference of ranged weap- onry and is in itself impressive as it fully reflects modern hunter-gatherer capabilities. From big game and even bears through small game and birds to fish and other seafood including seals and dolphins, Neanderthal subsistence strate- gies were varied and systematic—and in contrast to common misconception also heavily relied on nuts, mushrooms, and a multitude of other vegetation, though heavy reliance on meat was highly prevalent during cold climatic phases (Bocherens et al., 2016; El Zaatari, Grine, Ungar, & Hublin, 2011; Estalrrich, El Zaatari, & Rosas, 2017; Finlayson & Finlayson, 2016; Hardy et al., 2013; Morin et al., 2019; Power et al., 2018; Romandini et al., 2018; Stringer et al., 2008; Trinkaus, Samsel, & Villotte, 2019; Zilh~ao et al., 2020). Exploiting these resources in a specialized way resembling “logistic behavior” (Marín et al., 2020, p. 22), Neanderthals implemented “complex land use patterns, adapting to diver- sified landscapes and climates” (Moncel, Fernandes, Willmes, James, & Grün, 2019, p. 1) and thereby demonstrate “a complex scenario of extensive knowledge and intensive exploitation of the landscape” (de Soler et al., 2020, p. 1). Fur- ther, Neanderthals not only cooked, but also purposefully used naturally occurring medications and maintained mouth hygiene through the use of toothpicks (Estalrrich, Alarcón, & Rosas, 2017; Hardy, 2018; Hardy et al., 2012; Weyrich

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et al., 2017). Finally, the production and use of cords by Neanderthals has been documented, displaying another piece of a, according to the classical view, modern technology that allows for a range of possible behaviors mirrored by mod- ern hunter-gatherers (Hardy et al., 2013, 2020).

Beyond Neanderthal dietary strategies and technologies being comparable to their contemporary anatomically mod- ern humans and mirroring today's hunter-gatherer capabilities to a large extent, there is ground to be granted when it comes to resource management and symbolic behavior. First, raw material and color transport over dozens of kilome- ters for purposeful usage is documented to minimally 200,000–250,000 years ago (Roebroeks et al., 2012), giving this behavior a similar temporal depth as in the African archeological record. Further, Neanderthals perforated and colored seashells, apparently to string them, and behaved in similar ways with beads, geodes as well as prepared feathers and talons, sometimes carving geometric patterns on the materials and featuring some of this behavior by at least 130,000 years ago (Cârciumaru, Niţu, & Cîrstina, 2015; Caron, d'Errico, Del Moral, Santos, & Zilh~ao, 2011; Finlayson et al., 2012; Hoffmann, Angelucci, et al., 2018; Majki�c, d'Errico, & Stepanchuk, 2018; Majki�c, Evans, Stepanchuk, Tscelykh, & d'Errico, 2017; Morin & Laroulandie, 2012; Peresani, Fiore, Gala, Romandini, & Tagliacozzo, 2011; Per- esani, Vanhaeren, Quaggiogiotto, Queffelec, & d'Errico, 2013; Radoviči�c, Sršen, Radoviči�c, & Frayer, 2015; Welker et al., 2016; Zilh~ao et al., 2010). Jaubert et al. (2016) describe extended constructions more than 300 m deep into a cave system which possibly were used with a ritualistic purpose by Neanderthals at around 176,000 years ago and Hoffmann, Standish, et al. (2018a), Hoffmann, Standish, et al. (2018b), and Hoffmann et al. (2020) document some of the currently oldest known cave paintings to be of Neanderthal origin. Rodriguez-Vidal et al. (2014) argue that rock engravings in Gibraltar made by Neanderthals have a symbolic component to them and according to d'Errico et al. (2018), carvings on bones suggest that they were used to count, indicating an explicit, symbolic number sense beyond the innate num- ber sense that most animals feature (Petrazzini, Agrillo, Izard, & Bisazza, 2016; Scarf, Hayne, & Colombo, 2011; Wood- ruff & Premack, 1982). Hardy et al. (2020) also suggest a Neanderthal number sense by presenting evidence of Neanderthal cord production, which implies knowledge of numbers and sets in order to pair and intertwine individual fibers into the cord.

Another point to address is Neanderthals' attitude in social contexts. Hublin (2009), Thorpe (2016), Trinkaus and Villotte (2017), Spikins, Needham, Tilley, and Hitchens (2018), and Spikins et al. (2019) present or review evidence for social support by Neanderthal groups toward the hurt and sick and while Neanderthal burials have been a contentious issue since Solecki (1971, 1975), evidence is accumulating that “Neandertals seem to show a fairly complex mortuary behavior” and that “[i]t is […] likely that some Neandertal groups also purposefully buried their dead, i.e., that they practiced funerary behavior” (Gómez-Olivencia et al., 2018, p. 30; further see Arsuaga et al., 1997; Hovers, Kimbel, & Rak, 2000; Pettitt, 2002; Rendu et al., 2014, 2016; Schwarcz et al., 1989; Valladas et al., 1987). As for Berwick & Chom- sky stressing anatomically modern humans' crossing of open waters as evidence for their cognitively special status (Ber- wick & Chomsky, 2016, p. 38, a.k.a. the quote from the beginning of this section), there is some evidence that Neanderthals might have done seafaring of their own, though if this behavior was systematic or what exact form this behavior might have taken is unknown (Carter et al., 2019; Ferentinos, Gkioni, Geraga, & Papatheodorou, 2012; Papoulia, 2017; Strasser et al., 2010).

Overall, there seems to be no significant qualitative difference in behavior, culture or technology between Neander- thals and their contemporary anatomically modern humans right up to the replacement of the Neanderthals (Box 2 dis- plays a condensed summary of the evidence discussed so far). At the same time, this does not mean that there were no differences at all. As many of those authors that were cited as granting Neanderthals an extended, possibly full, human- ity readily admit, there seems to have been quantitative differences in the scale of for example, symbolic behaviors. How this is to be interpreted, however, is a point of contention. Some might fall back to the intuitive default position of positing an inherent difference in cognition between the species, while it could be once more an issue of ecology and demography. This is suggested by for example, the previously mentioned Australian archeological record, where the same patchiness can be observed far into modern human occupation. More perspectives of Neanderthal demography and its role will be discussed in the next section.

3.3 | Triangulating Neanderthal cognition beyond archeology

The previous sections established two main insights from the African and European fossil record: First, the emergence of behavioral patterns that reflect a basically modern cognition seems to have been a temporally and geographically extended process that began before the dawn of our species, and second, this process featured an overall slow cultural

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accumulation throughout a complex mosaic of local innovation and local loss of technologies and behaviors, involving both the African and Eurasian continents. This extended cultural patchwork, reconstructed from archeological and fos- sil evidence, leads Eren & Lycett (2012, p. 9) to the realization “that cognitive capacities in different species of Middle- Late Pleistocene hominins are not as sharply differentiated as previous generations of scholars postulate.” A growing number of researchers agree, either explicitly or implicitly, with this notion that demographic patterns and population dynamics were possibly or probably more important than biology in explaining technological-cultural patterns within the taxonomic complex that includes anatomically modern humans, Neanderthals and in extension the Denisovans as well as the common ancestors of these taxa (Bocquet-Appel & Degioanni, 2013; Carrión & Walker, 2019; Dediu & Levinson, 2013; Dediu & Levinson, 2018; Eren & Lycett, 2012; Hoffecker, 2018; Hosfield & Cole, 2018; Muller & Clarkson, 2016; Powell et al., 2009; Roebroeks & Soressi, 2016; Shea, 2011; Villa & Roebroeks, 2014). Adding to these insights are further data points from the fossil record as well as perspectives from paleogenetics, paleoclimatology, ecol- ogy, and modeling of population dynamics, as will be discussed in the following.

A first point to be made lies in the assessment that anatomically modern humans' population size was possibly up to an order of magnitude greater than the Neanderthals' and also featured a greater population density in at least some geographic areas, as supported by a number of studies (Bocquet-Appel & Degioanni, 2013; Castellano et al., 2014; Fabre et al., 2009; Mafessoni & Prüfer, 2017; Mellars & French, 2011; Prüfer et al., 2014; Vaesen, Scherjon, Hemerik, & Ver- poorte, 2019). This situation can be made sense of for ecological reasons alone as climatic conditions in Europe were comparably harsh and included climate cycles that regularly thinned out and pushed back Neanderthal subpopulations into Eurasian refuges, before climate conditions allowed them to spread back into a larger Eurasian territory (e.g., Bradtmöller, Pastoors, Weninger, & Weniger, 2012; also reflected in changing lithic strategies as in for example, Shipton et al., 2013). These cycles can be exemplified by the reoccurring colonization and abandonment of England by Homo antecessor, Homo heidelbergensis, and Neanderthals over the course of almost a million years (Bates, Pope, Shaw, Scott, & Schwenninger, 2013; Stout, Apel, Commander, & Roberts, 2014; Streeter et al., 2001; Stringer, Trinkaus, Roberts, Par- fitt, & Macphail, 1998), which is mirrored by anatomically modern human occupation, abandonment and recolonization that cycled multiple times before a stable population settled from around 12,000 years ago onward (Pettitt & White, 2012).

From such a fundamental demographic imbalance between Neanderthals and anatomically modern humans, fur- ther worsened by reoccurring demographic weaknesses within the Neanderthal population and a resulting decrease in Neanderthals' genetic fitness, completely neutral interpretations of demographic pressure, and migratory behavior on the side of anatomically modern humans are potentially sufficient to explain the Upper Paleolithic replacement (Degioanni, Bonenfant, Cabut, & Condemi, 2019; Goldfield, Booton, & Marston, 2018; Kolodny & Feldman, 2017; Mellars & French, 2011; Vaesen et al., 2019). Indeed, there is evidence that the dispersal, a.k.a. the demographic push, of anatomically modern humans that resulted in the Neanderthal demise matches up with such a climate cycle and the

BOX 2 OVERVIEW OF TECHNOLOGY AND BEHAVIOR SUPPORTING NEANDERTHALS' COGNITIVE COMPARABILITY

• Long-distance transport of materials and establishment of local workshops. • Free use of fire for cooking and technology, for example, production of hafting materials. • Long-time and hierarchical planning in tool production and resource management. • Production of tools classically seen as modern, like blade technology and ranged weapons in the form of throwing spears.

• Early pigment use and production of ornaments out of sea shells, beads, geodes, feathers, and talons. • Production of cave paintings and possibly construction of ritual sites. • Burial behavior, though possibly not fully compatible with modern funerary behavior. • Roughly modern, seasonal subsistence strategies including plants, mushrooms, nuts, small and big game, birds, sea food including fish and marine mammals.

• Social (health) care toward oneself as well as the group, including natural remedies as well as mouth hygiene via toothpicks.

• Seemingly crossing of open water.

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resulting demographic weakness on the side of the Neanderthal population (Alcarez-Castaño et al., 2017; Bradtmöller et al., 2012; Melchionna et al., 2018; Staubwasser et al., 2018; Wolf et al., 2018). A volcanic super-eruption during this glacial period, causing a volcanic winter in southern and eastern Europe and therefore impacting Neanderthal refuges, might have weakened the Neanderthal population further (Costa et al., 2012; De Vivo et al., 2001). Finally, slight differ- ences in fertility, reproductive or mortality rates, disease susceptibility, demands on caloric intake or cultural behavior might have also constituted significant factors without necessarily having cognitive implications (Degioanni et al., 2019; Goldfield et al., 2018; Pagano, Márquez, & Laitman, 2019; Trinkaus, 1995), though some of these aspects seem to be remarkably close to the contemporary anatomically modern human counterparts (e.g., Trinkaus, 2011).

The replacement itself then did not go without genetic intermixing, adding to the notion that the overall pattern looks more like later demographic events including, but not limited to, the aforementioned prehistoric hunter-gatherer replacements and the Neolithic dispersal into Europe (also see again Bradtmöller et al., 2012, for how later climatic cycles and correlating population replacements in Europe mirror the Neanderthal extinction). This genetic introgres- sion from Neanderthals as well as Denisovans is continually being explored and modeled, identifying both positive and negative selection pressures on introgressed DNA (Dannemann & Kelso, 2017; McCoy, Wakefield, & Akey, 2017; Gittelman et al., 2017; Gregory et al., 2017; Lipson & Reich, 2017; Nielsen et al., 2017; Racimo, Marnetto, & Huerta- Sánchez, 2017; Dannemann & Racimo, 2018; Enard & Petrov, 2018; Slon et al., 2018). According to these studies and reviews, the genetic impact of Neanderthals on modern Eurasian populations appears to be rather small, yet it should be considered that the genetic contribution of local populations in later replacements have been shown to be surpris- ingly small too (Again: Baralesque et al., 2010; Skoglund et al., 2012; Lazaridis et al., 2014; Fu et al., 2016; Hofmanová et al., 2016), and despite claims of selection against Neanderthal DNA in the genome of anatomically modern humans supposedly being evidence of Neanderthal inferiority (e.g., Berwick & Chomsky, 2017, pp. 167–168), studies propose that effective population sizes and gene flow between modern human populations sufficiently explain the pattern of genetic intro- gression and the selection against the introgression (Juric, Aeschbacher, & Coop, 2016; Petr, Pääbo, Kelso, & Vernot, 2019).

Overall, intermixing between anatomically modern humans, Neanderthals, and Denisovans seems to have been fre- quent and ultimately ended in the archaic populations being partly absorbed into the larger modern human populations (see the aforementioned, especially Slon et al., 2018). To quickly address Neanderthal contributions to modern Eurasian genetic variation: Identified areas include patterns of skull and brain dimensions (Gregory et al., 2017), skin tone, hair color, height, sleep patterns, mood, smoking behaviors (Dannemann & Kelso, 2017), reduced susceptibil- ity to certain pathogens (Enard & Petrov, 2018; Gittelman et al., 2017), reduced rate of miscarriage, less early pregnancy bleeding (Zeberg, Kelso, & Pääbo, 2020), metabolism, altitude tolerance, and a whole suit of further variables (as reviewed for example, by Dannemann & Racimo, 2018). The phenotypical relevance of these and other findings, includ- ing regulation of brain-related regions (McCoy et al., 2017), however, has as of yet not been sufficiently explored. This uncertainty leaves inferences on Neanderthal cognition from this area of research in a rather indecisive state that neces- sitates further exploration by future studies. Moving away from identifying Neanderthal traces in the modern genome, some additional paleogenetic evidence will be discussed below.

But first, an obligatory point to be addressed, though its implications for cognition are more limited than intuition would assume, lies in the fact that Neanderthals' brain volume was comparable to anatomically modern humans' (Buck & Stringer, 2014). A further discussion that is connected to modern cognition concerns the possibility of language abil- ity. It should be noted that there is some physiological and genetic evidence to suggest that Neanderthals perceived and used speech sounds in a similar manner as modern humans. The auditory capacities seem to have been fine-tuned to a basically modern constitution by H. heidelbergensis (de Boer, 2012; Martínez et al., 2004; Martínez et al., 2008; Martínez et al., 2013; Stoessel et al., 2016) and the Neanderthal hyoid bone is documented to be physiologically and biomechani- cally essentially modern-like, implicating the possibility of a roughly human speech capacity (Arensburg et al., 1989; Arensburg & Tillier, 1991; D'Anastasio et al., 2013). These physiological insights are complemented by stone tool ana- lyses as well as experimental data evaluating the demands on cognition in tool production and assessing the role of communication in learning, concluding that even pre-Neanderthal technologies likely necessitated active teaching and involved extensive planning (Lombao et al., 2017; Lycett, von Cramon-Taubadel, & Eren, 2016; Morgan et al., 2015; Stout, Hecht, Khreisheh, Bradley, & Chaminade, 2015; but see Cataldo, Migliano, & Vinicius, 2018, on how speech alone was also a poor model). Adding to this picture, the modern variant of the FOXP2 gene, which is, among a multi- tude of other functions (Nudel & Newbury, 2013), linked to a normative development of language, has been shown to be present in both Neanderthals and Denisovans and therefore seems to date back to at least their last common ances- tor with the lineage toward anatomically modern humans (Krause et al., 2007; Paixáo-Côrtez, Vicardi, Salzano, Hunemeier, & Bortolini, 2012; Reich et al., 2010).

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Maricic et al. (2013) document a relatively recent evolutionary change in the regulation of FOXP2 that is widespread in modern populations and therefore suspect to have found a plausible candidate for a positive selective sweep within anatomically modern humans, but then again, the ancestral allele occurs with up to around 10% frequency in some populations (p. 849) and therefore seems to fall within modern human phenotypic variation. Further, Atkinson et al. (2018) conclude that there seems to be no candidate for a selective sweep on or around FOXP2 that is exclusive to the evolutionary history of anatomically modern humans. Of course, there are other linguistically as well as cognitively relevant regions in the human genome. Concerning those, there exists further evidence of regulatory changes within the modern human lineage as well as downregulation of introgressed genes that are expressed in the brain (McCoy et al., 2017; Somel, Liu, & Khaitovich, 2013). Yet, until these findings can be substantiated and explored in more detail, the genetic comparison of Neanderthals and (anatomically) modern humans in regards to cognition and language has to be seen as tentative and inconclusive. Among others, Mountford and Newbury (2018), DeSalle and Tattersall (2018) and Martins, Marí, and Boeckx (2018) make the point that the relevant genomic landscape is highly complex and so far poorly understood, but that this exact complexity implies that a gradual, selective process was involved. This means that high-functioning, roughly modern cognition and language use were already in place when pressures on efficiency and easier bioavailability shaped the modern human genotype.

A last perspective to take comes from archeological finds that show a temporary presence of anatomically modern humans in southeastern Europe at around 210,000 years ago (Harvati et al., 2019) and genetic evidence that demon- strates contact between early Neanderthals and the line to anatomically modern humans between 460,000 and 219,000 years ago (Posth et al., 2017). Even before these recent insights, it has been known that anatomically modern humans dispersed out of Africa more than once and at much earlier points in time than traditionally thought, reaching China by roughly 100,000 years ago and Australia by 65,000 years ago (Cabrera, Marrero, Abu-Amero, & Larruga, 2018; Clarkson et al., 2017; Groucutt et al., 2015; Liu et al., 2015). Under the assumption that first, cognitive ability was the major function of such dispersals and replacements, and second, anatomically modern humans were cognitively and behaviorally clearly superior to Eurasian local populations, there is a question to be raised: Why did these earlier dis- persals not yield the same result as the later Upper Paleolithic dispersal? This dilemma cannot be untangled by trying to argue that full modernity was only reached later, as Khoe-S�an groups in Africa are undoubtedly fully modern humans, yet split from other African populations more than 100,000 years ago (Gronau, Hubisz, Gulko, Danko, & Siepel, 2011; Schlebusch et al., 2012). Ultimately, even while replacing the Neanderthals, anatomically modern humans may have adopted at least some survival strategies from the Neanderthals as there apparently has been a continuum in selected behaviors and technologies from Neanderthals to anatomically modern humans (Finlayson & Finlayson, 2016; Soressi et al., 2013; Tostevin, 2007). Closing this overview, Box 3 outlines the key takeaways from the review section, while some elaborations and broader conclusions follow below.

BOX 3 KEY TAKEAWAYS IN A ROUGH OUTLINE OF THE OVERVIEW

• Inferring cognition simply from documenting a replacement between populations is inherently implausible. • Independent archeological evidence shows Neanderthals as behaviorally and technologically qualitatively comparable to anatomically modern humans in for example, tool production, symbolic products, subsistence strategies, and social support.

• Genetically and physiologically, there were differences, but also considerable overlap, with for example, the modern variant of the FOXP2 gene, a biomechanically basically modern hyoid bone and inner ear and a com- parable cranial volume.

• Regular contact and local extinctions/replacements happened both ways between Neanderthals and anatomi- cally modern humans and further show cultural continuities.

• Demographic dynamics seem to explain the documented cultural patchwork that includes both Neanderthals and anatomically modern humans; demographic weakness could be the strongest contender to explain the eventual Neanderthal extinction.

• While the exact relationship between anatomically modern humans and Neanderthals is still uncertain, an at least rough comparability in cognition, culture and language ability is strongly suggested by the evidence.

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4 | CONCLUSIONS

The research on the inference of Neanderthal cognition features some inherent limitations and the pertinent literature shows itself to be rather indecisive, but some conclusions can nevertheless be reasonably well established. For once, inferring Neanderthals' cognitive inferiority from the fact that anatomically modern humans replaced them is overly simplistic in light of their shared history as well as the similar pattern featured by other replacements within both Neanderthal and anatomically modern human populations. Further, independently verifying this proposition of a cog- nitive inequality is, according to the current state of the interdisciplinary research, not sufficiently plausible. Instead, Neanderthal and anatomically modern human populations show comparable cultural and technological products up until the Neanderthal replacement and other dimensions, like relevant genetics and physiology, resemble each other to a significant extent too. Ecological and demographic factors seem to have played a major, if not the primary, role in the relationship between these early people. Overall, the current evidence suggests that Neanderthal mental lives were rich, probably not much unlike modern humans', and that the Neanderthal extinction is explainable without necessarily invoking a cognitive superiority on the side of anatomically modern humans. This does not mean, however, that Nean- derthals should be granted equality in every respect as if the species was indistinguishable from ours—which it of course was not. Any exact measure of Neanderthals' cognition lies beyond what is possible as of yet and so the specifics of the relationship between Neanderthals and anatomically modern humans as well as a possible impact of cognitive factors on the Neanderthal extinction will continue to be a valid point of contention.

Unfortunately, the degrees of freedom and the methodological difficulties of this research allows for plausible deni- ability whichever way the personal bias leans. Following this, the pertinent literature's biggest fault lies in its regularly one-sided way to handle the available evidence. Throughout the history of Neanderthal research, too many authors have tried to infer Neanderthal cognition, culture and language on the basis of single findings, which might come under scrutiny later in the light of new evidence, like with the instructive example of Solecki's flower burials. Similarly, too many more have tried to deny Neanderthals their humanity by contesting findings based on criticisms directed toward selected cases on the grounds that each one of them, taken on its own, cannot unequivocally demonstrate a modern- like Neanderthal cognition for one reason or another. Of course, it has to be acknowledged that almost none of the find- ings and interpretations are indisputable. This much is obvious from the nature of the inference itself, as has been made clear earlier in this overview. What indeed does show itself to be compelling is the emergent picture painted by the plethora of available data. Yes, conclusions may be contested on a case-by-case basis, yet the interdisciplinary aggregate research cannot be denied in its overwhelming entirety. In short, for those who clutch to the notion of Neanderthal inferiority, research on Neanderthal cognition, culture and language is a tale of not seeing the forest for the trees. A step back will yield a breath-taking view of a species that was not quite us, but nevertheless in many ways our ancestors' equals.

ACKNOWLEDGMENT This review was not written within a funded project. Employer of the author is the Ludwig-Maximilians-Universität München. Open access funding enabled and organized by Projekt DEAL.

CONFLICT OF INTEREST The author has declared no conflict of interest for this article.

ORCID Michael Breyl https://orcid.org/0000-0002-9736-9546

RELATED WIREs ARTICLES Cognition and art: the current interdisciplinary approach The thinking Neanderthals: What do we know about Neanderthal cognition? Coevolution of language and symbolic meaning: Co-opting meaning underlying the intial arts in early human culture The archeology of cognitive evolution

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How to cite this article: Breyl M. Triangulating Neanderthal cognition: A tale of not seeing the forest for the trees. WIREs Cogn Sci. 2021;12:e1545. https://doi.org/10.1002/wcs.1545

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