Updating Neanderthals: Understanding Behavioural Complexity in the Late Middle Palaeolithic

Updating Neanderthals: Understanding Behavioral Complexity in the Late Middle Paleolithic provides comprehensive knowledge on Neanderthals who lived throughout the European and Asian continents. The book synthesizes historical information about the study of Middle Paleolithic populations and presents current debates about their genetics, subsistence, technology, social and cognitive behaviors. It focuses on the last phase of Neanderthal settlements and presents the main patterns of modern humans across Europe. Written by international experts on the Middle Paleolithic who have conducted innovative studies in the last three decades, this book explores the implications of interactions between different human species, including Neanderthals, Denisovans and Sapiens.

In addition, the book discusses the diversity and variability of human adaptations and behaviors in the changing climate and environment of the Late Pleistocene, and the relationship between these behaviors, demography and cognitive capabilities.

• Offers a comprehensive update on the variability and diversity of Neanderthal behaviors during the Late Pleistocene
• Presents an interdisciplinary reconstruction of Neanderthals by assessing archaeology, paleontology, paleoecology, anthropology, genetics and cognition
• Reviews the reliability of archaeological data and the theoretical and methodological advances of the last 30 years
• Discusses the most debated Neanderthal themes, such as demography, diet, socio-economy and art

• Language: English
• Publisher: Academic Press
• Release date: Jun 28, 2022
• ISBN: 9780128214299

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Chapter 1: Updating Neanderthals: Taking stock of more than 160 years of studies

Francesca Romagnolia; Florent Rivalsb,c,d; Stefano Benazzie    a Departamento de Prehistoria y Arqueología, Facultad de Filosofía y Letras, Universidad Autónoma de Madrid, Madrid, Spain

b ICREA, Barcelona, Spain

c Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA), Tarragona, Spain

d Departament d’Història i Història de l’Art, Universitat Rovira i Virgili (URV), Tarragona, Spain

e Dipartimento di Beni Culturali, Università di Bologna, Ravenna, Italy

Abstract

This chapter aims is to briefly review the history of research on Neanderthals and to show how we have reached the current behavioural approach, including the complex scenario we have today. The main scientific debates that have marked Neanderthal studies since the discovery of skeletal remains at Feldhofer Cave in 1856 and the theoretical and methodological approaches that have progressively contributed to change and improve our image and understanding of this species are presented. Moreover, this chapter briefly introduces the main scientific issues and perspectives that have dominated the academic scene in the 21st century, which are further detailed in this book.

Keywords

Neanderthal; Middle Palaeolithic; Human evolution; Archaeological science; Cultural diversity; Material culture; Prehistory; Archaeology; Archaeological theory

Acknowledgements

Fr.R. research studies are supported by the Comunidad de Madrid and Universidad Autónoma de Madrid through the project SI1-PJI-2019-00488, which financed this work, and the Spanish Ministry of Science and Innovation through the projects ID2019-103987GB-C33 and PID2020-116732RB-I00.

Fl.R. research studies are supported by the Spanish Ministry of Science and Innovation through the project PID2019-103987GB-C31, the ‘María de Maeztu’ excellence accreditation (CEX2019-000945-M), and the Generalitat de Catalunya and AGAUR projects CLT009/18/00054, CLT009/18/00055, and 2017-SGR-836.

S.B. is supported by the European Research Council under the European Union’s Horizon 2020 research and innovation programme (grant agreement No 724046—SUCCESS) and the FARE programme 2018 (FARE Ricerca in Italia: Framework per l’attrazione e il rafforzamento delle eccellenze—MIUR).

"L’absence probable de toutes traces de préoccupations d’ordre esthétique ou d’ordre moral s’accorde bien avec l’aspect brutal de ce corps vigoureux et lourd, et cette tête osseuse, aux mâchoires robustes, où s’affirme encore la prédominance des fonctions purement végétatives ou bestiales sur les fonctions cérébrales"(Boule, 1911-1913).

Neanderthal—the first human fossil ever found—is the best pre-Sapiens hominid known today and the ancient species with the most extensive collection of bone remains. This human species occupied a vast area spanning Europe, the Near East, and extensive portions of Central Asia (Fig. 1.1), with variations in northern latitudes depending on ice covering due to Pleistocene climatic fluctuations, between approximately 400,000 and 40,000 years ago (Higham et al., 2014; Hublin et al., 2020; Meyer et al., 2016; Fig. 1.2).

Fig. 1.1

Fig. 1.1 Area of Neanderthal distribution.

Fig. 1.2

Fig. 1.2 Chronological span of Neanderthals and corresponding palaeoclimate curve.

Hundreds of caves, rock shelters, and open-air sites that were inhabited by Neanderthals have allowed us to yield many cultural remains and bio- and geological markers. Over the last decade, the application of different analytical methods to artefacts, bones, and sediments has opened novel perspectives in the understanding of technological and cultural strategies, subsistence and landscape dynamics, settlement patterns, and reconstruction of the environmental contexts in which these populations lived. These studies have changed our view of this human species and questioned our purported behavioural uniqueness.

Despite extensive data advances in laboratory analyses and techniques (Bocherens et al., 2022; Rivals et al., 2022), several aspects of Neanderthals’ day-to-day life and behaviours are still poorly understood. Furthermore, there is a long-standing juxtaposition in our approaches to Neanderthals and Sapiens. It is difficult for us to see ourselves as the result of a long and complex history. Our view is biased by the idea that our appearance in Africa at most probably 300 ka (Hublin et al., 2017) was disruptive and completely changed cognitive and behavioural patterns in human evolution. While most recent studies clearly state that it is increasingly difficult to detect specific and real differences between modern humans and Neanderthals, there remains a more or less conscious dichotomy between categories, such as ‘ancient/modern’, ‘simple/complex’ and ‘less/more’, which we use to portray the comparison between Sapiens and pre-Sapiens history. It has been emphasised that this should be related to our theoretical construction of ourselves as the only winners in the evolutionary process (Peeters and Zwart, 2020). Neanderthals, the closest fossil hominins to us from a chronological perspective, are somehow looked upon as what allows us to safeguard our own uniqueness and guarantee the philosophical distinction between us (‘humans’) and the others (‘non-humans’). It is not by chance that the label ‘Neanderthal’ is frequently used in the common language as a synonym for uncivilised and silly. The image of being brutish and unintelligent has characterised Neanderthals since the 19th century and endured, falling victim to political motivations and institutional settings as well as becoming part of our collective memory through illustrations and iconography in museums (Giacobini and Maureille, 2007; Moser, 1992; Sommer, 2006; Van Reybrouck, 2002).

1: Discovery, first studies, and the illustration of the dumb ape

The first Neanderthal remains were found in Germany in 1856 in Feldhofer Cave near Dusseldorf in the Rhineland. The name ‘Neanderthal’ comes from the name of this region, which was in turn derived from the family name of a 17th-century local composer: Joachim Neander (1650–1680). The meaning of Neander-thal, the Graeco-Roman translation of the original family name Neumann, literally means ‘the valley of the new man’. This name marked the Neanderthal’s paradox since the beginning. The first anatomical analysis of a skeleton highlighted the similarity of the skull’s shape with that of anthropoid great apes and some wild human groups, such as Negroes, Eskimos, and Australian Aborigines. According to the view of the time, these features were interpreted as belonging to individuals who were less advanced and less intelligent than Western modern humans (Drell, 2000; Madison, 2020). Schaaffhausen identified the bones as corresponding to a single skeleton and mentioned the possible antiquity of the remains dating back to the Ice Age, between the Miocene and the Quaternary periods (Schaaffhausen, 1858, 1888). Considered an archaic hominin, the remains fit perfectly with the prevailing notion of history based on the idea of a progressive and linear switch between different evolutionary stages, from savagery to barbarism and finally civilisation (Morgan, 1877).

Different studies published in the late 1800s and based on the anatomical traits of the remains found in Feldhofer Cave alternatively included this individual within the variability of modern humans yet in a subgroup of a primitive race thought of as either inferior to Western populations or within a savage, unknown, ancient human group (see Cohen, 2007). The first taxonomical definition was made by William King (1864), who appointed the remains the name Homo neanderthalensis. The Neanderthal individual was thus inserted into the genus Homo but excluded from our species. The first non-Sapiens hominin was defined. At that time, not all researchers agreed on the purported antiquity of these remains. Rudolf Virchow, a pathologist who was sceptical of the theories published by Darwin in 1859, proposed that the particular anatomical traits that were seen were attributable to diseases and malformations (Virchow, 1872). According to his interpretation, the Neanderthal was a modern human seriously affected by pathologies. This conclusion was based on comparisons with the skulls of ‘idiot’ individuals, as was customary at the time.

During the 60s and 70s of the 19th century, this debate created a great interest in the study of the human fossil remains that had been found in the first half of the 1800s in different locations (Stuttgart in 1700, Engis in 1829, and Gibraltar in 1847), which were now identified as being similar to the remains of the Neanderthal 1 individual at Feldhofer Cave. Furthermore, several novel fossils’ remains were yielded in the second half of the century, which increased the debate about Neanderthal traits and the evolutionary history of modern humans. In particular, two skeletons found in 1886 in Spy Cave in Belgium, in a well-defined geological context, allowed the first description of the direct association of human remains to a specific archaeological and palaeontological dataset, contributing to the acceptance of Neanderthals as an ancient hominin by the scientific community. The cultural data associated with Neanderthals were defined as ‘Mousterian’. This definition was given by Gabriel de Mortillet (1872) and inspired by the name of the Le Moustier in Dordogne—the site where they were found in 1860.

Thus, in the late 19th century, palaeoanthropological and archaeological studies significantly improved thanks to the systematisation of natural sciences and increasing systematic field investigations. These processes marked the early days of prehistoric archaeology as a recognised science. In the second half of the 19th century, the Stone Age studies developed worldwide (Nelson, 1937). Different ape-men were progressively identified and studied (Pithecanthropus erectus was identified in Java between 1891 and 1893), which unveiled different stages of human evolution from apes and provided an empirical basis for Darwin’s theory. In these studies, it was always clear, even when not explicit, that ape-men were inferior to us. A sign of this is the great emphasis systematically put on the Neanderthal skull rather than on postcranial skeletal remains. This was because, according to phrenology and physiognomy—common pseudosciences at that time that extrapolated personality traits from the shape of the skull and the face’s appearance—the cranium traits indicated a deviant and violent temper.

The most famous studies of the early 20th century (and the one on which the spread of the representation of the brutish and dumb Neanderthal lies upon the most) were conducted by palaeoanthropologist Marcellin Boule, who was then a professor at the Muséum National d’Histoire Naturelle in Paris. He analysed the almost complete skeleton of a male individual found at La Chapelle-aux-Saints, a cave in central France, and highlighted its physical similarities with apes, including the stooped posture that was considered to reflect a lack of intelligence (Boule, 1923). Regarding the non-human Neanderthal, a concept that has been raised since the first studies in this area, Boule emphasised the distance between Neanderthals and modern humans in temper (defining the former as being brute and violent) and cognitive capacities, stating that Neanderthals possessed a rudimentary psychic nature that was markedly inferior to any modern race (Boule, 1923, p. 237). His writings were complemented by the image of a hairy creature, hunched and ready to attack, as suggested by the creature’s hands holding a club and a boulder (Fig. 1.3). Boule’s study was a scientific, detailed analysis of the complete skeleton, and the scientific nature of the study as well as the prestige of the paleoanthropologist strongly affected the diffusion and consolidation of this image and interpretation (see Drell, 2000, pp. 10–12; Straus Jr and Cave, 1957, pp. 349–350). Therefore, the paradox of the ‘new man’ was definitively fixed.

Fig. 1.3

Fig. 1.3 The Neanderthal man ready to attack outside of La Chapelle-aux-Saints. (Drawing by Kupka published in L’Illustration (1909). Credit: agefotostock.com.)

2: The main scientific debates between 1930 and 1980: Anthropological and cultural data

During the first half of the 20th century, the academic debate continued, mainly focusing on evolutionary dynamics. Human evolution was thought of as a series of different human types that followed one another. However, it was not clear whether these types expressed different species that had appeared at different stages along the progressive evolution that resulted in us or were the expression of past racial diversity. The phylogenetic position of the Neanderthal was alternatively placed as an intermediate stage between Erectus and Sapiens or as a subspecies of Homo sapiens. The latter idea was expressed by Kleinschmidt (1938), who created the designation Homo sapiens neanderthalensis. This debate was fuelled by the intensification of fieldwork that increased palaeontological collections, showing a morphological variability between these individuals. Fossil remains identified as those of Neanderthals were progressively found in different locations in Eurasia. The widening of the geographical area that had been occupied by this population continued, and by the middle of the 1900s, it included the debate on intraspecific variability, nuclear areas, local evolution dynamics, and adaptations (e.g. Howell, 1957; Thoma, 1965). These were the years in which a shift in the interpretation of the Neanderthal is identifiable. Anatomical traits were no longer considered a reflection of an intellectual, primitive, and racial inferiority but were rather either the result of phenotypic responses to climatic conditions or caused by pathologies. Regarding the latter, it is notable that the palaeoanthropological evidence of La Chapelle-aux-Saints (more than others) contributed to the spread of the image of a brutish and dumb individual due to stooped posture, yet it indirectly led to the rehabilitation of the Neanderthal’s reputation. Novel studies suggested that the individual found in this cave in France was affected by several pathologies, including severe arthritis that caused deformities. He was standing and walking like modern humans affected by spinal osteoarthritis (Straus Jr and Cave, 1957, pp. 358–359). During these same years, other authors reached the same conclusions about the postural significance of Neanderthal vertebral morphology (see Straus Jr and Cave, 1957, pp. 360–362). Even if it was not immediately discussed and investigated at the time, this statement has relevant behavioural and cognitive implications: it indicates that other individuals within the group must have helped care for this man, which is at odds with the idea of a brutish and violent creature.

Since the end of the 19th century, publications about the Neanderthal often identified him as the ‘Mousterian man’, following the naming of his stone tool assemblages. Since the beginning of the 1900s, cultural classifications were based on the typological features of stone tools, which were determined through the study of the shape of the artefacts and the specific characters of the retouch, including position, delineation, morphology, etc. (Inizan et al., 1999). This approach was based on the ‘index fossil’ concept, which was borrowed from the natural sciences. Each cultural context was characterised by a specific tool or a specific set of tools, which were clearly identifiable. The presence of such elements in a stratigraphic layer guaranteed the direct attribution of a specific cultural context, and it was directly linked to a human species (Breuil, 1909). This approach, which required blind faith, allowed the comparison between sites and assemblages that ultimately created macro-regions in which the same cultural definition was applied and hence the same population dynamics.

Increasing fieldwork activities led to the growth of archaeological collections. Cultural materials associated with Neanderthal fossil remains were progressively collected across Eurasia and, in addition to anthropological traits, showed an internal variability. For example, Dorothy Garrod (Garrod and Bate, 1942) mentioned the presence of a ‘Levallois-Mousterian’ in the Levant assemblages according to the identification of Levallois knapping strategies for stone tools. In the first half of the 19th century, this variability was alternatively interpreted as the result of either chronologically successive technological stages (following a unilineal and stadial approach, such as in human evolution; Peyrony, 1930) or the existence of two lineal phyla. The latter model assumed the existence of a group of industries with bifacial tools evolving in parallel to another group without them, which was in turn subdivided into groups with and without Levallois elements (Breuil, 1929). From the mid-20th century onwards, the archaeological debate mainly focused on characterising and interpreting this techno-typological variability within the cultural category of the Mousterian. The debate persisted throughout the last century and is still current, albeit with some different perspectives. François Bordes systematised this variability in four Mousterian groups, which were differentiated according to the different ratio in the presence of specific retouched tools and a few technological traits in the lithic industries from sites in southwestern France (Bordes, 1953, 1961; Bordes and Bourgon, 1951; Table 1.1). The approach proposed by Bordes was the first attempt to systematise the definition of Mousterian’s typological variability through statistics and quantitative comparisons between the shapes of cumulative plots of tool types in each assemblage. Even if it still supported the ‘index fossil’ concept, Bordes’s classification (Bordes, 1953, 1961) was based on a larger selection of pieces from each accumulation to improve the objectivity of the analysis. This classification established a common vocabulary that is still used today and that constituted the basis for Neanderthal stone tool assemblages. However, technological traits were still viewed from a qualitative perspective, giving priority to the shape of retouched tools. Moreover, this chrono-cultural approach mainly focused on recording the presence and absence of diagnostic elements, while the typological approach limited the historical interpretation. Furthermore, the classification was based on a series of sites located in a limited geographical area and then applied to all of Eurasia without sufficient consideration for the possibility of differences determined by ecological adaptations and different technological knowledge outside western France (Dibble, 1991).

Table 1.1

Bordes (1950) suggested that the evolution of human technology was much more complex than previously thought and that it had had a bushy development (‘buissonnante’), which only further multidisciplinary studies—including typology, the study of ancient techniques, and a detailed stratigraphic analysis—could unravel. In Bordes’s view, the Mousterian types had cultural value in identifying different human groups with different technological traditions, each having been developed in limited areas over time.

The problem of how to interpret the Mousterian variability defined by Bordes soon generated a great debate. Binford and Binford (1966) suggested that the composition of stone tool assemblages yielded in a stratigraphic context was directly related to the human activities carried out at the site. The variability was not due to different tribes but rather to different social units performing different tasks. The idea was that each tool type had a different function (Binford, 1973), although progress in microscopic and use-wear analysis, experimental archaeology, and lithic technology disproved this theory (e.g. Anderson-Gerfaud, 1990; Beyries, 1987; Beyries and Boëda, 1983; Boëda and Bonilauri, 2006; Geneste and Plisson, 1996; Lemorini, 2000). In the late 1960s, Paul Mellars (1969, 1970, 1986) proposed that Mousterian variability was reflected technological evolution over time. Conversely to the idea of contemporaneity between facies expressed by Bordes and Binford, Mellars (1969, 1970, 1986) suggested a diachronic perspective. This model did not consider the interstratification of different Mousterian facies at the same site and their synchronic variability between different locations, as shown by further implementation of dating. That made the model unenforceable on a large scale (e.g. Jaubert, 1999; Jaubert et al., 2001; Turq et al., 1999).

Nicolas Rolland (1981, 1988, 1990) suggested that the variability reflected different modalities of landscape exploitation in terms of economy, mobility, raw material selection and use, and that this behavioural diversity was ascribable to the high adaptability of Neanderthal groups to a changing climate and environment. Harold Dibble (1984, 1988, 1995) proposed that typological variability could be attributable to the intense reduction of retouched tools. According to this theory, as the tool edge was worn out with use, the intensity and repetition of resharpening modified the final shape of the tool. Hence, the typological classes were not identifying different objects but rather different moments in the functional life of the same type of artefact. Dibble found that the understanding of Mousterian assemblages could not be reduced to a simple dichotomy between style and function and that other factors had to be considered. Looking for a dynamic perspective, Rolland and Dibble (1990) merged their ideas, proposing that the different stages of tool reduction were related to different accessibility to resources due to climate changes. From the 1980s onwards, many researchers emphasised the mobility strategies adopted by human groups in the territory in relation to the different sets of tools that they transported and to the different stages of the production sequence present at each location. Tool transportation and the fragmentation of technological activities in the landscape were affected by several factors, including the distance from raw material catchment areas, the abundance of abiotic resources and their specific constraints according to the knapping method, and the techniques that were applied (Geneste, 1985, 1989a, 1989b, 1991; Kuhn, 1992, 1995). Thus, it was necessary to look at the archaeological assemblages from a multidisciplinary and regional perspective, to integrate knowledge of the technological processes with the geological characterisation of the landscape and interpret human choices in raw material selection. Today, it is widely accepted that none of these interpretations alone is sufficient to understand Neanderthal behavioural variability.

3: The end of the 20th century

The taxonomic position of the Neanderthal in human evolution was debated throughout the 20th century. The debate advanced in the late 1990s thanks to paleogenetics. The first genetic analysis of ancient DNA was published in 1984 by Higuchi et al. (1984), with the molecular tissue being taken from an extinct member of the horse family. At the time, the combination of molecular biology and palaeontology was still considered futuristic. However, several improvements in molecular biology inspired ancient DNA research projects in the later decades of the last century (Hagelberg et al., 1991, 2015). The first studies of ancient DNA were focused on mitochondrial DNA (mtDNA), which is maternally inherited. Therefore, it is assumed that it does not recombine, and it is present in several copies in each organism located in the mitochondria—cellular organelles that produce chemical energy in most eukaryotic organisms (Cann et al., 1987; Kocher et al., 1989; Pääbo et al., 1988). Therefore, mtDNA is easier to study, and there is a higher probability of finding it, considering the degradation of DNA after the death of a living being. The first analysis of Neanderthal DNA showed that the mtDNA sequence of the individual found at Feldhofer Cave fell outside modern human variation (Krings et al., 1997). In this study, the researchers compared the mtDNA of a Neanderthal with sequences of present-day Homo sapiens. The comparison with mtDNA of ancient Homo sapiens confirmed that the Neanderthal was extraneous to our direct evolutionary history and that it went extinct without contributing to the mtDNA variation in Homo sapiens (Caramelli et al., 2003).

The confirmation by genetic analysis of Neanderthal’s segregation from Homo sapiens’ lineage offered the perfect framework in archaeology to reiterate our cultural and cognitive uniqueness. Thus, Upper Palaeolithic studies took a new approach. A common idea was that the Upper Palaeolithic transition was a major evolutionary event on a global scale, driven by Homo sapiens’ cultural and cognitive superiority, which marked the beginning of modern behaviour (Klein, 1995; Mellars and Gibson, 1996; White, 1982). It is significant that modern human origins were defined as a revolution that divided humans from all other primates (Bar-Yosef, 1998; Mellars and Stringer, 1989). The scientific debate was mainly focused on the final Mousterian assemblages, which are more or less contemporary to the entrance of Homo sapiens in Europe, and on the possible presence of innovations that were included in the list of modern behaviours (blade technology, microliths and weaponry, a wider range of exploited raw materials, hunting specialisation, a wider dietary spectrum, complex management of living space in campsites, and complex territorial mobility strategies). Emphasis was placed on evidence directly related to complex cognition, such as personal ornaments and the use of natural pigments. The attention to Middle/Upper Palaeolithic transition was justified by the purported contemporaneity between Sapiens and Neanderthals in time and space; thus, the origin of modern behaviour was discussed in terms of the capacity of acculturation of Neanderthal groups in the so-called ‘transitional industries’ (see d’Errico et al., 1998; Harrold, 2000; Pike-Tay et al., 1999; and bibliography therein). Some authors noted that the data suggested a more complex scenario and a gradual transition to modern behaviour. They also especially criticised the Eurocentric bias of the theoretical framework and the general preconceptions about the transition between archaic and modern humans, highlighting the more ancient evidence in Africa (Clark, 1997; McBrearty and Brooks, 2000; Straus, 1995).

4: The 21st century: Updating Neanderthals

The last 20 years have witnessed impressive advancements in the analysis of Neanderthal fossil remains by exploiting digital methods, dental histology coupled with chemical/physical methods and biomolecular analysis. Volumetric image data obtained from X-ray microtomography (microCT) and, more recently, synchrotron radiation have allowed the virtual reconstruction of the external morphology and internal architecture of bone and teeth (e.g. Macchiarelli et al., 2006; Olejniczak et al., 2008). Improvements in three-dimensional modelling techniques, geometric morphometric methods, and functional biomechanics took advantage of the digital models to improve our knowledge of unknowns, such as Neanderthal skeletal morphology (e.g. Gómez-Olivencia et al., 2018), taxonomy (e.g. Romandini et al., 2020), growth trajectory (e.g. Gunz et al., 2010), development patterns (e.g. Bayle et al., 2009), function (e.g. Wroe et al., 2018), and diet (e.g. Fiorenza et al., 2015, 2020). Moreover, histological analyses of Neanderthal teeth were used to determine their pace of growth (e.g. Nava et al., 2020), and the use of laser ablation analysis on tooth enamel, even on thin dental sections, enabled high-resolution mapping that exposed the internal differences of trace elements and isotopic ratios to improve our understanding of Neanderthal’s diet and mobility (e.g. Austin et al., 2013; Nava et al., 2020; Smith et al., 2018).

Undoubtedly, one of the most striking achievements of the last two decades was the extraction of endogenous DNA from Palaeolithic human groups, including Neanderthals, not only directly from fossil remains (e.g. Green et al., 2010; Hajdinjak et al., 2018; Prüfer et al., 2014) but also from sediment samples (Slon et al., 2017; Zhang et al., 2020). Palaeogenetic studies have confirmed the biological distance between Neanderthals and Sapiens, and the molecular clock has placed the time of their divergence from a common ancestor around 700,000–600,000 years ago (e.g. Prüfer et al., 2017). Moreover, an ancient genome unveiled the existence of a hominin species—Denisovans, a sister group of Neanderthals—which had not been identified by fossil remains alone (Krause et al., 2010; Meyer et al., 2012; Reich et al., 2010; Viola, 2022). Interestingly, the nuclear genome suggested interbreeding between Neanderthals, Denisovans, and Sapiens (e.g. Fu et al., 2015; Green et al., 2010; Kuhlwilm et al., 2016; Slon et al., 2018), and a small percentage of their DNA currently survives in modern humans (e.g. Massilani et al., 2020; Prüfer et al., 2017; Reich et al., 2011), which could potentially affect human health (e.g. Dannemann et al., 2016; Quach et al., 2016; Racimo et al., 2015; Simonti et al., 2016). More recently, ancient proteins (i.e. palaeoproteomics) were used either to support palaeogenetic studies or to overcome some of the limits of ancient DNA because proteins survive longer. For example, the collagen protein COL1 was used to identify human genetic fingerprints from morphologically undiagnostic bone fragments, which were then analysed for ancient DNA to confirm their Neanderthal affinity (e.g. Welker et al., 2016). In other circumstances, ancient proteins were used directly for taxonomic discrimination (Chen et al., 2019).

The reconstruction of the palaeoenvironmental and palaeoecological contexts of the Neanderthal groups (Sánchez Goñi, 2022) and their mobility and land use patterns is now well established as a multipronged and transdisciplinary task. In recent years, the advantages of combining approaches from different fields of research that were previously employed independently have emerged. This strategy has allowed improved palaeoenvironmental reconstructions and inferences on Neanderthal subsistence strategies, mobility, and seasonality. The combination of evidence gathered from different disciplines and high-resolution techniques allows the identification of the territories exploited by Neanderthals and the strategies adopted in doing so. Such studies involve the combination of two or more of the following disciplines and techniques: zooarchaeology sensu stricto, lithic technology and the analysis of raw materials, stable isotopes, cementum, dental calculus, dental micro- and mesowear, and macro- and microbotanical remains (pollens, phytoliths, and other vegetal remains). Neanderthals had complex mobility and land-use patterns that were adapted to changing climates and diversified landscapes (e.g. Delagnes and Rendu, 2011; Moncel et al., 2019; Sánchez-Hernández et al., 2019). Neanderthal mobility patterns are mostly related to the provisioning of abiotic resources (mainly raw materials) and to the seasonal availability of biotic resources (both vegetal and animal; Blasco et al., 2022; Hardy, 2022; Rendu, 2022). Mobility strategies were found to be highly diverse and to vary regionally depending on the characteristics of each area (e.g. Delagnes and Rendu, 2011; Turq et al., 2017). In terms of subsistence strategies, zooarchaeology and taphonomy allowed exploring prey selection, transportation strategies, seasonality of occupations, carcass processing, food exchange, or group cooperation among Neanderthals. The significant role of large- and medium-sized ungulates in the Neanderthal diet has been largely accepted, but recent studies have reported a highly diverse diet that also included other food items (plants, small mammals, and aquatic resources) and high dietary flexibility, with significant differences across the Neanderthal world (e.g., Fiorenza et al., 2015, 2020).

The improvement of combined technological and geological analysis, and Geographical Information System (GIS) applications, allowed the reconstruction of the supply areas of abiotic resources. Although Neanderthals were mainly collecting resources in a local range (less than 10 km far from the archaeological site), the evidence of mobility in a regional range (within 30 km of the site) is quite substantial (e.g., Browne and Wilson, 2011; Gómez de Soler et al., 2020; Villeneuve et al., 2019), and data showing the collection of raw materials located further afield at more than 50 km and up to 200 km also suggested complex mobility strategies and an interconnection between different territories and human groups (see Eixea et al., 2020; Minet et al., 2021; Neruda, 2012; Romagnoli et al., 2016a; Turq et al., 2017; and the bibliography therein). These studies suggest that Neanderthals knew the landscape well, selected resources according to different criteria, including not only availability and good quality (i.e. a homogeneous texture for good control of knapping fracture) but also social and cultural matters, and adapted their knapping methods to raw material constraints. Furthermore, Neanderthals accurately selected their personal gear—a kit of tools and cores that were transported by individuals—to cope with possible needs during their daily and seasonal movements and all planned tasks for the different locations and camps where they lived and to dispose of raw material on hand (e.g., Martín-Viveros et al., 2020; Turq et al., 2013).

Neanderthals created a diversified and versatile stone tool technology that goes beyond the original Mousterian definition based on stone tool assemblages in Western Europe and include diversified bifacial industries in Central and Eastern Europe and large portions of Asian regions (Romagnoli et al., 2022). Furthermore, micro-productions, traditionally related only to modern humans, have recently been identified as a relevant component of Neanderthal technology (see Mathias and Bourguignon, 2020; bibliography therein). Neanderthals also used wood (Aranguren et al., 2018; Carbonell and Castro-Curel, 1992; Rios-Garaizar et al., 2018), shells (Romagnoli, 2018; Romagnoli et al., 2016b), and animal bones and teeth to manufacture their tools (Blasco et al., 2013; Burke and d’Errico, 2008; Hutson et al., 2018; Majkić et al., 2017; Soressi et al., 2013).

The behavioural complexity of Neanderthals has been shown not only in landscape mobility but also in the organisation of living space within their campsites (Vaquero, 2022). Activity areas were carefully organised around fireplaces (Vaquero and Pasto, 2001), diversified according to the tasks that were realised (Spagnolo et al., 2019; Vallverdú et al., 2010), and functionally and socially interconnected (Vaquero et al., 2019). Neanderthals maintained a collective memory of the use of living spaces across time and simultaneously diversified the intra-site activities and their locations according to their mobility strategies, duration of occupation at the site, and seasonal movements (Marín et al., 2019; Zilio et al., 2021).

An issue that has recently increased the scientific debate concerns the cognitive complexity of Neanderthals as expressed by symbolic-mediated elements (García-Diez, 2022; Jaubert et al., 2022). Within this body of evidence, several authors include the extraction of bird feathers (e.g. Blasco et al., 2016), the exploitation of raptors’ talons for non-utilitarian purposes (Rodríguez-Hidalgo et al., 2019; Romandini et al., 2014), the use of shells, often associated with colouring substances, and the use of natural pigments (Peresani et al., 2013; Zilhão et al., 2010). This last category of evidence also includes the dating of carbonate crusts overlying schematic rock art at three sites in Spain (Hoffmann et al., 2018a) that recently generated a heated debate (see Hoffmann et al., 2018b, 2020 and the bibliography therein). Furthermore, Neanderthals are associated with burials and the evidence of post-mortem body treatment that could be related to a novel perception of individuals as part of the social group. Complex social dynamics can also be seen in the increased survival of pathological individuals that was revealed in the fossil record, suggesting social care (Trinkaus, 2018).

5: From brutality to rehabilitation and narration

Middle Palaeolithic research in the 21th century has shown the behavioural and cognitive complexity of Neanderthal groups, their great capacity for adaptation and resilience across Eurasia, and their complex genetic history. However, we are still unable to interpret these data from an evolutionary perspective, to understand their demise, and several related topics are still strongly debated. Over the last two decades, the concept of ‘behavioural modernity’ has progressively lost support (see Roberts, 2016; Zilhão, 2018, and the bibliography therein). The most recent studies clearly show a composite mosaic of ‘modern’ characters that appeared in different locations at different times as the result of local complex dynamics, which are still unknown in terms of adding many details. Recent data show that Neanderthals and modern humans were culturally and cognitively closer than previously thought (Fig. 1.4). However, our view of this species is still biased by our evolutionary ego and our dualistic view of ‘us’ opposed to ‘others’, with a qualitative perception in which, obviously, the best qualities have to be assigned to us. From this perspective, it is significant that the word ‘Neanderthal’ is often associated with ‘rehabilitation’ (Françozo and Soressi, 2018), indicating the effort to normalise the perception of someone after a long period of portrayed inferiority or impairment.

Fig. 1.4

Fig. 1.4 The data generated in the last two decades clearly show that Neanderthals were culturally and cognitively close to us and a dualistic way of thinking by noting that juxtaposing humans versus non-humans is no longer acceptable. (Drawing created by Alessandro Bartoletti, Cooperativa Archeologica ARA and based on an idea by Prasildo Brilli.)

Studies of Neanderthals in the last 20 years have probably changed our knowledge of the species more than that of any other ancient population. This is because we have become increasingly more aware of the processes that affected their sites and assemblage formation; consequently, the cultural inferences drawn from archaeological data are now more accurate (Romagnoli et al., 2018). We have refined the chronological framework, especially for the Late Middle Palaeolithic prior to Neanderthals’ demise (Higham et al., 2014). We have systematised multidisciplinary research, developed archaeological science, and strongly improved laboratory analyses and techniques, opening novel perspectives in several domains and paving the way for research questions that were unforeseen 30 years ago. We are able to investigate growth rates, early-life metabolic constraints and language capacities (Dediu and Levinson, 2018; Nava et al., 2020), and mating behaviours (Lalueza-Fox et al., 2011). Neanderthals’ strengths were in repetitive movements, an effective power grip in precision tasks (Faivre et al., 2014; Karakostis et al., 2018), diet diversity (e.g. Blasco et al., 2010, 2016; Marín et al., 2017; Zilhão et al., 2020), knowledge and use of medical plants (Hardy, 2018), and symbolic thinking (e.g. Hoffmann et al., 2018a; Jaubert et al., 2016; Majkić et al., 2017; Peresani et al., 2011; Zilhão et al., 2010). We are now ready to move on from the rehabilitation phase to the narration of Neanderthal lifestyle, knowledge, and capacities to depict the diversity of Neanderthal adaptations in their world and test the evolutionary implications on both local and larger scales. We need to consider the overall plethora of published case studies, including the different perspectives and theoretical frameworks. The collective narrative in this book is a pathway to tentatively review where we are and what has changed about our understanding of Neanderthals while considering the methodological constraints that still limit comparisons in space and time and lay the groundwork for the next steps in Neanderthal studies. It could be a pathway to definitively overcome the paradox of the ‘new man’.

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