Neandertal Demise: An Archaeological Analysis of the

Modern Human Superiority Complex

Paola Villa1,2,3*, Wil Roebroeks4
1 University of Colorado Museum, Boulder, Colorado, United States of America, 2 Unité Mixte de Recherche 5199, De la Préhistoire à l’Actuel, Culture, Environnement et
Anthropologie (PACEA), Université Bordeaux 1, Talence, France, 3 School of Geography and Environmental Studies, University of the Witwatersrand, Johannesburg, South
Africa, 4 Faculty of Archaeology, Leiden University, Leiden, The Netherlands

Abstract
Neandertals are the best-studied of all extinct hominins, with a rich fossil record sampling hundreds of individuals, roughly
dating from between 350,000 and 40,000 years ago. Their distinct fossil remains have been retrieved from Portugal in the
west to the Altai area in central Asia in the east and from below the waters of the North Sea in the north to a series of caves
in Israel in the south. Having thrived in Eurasia for more than 300,000 years, Neandertals vanished from the record around
40,000 years ago, when modern humans entered Europe. Modern humans are usually seen as superior in a wide range of
domains, including weaponry and subsistence strategies, which would have led to the demise of Neandertals. This
systematic review of the archaeological records of Neandertals and their modern human contemporaries finds no support
for such interpretations, as the Neandertal archaeological record is not different enough to explain the demise in terms of
inferiority in archaeologically visible domains. Instead, current genetic data suggest that complex processes of interbreeding
and assimilation may have been responsible for the disappearance of the specific Neandertal morphology from the fossil
record.

Citation: Villa P, Roebroeks W (2014) Neandertal Demise: An Archaeological Analysis of the Modern Human Superiority Complex. PLoS ONE 9(4): e96424. doi:10.
1371/journal.pone.0096424
Editor: Michael D. Petraglia, University of Oxford, United Kingdom
Received February 26, 2014; Accepted April 7, 2014; Published April 30, 2014
Copyright: ß 2014 Villa, Roebroeks. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: Research by P.V. was funded by the National Science Foundation grant BCS 1118143. Both P.V. and W.R. were supported by the Netherlands
Organization for Scientific Research (N.W.O, SP128-548). The funders had no role in study design, data collection and analysis, decision to publish, or preparation
of the manuscript.
Competing Interests: The authors have declared that no competing interest exist.
* E-mail: [email protected]

Introduction in Ethiopia suggesting that the early modern human morphology

emerged in East Africa possibly as early as 195,000 year ago [8–
The demise of Neandertals is one of the most debated issues in 10]. There is now general agreement that modern humans
paleoanthropology. Their disappearance in the fossil record originated in Africa, and subsequently expanded their range into
constitutes the biological part of a process of change that occurred the Near East and later into Europe. This is the core of the so-
in Europe and in the Near East between approximately 45 and 35 called Out-of-Africa hypothesis [11].
thousand years ago (ka) [1–2]. In western Eurasia, the process led In tandem with these developments, archaeologists began
to the replacement of an archaic population (Neandertals) with looking for modern behavioral markers in African sites dated
Middle Paleolithic technologies by a population of modern between 200,000 and 60,000 years ago. Many (see below) would
humans (Homo sapiens) with Upper Paleolithic ones [3–5]. The now suggest that there is indeed evidence for significant behavioral
study of this process of transition integrates data and scientists and cognitive differences between Neandertals and their African
from a wide range of disciplines, including archaeologists, physical contemporaries, and that when early moderns encountered
anthropologists, dating specialists, and increasingly so, geneticists. Neandertals in Western Eurasia, these differences would have
Into the 1980’s many paleoanthropologists argued that the entailed the demise of the Neandertals.
Neandertals had evolved into modern humans (or modern
Europeans) and that the Upper Paleolithic derived from the
Hypotheses for the Demise of Neandertals
Middle Paleolithic Neandertal culture. The opposite view assumed
Virtually all explanations for the disappearance of the
a single origin of modern humans and replacement of archaic
Neandertals from the Eurasian record point in one way or
populations, including Neandertals, by modern humans immi-
another to the arrival of Homo sapiens, anatomically modern
grating from an unknown source area [6]. This view became
humans (AMH), in Europe and western Asia. Late Pleistocene
widely accepted with advances in genetic studies and dating of
dispersal events brought AMH into the ranges of other hominin
fossils and sites in Africa, Europe and the Near East. In 1987 the
populations outside of Africa. In recent years we have seen a series
work of Cann and colleagues [7] provided compelling mitochon-
of publications with detailed maps purported to show the progress
drial evidence for a recent African origin of all modern humans.
of modern humans and their new technology across various
Later, the genetic evidence was supported by fossils which showed
Eurasian landscapes populated by ‘‘archaic’’ hominins, including
that Africans were far more modern looking than their Neandertal
Neandertals [12–15]. The source populations [16–19] and the
contemporaries, with dates for the Omo Kibish 1 and Herto skulls

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 An Archaeological Analysis of the Neandertal Demise

routes supposedly taken by the advancing modern humans vary Non-archaeological data have also been called upon to explain
widely in these papers though, as do the chronologies for the the outcompeting of the large-bodied and big-brained Neandertals
supposed Out of Africa dispersal(s) of modern humans. Some by modern humans, but these fall in first instance out of the scope
archaeological estimates hypothesize an age of around 125 ka for of this review (but see Discussion). The goal of this paper is to test
the first AMH dispersals into the Arabian peninsula [20] and the strength of the archaeology-derived hypotheses for Neandertal
around 77 ka for India [21], while others suggest that AMH extinction referred to above. While some of these hypotheses have
dispersal, thought to be associated with distinctively African been evaluated individually [47], ours is the first systematic study
technologies analogous to the ‘‘Howiesons Poort’’, occurred only of a wide range explanations. It is timely too, given the large
around 50–60 ka, i.e. after the Toba volcanic eruption at around amount of new data generated by fieldwork in Africa, the resulting
75 ka [22]. Genetic dates for the Out-of-Africa dispersal(s) of speculations on modern humans cognitive modernity [28,30], and
AMH also vary widely, between approximately 45 to 130 ka [23– new insights into Neandertal behavior and biology, including their
25]. biological affinity with modern humans. Genetic studies now
The disappearance of the archaic populations, including suggest that the debate on the demise of the Neandertals needs to
Neandertals, is routinely explained in terms of the ‘‘superiority’’ be reframed in terms of some degree of interbreeding [23,48,49].
of modern humans, who had developed in Africa the ability to In that sense, Neandertals did not go extinct, even though their
evolve complex cultural traditions and had become equipped with distinctive morphology did disappear. We will return to this topic
cognitive capacities which allowed them to expand globally and at the end of this paper.
replace all other hominins [26,27]. Such interpretations have
increasingly become based on proxies in the Middle Stone Age Methods
(MSA) archaeological record of sub-Saharan Africa which,
compared to the Middle Paleolithic record of Europe and western Our evaluation of the key archaeology-derived explanations for
Asia, would testify to superiority in a wide range of domains, either the demise of the Neandertals entails a comparative study of the
in Africa and/or upon arrival of Homo sapiens in the Neandertal archaeological record of Neandertals and contemporary modern
geographical ranges. These include inventiveness and capacity for humans, i.e. AMH in Africa and Southwest Asia between 200 and
innovation [11,28], complex symbolic and linguistic abilities 40 ka. To include younger periods would disregard the effects of
[29,30], more efficient hunting strategies [31], exploitation of a cultural and technological evolution after the demise of the
broader range of resources including plants and aquatic ones [32], Neandertals. The various competing models regarding the
projectile technology [33–35], heat treatment of lithic raw evolutionary disadvantages of Neandertals are listed in Table 1
materials [36], hafting technology [37,38], planning capacities and are reviewed in detail in (Text S1 Hypotheses 1–11), where
including larger scale social networks as shown by large transport they are systematically described listing the specific hypothesis and
distances of raw materials [39], environmental flexibility [40], supporting as well as refuting evidence.
memory capacity [41] as well as larger population sizes [42].
Inferiority in one or more of these domains has been at the core of Transitional Industries
many explanations for the demise of the Neandertals. The so-called ‘‘transitional industries’’, which show some
Prior to the last decade the cultural attributes listed above were similarities to late Middle Paleolithic (Mousterian) industries but
generally considered as exclusive manifestations of the western also contain Upper Paleolithic forms and whose time range falls
Eurasian Upper Paleolithic, as the result of a major behavioral within a 45235 ka interval, will not be discussed in detail here, for
revolution compared to the preceding Middle Paleolithic. Seen the following reasons:
from a European or Near Eastern perspective, the Upper
Paleolithic witnessed the introduction of new technologies, the a) the makers of the Bohunician, Bachokirian, Szeletian and
ability to communicate symbolically, systematic use of body Streletskayan [in Central and Southeastern Europe and
ornaments and various forms of mobile and rock art, by modern Russia) are not known yet (late Neandertals or AMH?) and
humans expanding from Africa into Eurasia, leading to the hence their status is ambiguous [50].
gradual replacement of non-modern populations, such as the b) Neandertals are accepted by many–though not by all [51] - as
Neandertals [29,43–45]. It was acknowledged that some of those the makers of the Châtelperronian, best known from the
features had emerged earlier in Africa, but the most complex Grotte du Renne at Arcy-sur-Cure in France, excavated by
technologies and art forms were seen as characteristic of the Leroi-Gourhan and his team between 1949 and 1963 [52].
European Upper Paleolithic, thus clouding the issue of the source The interpretation of the industry, rich in distinctively
area where these innovations had taken place. ‘‘modern’’ cultural features such as ornaments and bone
In 2000 McBrearty and Brooks [27] forcefully argued that the tools, has been the subject of heated debates. The
components of this ‘‘Upper Paleolithic revolution’’ were already controversies about whether the ornaments and bone tools
visible in the African MSA, tens of thousands of year earlier. They were (i) an invention of Neandertals [46,53], (ii) the result of
suggested a gradual assembling of a package of modern human stratigraphic admixture of Neandertal remains and Upper
behavior in Africa, which was later exported to other regions of the Paleolithic artifacts [54–57], or (iii) due to acculturation
Old World: a view contested by Klein [11], who stressed a later [28,58,59] have been going on since the acculturation
and punctuated emergence of ‘‘modern human behavior’’. In hypothesis was most explicitly discussed in 1998 by D’Errico
2003 D’Errico [46] reviewed the cultural attributes which et al. [60]. The stratigraphic integrity of the Châtelperronian
McBrearty and Brooks saw as defining modernity. He argued layers at the site has been reaffirmed in a recent paper [61],
that comparable traits also occur in the Neandertal record and contra [62]. New radiocarbon dates of 44,970244,520 cal
rejected the theory that behavioral ‘‘modernity’’ indicators are BP for the start and 41,300240,570 cal BP for the end of the
uniquely associated with Homo sapiens. Nevertheless, the behavioral Châtelperronian at Arcy and of 41,950240,660 cal BP (all
markers described by McBrearty and Brooks have in recent years dates with probability at 68.2%) for the Saint Césaire
increasingly been used to explain the demise of the Neandertals Neandertal suggest that the makers of the Châtelperronian
when modern humans expanded into their territories. ornaments were indeed Neandertals [63]. However, the

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 An Archaeological Analysis of the Neandertal Demise

Table 1. Hypotheses for the demise of Neandertals (a).

1. AMH had ‘‘complex symbolic communication systems’’ and ‘‘fully syntactic language’’, while Neandertals did not.
2 Neandertals had limited capacity for innovations.
3. Neandertals were less efficient hunters.
4. Neandertal weaponry was inferior to AMH projectile technology.
5. Neandertals had a narrow diet, unsuccessful in competition with AMH with their more diverse diets.
6. The use of traps and snares to capture animals was the exclusive domain of AMH.
7. AMH had larger social networks.
8. The initial AMH populations entering Neandertal territory were significantly larger than regional Neandertal populations.
9. Hafting by AMH required complex procedures indicative of modern cognition, while Neandertals hafting was a simple procedure using naturally available glues.
10. Cold climate around 40 ka was a factor in Neandertal decline.
11. Eruption of Mount Toba volcano at 75 ka played an indirect role in Neandertal extinction.

(a) See Text S1 Hypotheses 1–11 for details.

doi:10.1371/journal.pone.0096424.t001

conflicting hypotheses of acculturation versus independent technological and in some cases fossil) data before we can make
invention persist, as the dates appear to postdate or overlap in accurate assessments of the evidence, e.g. in terms of the type of
time with the arrival of early modern humans in Italy [5] and hominin authorship [62]. In the case of the Châtelperronian,
with dates for the Aurignacian in Germany [63; contra 2]. attributed to Neandertals by several scholars, we will review recent
evidence from sites where stratigraphic admixture can be excluded
Early descriptions of Châtelperronian assemblages stressed a and are less controversial than Grotte du Renne.
Mousterian component but the industry is now considered Upper
Paleolithic in technology, although different from the Aurignacian,
Results
and the presence of Mousterian tools as due to syn-or post-
depositional mixing [51]. The Châtelperronian lithic industry Explanations for the demise of Neandertals have been
recently studied at the open-air site Canaule II in France [64] is developed at various levels of abstraction, and include topics
also described as fully Upper Paleolithic, based on its technology notoriously difficult to study in the archaeological record, such as
and almost exclusive production of blades and backed points. In ‘‘complex symbolic communication systems’’ [28], ‘‘fully syntactic
contrast to the Arcy site, the very large assemblage of Canaule II language’’ [67] or ‘‘cognitive capacities’’ in general. Other
comes from a thin and unique layer, with its integrity and hypotheses refer to behavioral domains which do leave clear
homogeneity confirmed by refitting. The absence of any Middle traces in the archaeological record, provided the right taphonomic
Paleolithic elements in this Châtelperronian assemblage again conditions prevail (Table 1; Text S1, Hypotheses 1–11).
strongly suggests that the Châtelperronian, chronologically inter-
mediate between the Middle and the Upper Paleolithic, is a Language and Symbolism
unique entity, not the result of a mix of Middle and Upper The archaeological record has been mined in various ways to
Paleolithic artifacts. produce evidence for symbolic aspects of human culture, with a
strong focus on the emergence of language. Archaeological finds
c) The Uluzzian, an Italian transitional industry also present
in Greece and previously attributed to Neandertals [60,65], from the MSA have been used to build scenarios for the timing
is now seen by some as a product of modern humans, on and location of the origin(s) of language. Several of these finds
the basis of a study of two deciduous teeth from Grotta del come from South Africa and include engraved pieces of ochre
Cavallo in southern Italy [5]. AMS dates on shell beads from Blombos Cave [68–69], Nassarius shells from the same
from Grotta del Cavallo yielded 45.010243,380 cal BP for location [70], and heated silcrete artefacts from the site of Pinnacle
the lower Uluzzian layer. If the dates and the taxonomic Point, said to testify to sophisticated pyrotechnological know-how
attribution are accepted, they would extend the period of by early modern humans [36]. Botha has shown the assumptions
Neandertal-modern human coexistence to some millennia. and series of inferential steps some of these authors had to make
Neandertals are thought to have persisted in southern before being able to squeeze ‘‘language’’ out of their mute artefacts
Iberia until 37 ka, based on the dates for Middle Paleolithic [71–72], see also [73–74] pinpointing the weak spots in the steps
assemblages there [66] and at other sites in Europe based leading from observations about archaeological phenomena to
on dates for Neandertal remains at Spy (Belgium) and statements about the presence of ‘‘fully syntactical language’’.
Vindija (Croatia). Elsewhere the dates for the Campanian Moreover, recent data on Neandertal use of ochre and manganese
Ignimbrite ash horizon, stratigraphically above several as well as on Neandertal production of pitch, the presence of
Proto-Aurignacian layers, situate the end of the Middle transported and ochre-smeared shells, of ornaments such as eagle
Paleolithic at about 40 ka (see The date of the demise). claws and perhaps bird feathers [75–78] (Text S1, Hypothesis 1),
and the production of the specialized bone tools recently reported
The ‘‘transitional’’ industries are extremely relevant to under- from two late Middle Paleolithic sites [79] (Text S3, Lissoirs)
stand the routes of migrations and expansion of AMH in Europe, indicate no significant differences between the MSA data
the nature of cultural contacts between the local and immigrant commonly used to create these more abstract explanations and
populations and the onset of the Upper Paleolithic in those the later Middle Paleolithic record.
regions. However, we need more contextual (i.e. stratigraphic,

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 An Archaeological Analysis of the Neandertal Demise

Table 2. Dates of technological phases in the late Middle Paleolithic of Europe and in the late Middle Stone Age of South Africa (a).

Technological phases Start (ka) End (ka)

Still Bay
Blombos ca.75.5 (OSL) 67.8 (OSL)
Sibudu 70.562.0 (OSL)
Diepkloof 109610 (TL)
Howiesons Poort
Sibudu 64.762.3 (OSL) 61.762 (OSL)
Klasies River Main Site, Cave 1A 64.162.6 (OSL) 5663 (TL)
Border Cave 7464 (ESR) 6063 (ESR)
Diepkloof 105610 (TL) 52.565 (TL)
Post-Howiesons Poort/MSA III (b)
Klasies, Cave 1A 6065 (TL) –
Klasies, Cave 1A 57.965.3 (OSL) –
Border Cave 6063 (ESR) 44242 (14 C cal BP)
Sibudu 58.561.4 (OSL) 38.661.9 (OSL)
Boomplaas ca 5666 (U-series) 38236 (14 C cal BP)
Klein Kliphuis 57.862.4 (OSL) 33.361.3 (OSL)
Rose Cottage 56.062.3 (OSL) –
Mousterian in Western Europe (c)
Mousterian of Acheulian Tradition 70 40
(six sites in SW France)
Quina Mousterian 73 40
(six sites in SW France)
The Keilmessergruppen 80 50
(13 sites in Germany, Poland and the Czech Republic)

(a) After [89,91–92,94,96–105]. We have excluded assemblages with uncertain stratigraphy (Umhlatuzana, HP layers at Klein Kliphuis) or unpublished dates (Hollow Rock
Shelter).
(b) The term Post-Howiesons Poort is equivalent to MSA III at Klasies River Main Site. It includes informal designations of the Sibudu sequence such as late MSA and final
MSA. We have not included several TL and OSL dates for the HP and Post-HP of Rose Cottage because they are inconsistent or only informative for the middle part of
the sequence [105–106]. The Post-HP OSL date reported here for Rose Cottage [89] is of layer LIN which is toward the base of the Post-HP sequence but above its oldest
layer.
(c) The Middle Paleolithic technocomplexes are dated by TL, ESR, 14C (calibrated BP) and chronostratigraphy.
doi:10.1371/journal.pone.0096424.t002

The same applies to explanations regarding behavioral domains there is no solid archaeological evidence in its support (Text S1,
which do leave clear traces in the archaeological record. In our Hypothesis 4.3).
study none of the explanations listed in the introduction and in
Table 1 proved to be supported by adequate archaeological data. Organized Use of Space
The same applies to purported differences in the use of space at
Hunting Methods and Diet the level of camp sites by AMH and Neandertals, with organized
With the demise of the idea that Neandertals were scavengers use of space seen as typical for AMH. The South African MSA
and ineffective hunters [80–82], the former interpretive framework record has some cases of excellent preservation of plant materials
has to some degree been reformulated in terms of Neandertals in dry conditions, including possible bedding material recovered
inferiority in subsistence strategies and hunting weaponry for from 77 to 58 ka old deposits at Sibudu [85–86]. Some researchers
which, again, there is no support from the archaeological record have taken the presence of bedding material and ‘‘the deliberate
(Text SI, Hypotheses 3–4). Neandertals were by all means use and organization of living space’’ to be ‘‘an important trait of
accomplished large game hunters, who survived in a wide range culturally modern behavior’’ [87]. However, there exists good
of environments subsisting by hunting a wide range of animals in a evidence for well-delimited activity areas at Neandertal sites such
variety of topographical settings. In contrast to prevailing ideas as Kebara, Amud (Israel) and Tor Faraj (Jordan) as well as from
[31,83], their diet was not restricted to large and medium size several European sites where the task-specific areas are docu-
herbivores only. Several sites document a broader diet, including mented by refitting (Text S1, Hypothesis 3). Furthermore, bed
aquatic foods, small fast game (birds, rabbits) as well as plant building by great apes is a well-documented learned behavior,
resources (SI Hypothesis 5). Likewise, the idea that spear throwers dependent on appropriate early experiences [88].
and bow and arrow were first developed in the MSA of South
Africa before 60 ka and conferred substantive advantages on Capacity for Innovation
modern humans as they left Africa and encountered Neandertals Another prominent scenario suggests that the archaeological
equipped with only hand-cast spears [33,84] may be correct, but record of sub-Saharan modern humans, to wit of the two main

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 An Archaeological Analysis of the Neandertal Demise

technocomplexes of the South African late MSA, the Still Bay (SB) 70 ka in South Africa. Replication experiments suggest that HP
and the Howiesons Poort (HP), indicates very dynamic and hunters used a mixture of plant gum, beeswax and powdered
innovative phases which lasted less than 10,000 years each [26,89]. ochre to produce an adhesive that had to be carefully dried using
These would constitute a striking contrast to the record of the fire [37]. However, from 200,000 years ago onward, European
Neandertals, who supposedly lacked the capacities to innovate and Neandertals used fire to synthesize pitch from bark, through a
‘‘made the same kinds of tools for 200,000 years without ever process that involved distillation in the absence of oxygen and
tinkering with the basic components’’ [90]. Recently reported within a temperature interval of 340uC–400uC [108]. Pitch is not a
dates from the Diepkloof site (South Africa) are significantly naturally occurring glue; it is a man-made material produced using
complicating our views on cultural change in the Late Pleistocene fire as a tool. Birch park pitches have been experimentally
there, however. According to these new dates [91–92] the SB and produced in small dug out and subsequently covered pits beneath
HP technocomplexes would have a much longer duration than camp fires [109], though in very small quantities only, leaving
previously envisaged [89], comparable to those of broadly open the question how exactly Neandertals produced their pitches.
contemporaneous Middle Paleolithic industries in Europe, which Two flakes associated with elephant remains at the Italian site of
show clear spatio-temporal distributions (Table 2, Text S1 Campitello (Tuscany, Italy) were found enclosed in blackish
Hypothesis 2). Jacobs’ OSL age estimates for the SB and HP are organic material that was analyzed by gas chromatography/mass
considered controversial by some [93]. More dating work is clearly spectrometry and identified as a pitch obtained by a pyrolysis-type
required, while systematic technological and typological analyses process of birch bark for hafting the flint flakes [110–111]. The
are necessary to dispel doubts about assemblage definition, Campitello finds date to the end of MIS 7. Comparable finds of
especially for the MIS 5 occurrences. birch bark pitch come from the German site Königsaue A, with an
In contrast it is clear that the Post-HP technocomplex, estimated age of 80 ka [112–113]. On basis of the stratigraphy of
characterized by unifacial points on flakes (Sibudu) or blades the site, the AMS dates of 43,80062100 BP and 48,40063700 BP
(Border Cave, Klasies), hard hammer percussion, rare presence of cited in ref [113] should be considered minimum ages. Mania’s
the Levallois technique and of formal tools on blades (Klasies) and fieldwork at the site produced two pieces of pitch, one with
flakes (esp. Rose Cottage and Sibudu), has a duration of about fingerprints as well as the imprint of a stone tool and a wooden
20,000 years; even more if ‘‘transitional’’ or late MSA sites in haft. Experimental studies show that production of pitch in the
South Africa, dated between 40 and 20 ka, are taken into account. absence of air-tight pottery containers requires a high degree of
These include three layers at Rose Cottage, dated between ca 30.8 technical knowledge.
and 27 ka, and Strathalan Cave B, with two layers dated between According to Brown et al [36] heat treatment of silcrete at the
29 and 25.7 ka [94]. OSL and ESR dates for post-HP assemblages South African site of Pinnacle Point at c. 72 ka and possibly as
are supported by AMS radiocarbon dates. Thus the pace of early as 164 ka indicates sophisticated knowledge of fire and
change and the evolutionary patterns of the European Upper elevated cognitive abilities that may have been a behavioral
Pleistocene record, which shows regional differentiation, cultural advantage on Neandertals as early modern humans moved to
traditions and technological changes through time, are compara- Eurasia. The evidence of pitch production as early as 200 ka by
ble to what is known from the African record. Technological and European Neandertals shows that those ‘‘elevated cognitive
tool-type changes in the Mousterian industries precede by far the abilities’’ were not the exclusive domain of modern humans.
advent of Proto-Aurignacian and Aurignacian industries. What- The straightforward scenario of superior AMH moving into
ever dates are accepted for these industries [95], changes in Neandertal territory is also complicated by the Late Pleistocene
Mousterian industries occurred long before 50 ka. occupation history of the East Mediterranean Levant. AMH were
present in that region between 80 and 130 ka, and created the
Size of Social Networks Skuhl and Qafzeh record with its burials, pigments and personal
Other workers have suggested that Neandertals and AMH ornaments [114], associated with a Middle Paleolithic lithic
differed significantly in the sizes of their social networks. AMH technology. Between 80 and 47 ka however, only Neandertals are
larger-scale social networks are supposed to have acted as a buffer known from the fossil record of the Levant [115]. If the absence of
against environmental downturns, thus fostering long term fossil AMH in the record represents a true absence from the
survival. Such inferences are based on the translation of distances region, this could indicate that the Skuhl/Qafzeh hominins and
over which artifacts were transported in the deep past into their immediate descendants indeed may have ‘‘lacked the
statements about former mobility strategies, exchange systems and behavioral capacities that enabled subsequent modern humans
sizes of social networks. Yet it is almost impossible to differentiate to compete successfully against the Neanderthals’’ [115].
between long distance transport as a signature of direct
procurement as opposed to indirect acquisition, such as through
Discussion
trade or exchange networks [107]. Our review of the evidence
(Text S1, Hypothesis 7) shows that as far as the archaeological We conclude that all the ‘‘archaeology-based’’ explanations for
record for raw material transfer distances is concerned, the MSA the demise of the Neandertals reviewed here (Table 1, Text S1,
and the Middle Paleolithic record are not significantly different, Hypotheses 1–11) are flawed. They were based on much less data
despite of the obvious ecological differences between western than we have available today and were at least in part the result of
Eurasia and Africa. a long tradition of thinking in terms of Neandertals-AMH
dichotomies, steered by overstressing developments within the
Hafting Procedures, Heat Treatment and Cognition Upper Paleolithic of Europe, the record of which has become
According to another hypothesis Neandertals hafting of tools almost like a yardstick for modern human behavior (Text S2).
was a simple procedure, only using naturally available glues. Early While the debate about AMH dispersal times and routes out of
modern human hafting techniques entailed complex procedures Africa is intense, based on a range of archaeological as well as
which required ‘‘abstract reasoning’’ and are hence indicative of genetic data, the archaeological record from the various continents
modern cognition. According to Wynn and Coolidge [41] does not provide strong support for any of the suggested routes nor
evidence for complex hafting procedures dates back to about any of the suggested factors in the demise of the Neandertals. The

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very fact that the migration time estimates vary so widely suggests frequency, not cognition or technical competence, that distin-
that we simply have no solid data; perhaps there was more than guishes AMH bladelet production from that of Neandertals [120].
one migration event (in addition to the Last Interglacial limited The techniques and methods of bladelet making in the Mousterian
expansion in the Levant), and in all probability the migrating are different from those of the Protoaurignacian, just as the kind of
groups did not have a strong cultural homogeneity. This may possible symbolic objects are also different (use of raptor claws; on
explain why we do not see clear archaeological signatures for perforated or grooved animal teeth (see Text SI, Hypothesis 1).
AMH on the move. Perhaps the nature of the contacts should be seen in terms of
Interestingly, the widely accepted date of 60 or 50 ka for the diffusion of ideas rather than as face to face interaction and the
modern human expansion into Eurasia (following the earlier short- copying of specific objects [122]. The occurrence of Dufour
lived exodus in the Levant documented at Skuhl and Qafzeh) bladelets (often used as projectile elements in the Aurignacian and
would rule out South Africa as the location for source populations the Protoaurignacian) with very specific techniques of manufacture
for two reasons: (i) by 60 ka the HP tradition of backed tools made in the Châtelperronian of Quincay is interpreted in a similar way,
on blades and bladelets produced by soft stone hammer as a form of low-degree social interaction between Neandertals
(supposedly associated with the AMH expansion) had given way and modern humans [123].
to the post-HP assemblages characterized by a variety of flake tools
and blades produced by hard hammer percussion but without The Date of the Demise
backed blades [94,103,105,116]; (ii) the Still Bay and HP Various new dates support the idea of some chronological
populations were not larger than other MSA populations and overlap between AMH and Neandertals, which may have enabled
might even have been smaller, thus excluding population pressure interbreeding and cultural interaction in western Europe: AMS
as the prime mover of the migration [117]. According to Klein dates on ultrafiltered bone collagen from the Châtelperronian
[11] the Out of Africa expansion was underlain by a neural layers X and IX of Grotte du Renne at Arcy, c. 44 to 41 kyr cal
mutation that promoted the final development of the modern BP; the date of the Saint-Césaire Neandertal at 41.9240.6 kyr cal
human brain. Direct evidence for this hypothesis may come from BP [63]; the fact that the Protoaurignacian at the Italian sites of
comparisons of Neandertal and modern human genomes. Castelcivita and Serino is overlain by the Campanian Ignimbrite
In the recent past, much debate has been generated from the tephra, dated to 39.2860.11 ka by 40Ar/39Ar [124]; the modeled
observation that Neandertals began to produce a richer archae- age ranges of c. 41.5239.9 kyr cal BP of several radiocarbon-
ological record, including bone tools, personal ornaments and use dated Proto-Aurignacian sites [95]; the date of the Oase 2 early
of manganese and ochre, at the time when AMH started modern cranium at c. 40 ka [125]; the AMS dates for the
colonizing Europe. Some interpreted this change in the record Neandertal child from Spy cave (Belgium), 36,870 to 38,494 and
as the result of Neandertal absorption of ideas and techniques from 37,297 to 40,490 cal BP [126]; the AMS dates for the Vindija
the incoming AMH. After having produced a rather monotonous (Croatia) Neandertal remains at c. 38 kyr cal BP [127,128]; the
record for almost 300,000 years, an independent invention of these 37.4 ka cal BP date for the final Mousterian level of Cueva Antón
new items just at the time of the arrival of AMH would have to be in southeastern Spain [75]. Even if we do not consider dates
seen as an ‘‘impossible coincidence’’ [28]. However, as reviewed judged by some as controversial such as (i) the AMS dates on shell
here, use of ochre, of personal ornaments, production of beads for the layer containing the modern human teeth at Grotta
specialized bone tools and complex hafting techniques were part del Cavallo at 45.010243,380 cal BP [5, contra 2]; (ii) the dates
of the Neandertal repertoire already before the arrival of AMH in for the Kent’s Cavern modern human maxilla [3, contra 2, 129];
western Eurasia. and (iii) the dates for the Aurignacian at Geissenklösterle at c.
The present review also suggests that some of the innovative 42 kyr cal BP [4, contra 2], some millennia of overlap are
technologies of the Protoaurignacian and of the Aurignacian may indicated The latest Neandertal currently known from the Levant
have developed out of a Middle Paleolithic base (for a similar is the adult male skeleton from Amud Cave (Israel) with an ESR
viewpoint, see [118]). Some components that occur sporadically or date of 5368 ka on tooth enamel [130].
episodically in Neandertal and late MSA assemblages become
much more common later, like pigment use, symbolic objects, Interbreeding and Assimilation
extensive transport of raw materials and even specialized bone For some authors replacement and supposedly rapid extinction
tools [79]. The same goes for another element, the intentional of Neandertals can be explained only in terms of substantial
production of bladelets (,4 cm in length) from bladelet cores. cognitive, technological and demographic differences between the
Bladelets have been considered a discriminant factor between the Neandertals and AMH [42,131]. But, as we tried to show here, the
Upper and Middle Paleolithic and therefore between AMH and Neandertal archaeological record was not different enough to
Neandertals [119]. Production of bladelets has been securely explain their demise in terms of inferiority in archaeologically
identified in French Mousterian assemblages, e.g. at Combe visible domains. Thus, if Neandertals were not technologically and
Grenal (layers 30–29 and layers 16 and 14), Champ Grand and cognitively ‘‘disadvantaged’’, how can we explain that they did not
Grotte Mandrin, and in Spain at sites such as El Castillo and survive?
Cueva Morin [120,121]. All these assemblages belong to the final Some modern human-like anatomical characteristics are said to
Mousterian, with the exception of Combe Grenal and Grotte occur in late Neandertal fossils (as in the Vindja, St. Césaire and
Mandrin; at the latter site, a layer with blades, bladelets and Riparo Mezzena late Neandertals [132,133] and refs therein) and
microlithic points is overlain by five layers with flake-based vice versa some Neandertal features are present in early specimens
Mousterian assemblages [121]. At Combe Grenal layers 29–30 of modern humans in Europe [134,135] supporting a hypothesis of
have an estimated age of late MIS 4, i.e. around 60 ka. Bladelets some degree of admixture between the two groups. However, until
and bladelet cores are not abundant (5% of the assemblage at recently the morphological evidence of admixture was often
Combe Grenal layers 29–30), yet they show that Neandertals, like dismissed. In 2010 a draft sequence of the Neandertal nuclear
late MSA humans and the makers of the Protoaurignacian, DNA provided clear evidence of interbreeding between Neander-
mastered the technology of bladelet production, albeit using tals and modern humans [48], estimating that Neandertal
methods different from the HP small blade technology. It is their inheritance makes up 1–4% of the genomes of people outside of

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 An Archaeological Analysis of the Neandertal Demise

Africa. A revised estimate based on a high-coverage sequence of a resulted from a complex and protracted process [147] including
Neandertal from the Altai Mountains now suggests 1.5–2.1% [49]. multiple dynamic factors such as low population density,
Genes of Neandertals may have been favored through natural interbreeding with some cultural contact, possible male hybrid
selection, and possibly played a role in the development of the sterility and contraction in geographic distribution [148] followed
immune system of modern humans [136] or in UV-light by genetic swamping and assimilation by the increasing numbers
adaptations [137]. According to [138] gene flow from Neandertals of modern immigrants.
to modern humans occurred between 47,000 and 65,000 years
ago, and most likely happened at the time when Neandertals and Conclusion
modern humans encountered each other in Europe and the
Middle East around 50,000 years ago. In a review of the MSA and Middle Paleolithic archaeological
In sum, interbreeding and assimilation, the tenants of a model record we have shown that inferred markers of modern human
first proposed by Fred Smith [139] are now supported by genetic cognitive and behavioral capacities have a greater time depth in
data [134,140]. It can be argued that the level of interbreeding the Middle Palaeolithic record than commonly acknowledged. We
may have been too limited to support an assimilation scenario. An have found no data in support of the supposed technological,
interestingly parallel to this complex situation can be found in social and cognitive inferiority of Neandertals compared to their
another ‘‘revolution’’, the so-called Neolithic Revolution [43,141], AMH contemporaries. The results of our study imply that single-
which does not feature explanations in terms of ‘‘cognitive’’ factor explanations for the disappearance of the Neandertals are
differences. The first farmers swept into Europe from the Near not warranted any more, and that their demise was clearly more
East at about 7500y BP displacing the local Late Mesolithic complex than many archaeology-based scenarios of ‘‘cognitive
hunter-gatherers. But the Mesolithic hunter-gatherers, who cannot inferiority’’ reviewed here seem to suggest. This has implications
be described as cognitively inferior, were not submerged by hordes beyond the field of archaeology per se: archaeologists’ character-
of farmers. Farmers and foragers coexisted for thousands of years izations of Neandertals as cognitively inferior to modern humans
in NW Europe; in Central Europe local hunter-gatherers adopted [149] have created an interpretive framework within which subtle
farming but in southern Scandinavia local foragers retained the biological differences between Neandertals and modern humans
Mesolithic lifestyle for c. 1500 years after farming arrived in tend to be overinterpreted (see for instance [150].
Central Europe [142,143]. Cultural contact is suggested by clear After 40,000 years and 2000 generations the Neandertal
continuities in flint technology between the Mesolithic and early fraction in non-African modern human genomes still constitutes
Neolithic in the region. After a very complex pattern of expansions a substantial legacy from these ancient hominins who differed
and genetic shifts of the last 8,000 years the hunter-gatherer from contemporary AMHs in both geno- and phenotypes [151]
mitochondrial DNA haplogroups form 16% of the present-day but whose archeological record was not different enough to
Central European genetic composition [143]. It would take at least support the purported cognitive ‘‘gap’’ between them and their
one millennium between the first arrival of immigrants and a contemporary modern humans.
notable increase in their population size.
The original Neandertal contribution to modern human biology
Supporting Information
may have been larger 40,000 years ago - equivalent to 2000
generations (with generation time at 20 years) – than estimates Text S1 Hypotheses 1–11. Data on the various competing
based on genomic regions of present-day humans suggest models on the evolutionary disadvantages of Neandertals,
[144,145]. Interbreeding of Neandertals and modern humans presented as Hypotheses 1 to 11, systematically described listing
may have helped modern humans to adapt to non-African each specific hypothesis and supporting as well as refuting
environments but also introduced alleles that were not tolerated evidence.
and contributed to male hybrid sterility thus reducing the (DOC)
proportion of Neandertal ancestry of the period of contact to that
seen today [144]. Text S2 A single package of modern behavior?
Mitochondrial genetic diversity of eight early modern European (DOC)
humans dated to ca 38,000 to 4,500 (14C cal BP, from Kostienki Text S3 Lissoirs.
14 to Őtzi The Iceman) is 1.5 times higher than that of five (DOC)
European Neandertals spanning the time to 38 to 70 ka [23,146].
The high coverage genome of the Altai Neandertal [49] also Acknowledgments
suggests low genetic diversity which could indicate small popula-
tion sizes (see Text S1 Hypothesis 8 for archaeological data). These We thank three reviewers for their constructive comments. P.V. is grateful
genetic data suggest that differences in population sizes between to Payson Sheets, John Hoffecker, Douglas Bamforth and Gerardo
the ‘‘resident’’ Neandertals and incoming AMH populations may Gutierrez for useful suggestions. Marie Soressi and Alexander Verpoorte
gave valuable feedback on previous versions of the paper.
have been a contributing factor in the absorption of Neandertal
populations [23]. The momentous cultural changes that followed
the arrival of AMH in Western Eurasia were not uniquely due to Author Contributions
the residents’ cognitive or technological inferiority causing rapid Conceived and designed the experiments: PV WR. Analyzed the data: PV
and total replacement. The Neandertal demise appears to have WR. Wrote the paper: PV WR. Performed research: PV WR.

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 1

Neandertal Demise: An Archaeological Analysis of the Modern Human

Superiority Complex

Paola Villa, Wil Roebroeks

Supporting Information

Text S1. Hypotheses 1-11.

1. AMH had complex symbolic communication system and fully syntactic language
while Neandertals did not.
Hypothesis. In 1989 Paul Mellars [1] wrote a comprehensive review of facts and ideas,
current at the time, on links between modern behavior and the emergence of complex
forms of symbolism and language in the Upper Paleolithic. He noted the occurrence of
coloring materials such as ochre and manganese, some reportedly showing scraping or
facets due to use, at Mousterian sites in southwestern France [eleven Mousterian sites
with pieces of manganese or ochre were listed by Demars [2] for a total of 37 layers) and
the occasional specimens of transported fossil shells [3, see also 4]. However neither
pigments nor fossils were documented in any detail. Mellars also noted a few doubtful
examples of non-utilitarian objects such as perforated bones (i.e. “pendants”) and bones
and stones with supposed engravings. He emphasized the contrast between these rather
ambiguous cases of symbolic artifacts and the large numbers of clearly man-made
ornaments and figurative art documented since the early stages of the Aurignacian.
Aware of the limitations of the data, but aiming at a general explanation, he concluded
that evidence of clearly symbolic behavior amongst Neandertals was sparse, and that they
may have possessed some form of language although less complex than that documented
for anatomically modern humans. The emergence of complex language and other forms
of symbolic communications, in addition to more advanced technology, gave modern
humans a crucial adaptive advantage and led to their successful dispersal across Eurasia.
This remains a commonly accepted explanatory model for the Neandertal demise,
although the cause and tempo of this behavioral shift remains controversial [5-9].
Supporting evidence. Many of the controversial symbolic artifacts mentioned by Mellars
have now been shown to be natural. This is the case of the “engraved” rib from the MIS 6
layer of Pech de l’Azé II (France) of eleven bones with meandering marks from Cueva
Morín (Spain), of grooves on an elephant vertebra from Stránská Skála (Czech
Republic), and of perforated bone and phalanges, supposedly “pendants”, from Pech de
l’Azé II, Kulna (Czech Republic) Bois Roche (France) and Bocksteinschmiede
(Germany) [10] The Divje Babe flute has been shown to be carnivore damage [11].
In contrast, archeological finds from the South African record have been used as building
blocks for scenarios on the timing and location(s) of the origin(s) of language, suggesting
that the anatomically modern southern African populations had fully modern cognitive
abilities and syntactical language 75,000 years ago [12]. These finds include engraved
pieces of ochre from Blombos Cave, South Africa [13, 14], Nassarius shells from the
same location [12], and heated silcrete artefacts thought to testify to sophisticated
pyrotechnological know-how by early modern humans in South Africa [15].
 2

However, connecting archaeological objects to inferences regarding cognitive abilities

and the possible presence of “fully syntactical” language remains a very controversial
issue. In a dissection of the conceptual anatomy of such bridging attempts, Botha has
shown the (often implicit) assumptions and series of inferential steps archaeologists have
to make before being able to squeeze “language” out of mute artefacts [16, 17, see also
18, 19], stressing the weak spots in those arguments. However, the engravings on the
ostrich eggshell containers documented in the Howiesons Poort of Diepkloof [20, 21]
indicate the existence of a graphic tradition of communicating among members of the
MSA groups inhabiting the site.
Refuting evidence. A recent analysis of the microstructure and bio-mechanical
performance of the hyoid bone from the 60,000 years old Neandertal skeleton at Kebara
[22], concludes that this hyoid is not only similar to those of modern humans, as already
suggested in 1989 [23], but also suggests that it was used in a very similar way. The
hyoid bone plays an active role in speech, by providing support for the larynx and the
tongue, and it seems that Neandertals were capable of speech. But did they possess the
“the critical thought and syntactic ability necessary for complex language?” [24]. Modern
human language is syntactical, that is it consists of discrete components that can be
combined to produce an almost unlimited number of sentences to communicate about
objects, events and ideas [24]. Given the difficulties of linking speech to language, we
prefer to refrain from speculation about the evolution of language per se and provide
instead well-documented recent evidence of Neandertal non-utilitarian, symbolic
behavior, comparable to the contemporary MSA record. For a similar approach see
d’Errico et al. [11] and Dediu and Levinson [25]. We review here recent research on
possible pigments and objects which may have been used for personal decoration. The
minerals from the 11 Mousterian sites cited by Demars [2] for a total of 37 layers remain
to be verified by modern analyses; of these sites only two, Pech de l’Aze 1 and IV
(France), have been documented by recent research (see below).

Pigments and objects for personal decoration.

250 pieces of manganese with wear-facets and scraping documented by microscopic
analysis have been found in layer 4 (Mousterian of Acheulian Tradition) at the site of
Pech de l’Azé I. Based on OSL dating of sediments, layer 4 has a weighted mean age of
51.4 ± 2.0 ka [26, 27]. The site of Pech de l’Azé IV has yielded 26 pieces of manganese,
of which 15 bear traces of modification [4]. Several pieces of manganese with traces of
being worked come from the Abri Peyrony in France [28]. The interpretation of
manganese as a coloring material is not as sure as that of ochre due to the scarcity of
ethnographic and other archaeological sources on the use of manganese as a pigment. It
does, however, find some support in the documented use of manganese oxides in Upper
Paleolithic cave paintings [29, 30]. Given its flammable qualities, the use of manganese
dioxide powder to rekindle or maintain a fire cannot be excluded [31].
At the MIS 3 open-air site of Les Bossats in Northern France there are 83 pieces of red
ochre with multiple striations and scraping marks resulting from extracting red powder
[32]. Further evidence for Neandertal use of pigments comes from the finds [33] of three
perforated marine shells from Cueva de los Aviones (southeast Spain); one of the shells
had stains of a red colorant identified as hematite around the perforation. A Spondylus
shell contained residues of red and black pigments mixed with charcoal and pyrite,
 3

suggesting use as a paint container. Lumps of red and yellow colorants were found during
the excavation; the sources of the material are located 3 to 7 km from the site. The
Neandertal occupation is dated at between 45 and 50 ka. At the time of occupation the
site was at some distance from the shore, about 1.5 to 7 km (the coast is in a subsidence
area). There is also abundant evidence of the use of mollusks as food; the number of
identified specimens is 785.
At Cueva Antón, in the same region but 60 km inland, level I-k yielded a perforated
upper valve of Pecten maxima (scallop shell) which on its external side was covered with
a mixture of yellow goethite and red hematite (sources for which are about 5 km from the
site). The perforation may be natural but the hole existed before the application of
colorant. The level was radiocarbon dated to 37-38 ka cal BP [33], but more recent
excavations confirm that the small lithic assemblage represents a Middle Paleolithic
occupation. While there is no clear evidence that the holes in the shells were
anthropogenic, the authors conclude that Neandertals selected beached shells with holes
4.5-6.5 mm in diameter, convenient for stringing.
At the cave of Fumane (northern Italy) a Mousterian layer dated between 47.6 to 44.8 ka
cal BP has yielded a marine shell (Aspa marginata) collected by Neandertals at a fossil
exposure more than 100 km from the site. The shell outer surface was smeared with
ochre, suggesting use as a pendant [34]. Fragments of non-local hematite have been
found at Maastricht- Belvédère Site C, well-dated to minimally MIS 7 by TL and ESR
[35]. Eight containers made of stalagmite fragments with traces of ochre and scraping
marks have been found in two layers of Cioarei Cave in Romania, dated to MIS 3 [36]
Thus two sites (Cueva Antón, Fumane) document transport and coloring of exotic objects
and their possible use as pendants, while the French sites, Ciaorei Cave and Maastricht-
Belvédère site C show that use of red ochre was a component of Neandertal behavior,
since at least 200-250 ka.
Extraction of large feathers from raptors, corvids and pigeons is documented by cut
marks on distal wing and foot bones (elements that have no food value) in Mousterian
layers from the site of Fumane (northern Italy [37] and Gorham and Vanguard Caves
(Gibraltar [38]. Stronger evidence of the systematic use of bird elements is provided by
cut marks on terminal phalanges of eagles at five sites in France and Italy (Fumane, Pech
de l’Azé 1, Combe Grenal, Les Fieux, Pech de l’Azé IV) indicating the removal of the
claw. Photos are available in the publications of first four sites [26, 39-41].
The use of raptor claws is supported by the similar find of a terminal phalange of a raptor
(probably a vulture) with a deep notch, interpreted as a pendant, from the cave site of
Üçağizli (Turkey). The Initial Upper Paleolithic (cf. Emiran) and the Ahmarian layers at
the site have yielded a large assemblage of ornaments, mostly pendants made from
marine and freshwater mollusk shells. The raptor claw, the only non-shell ornament,
comes from layer B (Ahmarian) dated to c. 41 ka [Fig 17 in [42]. The find of Üçağizli in
an Upper Paleolithic context provides support for the hypothesis that raptor claws could
have been used as ornaments by Neandertals. Raptor claws and wing elements are
overrepresented at two Epipaleolithic sites in Israel, Meged and Ohalo II and in both
cases an interpretation of use of claws and feathers as ornaments was suggested [43] .
 4

Social interaction between Neandertals and modern humans.

As noted in the text, there are components of Neandertal cultural behavior that were
demonstrably developed independently, such as the use of red ochre and manganese,
personal ornaments like eagle claws, and bladelet production. In a recent paper [27) it has
been suggested that late Middle Paleolithic lissoirs, a type of specialized leather working
bone tool, may constitute evidence for cultural diffusion from Neandertals to modern
humans. One lissoir comes from layer 4 at Pech de l’Azé I dated to 51.4 ± 2.0 ka, thus
predating the use of similar objects by AMH in Europe (see below “Lissoirs”). In the case
of the Dufour bladelets in the Châtelperronian of Quinçay, where intrusions from
overlying Aurignacian must be excluded by the absence of such levels at the site, a
stimulus diffusion from the Proto-Aurignacian to Neandertals has been suggested [44].
It is interesting that data from Quinçay suggest that diffusion from Neandertals to modern
human groups is also conceivable for another kind of artifact. In the same
Châtelperronian levels of Quinçay there are six perforated animal teeth of different
species (fox, wolf and cervids, see ref [45] for photos of two). A similar conclusion was
reached by Zilhao [46] on the basis of the Arcy Grotte du Renne ornaments. Twenty
eight grooved or perforated animal teeth occur in the Châtelperronian levels [X-VIII) of
Grotte du Renne [47]. Three perforated teeth (possibly of a bear) have been reported from
Bacho Kiro (Bulgaria) in level 11 (the type assemblage of the Bachokirian [48]). The
Bachokirian is considered broadly coeval with the Châtelperronian but its makers remain
unidentified.
Perforated or grooved animal teeth occur in the Proto-Aurignacian but they are not
common. Only one (a bear tooth) has been found in the Proto-Aurignacian level VII of
Grotte du Renne [49: figs. 157-158], one pierced red deer canine comes from the
Rothschild rock shelter (SE France) and two pierced incisors of a herbivore from Isturitz
[46]. To our knowledge none have been reported from pre-Aurignacian levels in the Near
East, including the Initial Upper Paleolithic and Ahmarian levels of Üçağizli cave
(Turkey), which yielded a large number of marine shell ornaments and one grooved
raptor claw [50]. As noted by Zilhao [46] pendants of animal teeth are unknown in Africa
and the Levant, they seem to be a European novelty. Thus if the dates for the
Châtelperronian at 45 to 41ka cal BP are accepted and the Protoaurignacian is dated to
39.9-41.5 ka cal BP, one could conclude that pendants of animal teeth were a
Châtelperronian (Neandertal) invention transmitted to the Protoaurignacian [51].
It is important to note that perforated or grooved animal teeth used as pendants are very
common in the Aurignacian: 486 teeth are recorded from 37 sites in Europe, two of
which also contain pierced human teeth: Les Rois and Grotte des Hyènes at Brassempouy
[52, 53]. Of course, the duration of the Châtelperronian and of the Protoaurignacian is
much shorter and their spatial distribution more limited than that of the Aurignacian. Still
this seems again to be a case of a novelty that was episodically present in Neandertal
assemblages and later became a stable component of cultural behavior.
In conclusion the archeological evidence shows that personal ornaments and
manipulation of ochre and manganese existed in the Neandertal world, well before the
Châtelperronian.
 5

2. Neandertals had limited capacity for innovations

Hypothesis. This observation was first explicitly formulated by Mellars [54], who argued
that rates of change in technology and regional variability were much more rapid in the
Upper than in the Middle Paleolithic. Since technological change is slow throughout most
of prehistory, increases in rates of technological change are thought to signal the
emergence of abilities to develop new techniques and/or devices to cope with adaptive
problems. This capacity is widely viewed as symptomatic of “modern” behavior and
possibly signaling the emergence of a highly structured language [8]. Klein [6] also
emphasized rapid increase in the rate of artifactual change through time and in the degree
of artifact diversity through space in the Upper Paleolithic. This point of view was readily
adopted by other scholars: Wynn and Coolidge [55] for instance state that “Neandertals
lacked the inventiveness, characteristic of people today”.
Supporting evidence. The Upper Paleolithic technocomplexes of Western Europe, each
characterized by distinctive “type fossil” forms both in stone and bone or antler
implements, are confined to certain geographical areas and time periods. Each seems to
last less than 10,000 years. The two main technocomplexes of the South African late
MSA, the Still Bay (SB) and the Howiesons Poort (HP), considered to be two very
dynamic and innovative phases with a large geographical distribution, were until recently
thought to last less than 10,000 years each. The SB is dated at Blombos from about 75.5
to 67.8 ka [56] and to about 70 ka at Sibudu [57]. The HP at Klasies River, Sibudu and
Rose Cottage has dates from 65 to about 60 ka [57-59].
Refuting evidence. Recent research at the site of Diepkloof (Western Cape, South Africa)
has shown that these two MSA technocomplexes have a significantly longer duration. In
particular, the HP has now been subdivided in three phases: Early, dated by OSL to
109±10 ka and by TL to 105±10 ka; Intermediate, between 85±9 ka and 65±8 ka; and the
Final HP (corresponding in time to and resembling the occurrences of Sibudu, Rose
Cottage and Klasies, hence called the “classic” HP) dated to 52±5 ka [20,60]. The SB is
dated to 109 ±10 ka. The implications seemed clear: the two technocomplexes have a
much longer duration than previously envisaged and they are not homogeneous in space.
However the OSL method used by Jacobs at the MIS4 - MIS3 sites has been criticized
and the dates are judged unreliable [61]. Moreover the nature of the Diepkloof SB and its
relationship to the much younger assemblages of Blombos, Sibudu and Hollow Rock
Shelter remain to be explored. The same is true for the HP assemblages dated to MIS 5 at
Diepkloof. In contrast the dates for the Post-HP technocomplex based on several dating
methods (TL, OSL, ESR, 14C, U-series) are more secure and can be compared to the
dates of Upper Pleistocene technocomplexes in Europe, as in Table 1 (text).
Middle Paleolithic industries in Europe dated between MIS 5 to MIS 3 show clear spatio-
temporal distributions [62, 63]. The Mousterian of Acheulian Tradition of Southwest
France dates to between 70 and 40 ka, based on dates obtained from six sites, and hence
lasted approximately 30,000 years. The Quina Mousterian dates to between 73 and 40 ka,
based on dates obtained from six sites [64]. The Keilmessergruppen (KMG) in Central
Europe have bifacial tools which are technologically and typologically well-defined
guide fossils. The KMG can be subdivided into separate inventory types in a clear
chronological succession and with regional differentiation [63, 65].
In conclusion, the pace of change in the last phases of the Middle Paleolithic, prior to the
arrival of modern humans, is comparable to that of the late MSA in South Africa.
 6

3. Neandertals were less effective hunters

The idea that scavenging was an important pattern of subsistence behavior of early
hominins and the Neandertals was first suggested by Binford [66, 67]. Later Stiner [67]
argued that Neandertals in Central Italy had a flexible adaptation alternating scavenging
with hunting, which became prevalent only in late Neandertal times. These ideas have
been disproven by a number of studies [69, 70], and accumulated evidence shows that
hunting was the main method of meat procurement by Neandertals [71, 72]. The
scavenging hypothesis is dead and we will not discuss it any further. However the idea
that Neandertals were less effective hunters has been reformulated in terms of their
hunting methods and weapon technology.

Hunting methods
Hypothesis. Neandertals hunted in small groups in long dogged pursuits across the
landscape, targeting the largest prey animals. Their killing ability was limited to thrusting
spears equipped with simple stone points [73-75]. Their encounter technique involved
high risk close contact with large prey animals resulting in high trauma rates; patterns and
frequencies of Neandertal bone breakage fall outside the norm for modern humans [74],
except for twentieth-century rodeo riders [76]. In contrast to modern humans,
Neandertals did not focus on communal hunts and herd intercept at times of migration
and did not make efficient use of the landscape to hunt their prey. Their hunting
efficiency, as measured by time and effort expended to kill their prey was, less than that
of modern humans in Africa and in Europe [74].
Supporting evidence for organizational techniques and killing abilities. Neandertal
skeletons show extreme physical activity and healed upper-body fractures [76] indicating
close combat with spear thrusting. Their sites were small, focused around a single hearth
with little structured use of space [67, 77]. The implication is that the Neandertal small
sites do not indicate large group organization of the type typical for groups with
communal hunts.
Refuting evidence.
Traumas. This reading of trauma in Neandertals versus modern humans has now been
retracted by Trinkaus himself [78], stating that “assessment of Late Pleistocene modern
humans indicates a similar pattern to that of Neandertals…explaining the trauma pattern
solely as a result of Middle Paleolithic hunting weaponry or conversely using the
Neandertal trauma pattern to argue for ineffective Middle Paleolithic weaponry, should
be abandoned”.
Communal hunts and tactical thinking. A number of sites dated between MIS 9 and MIS
3 show evidence indicative of organized hunts, using topographic setting for killing one
or two species of large mammals (Table S1). Four sites show the use of cliff faces and
rocky barriers associated with karstic systems for bringing down large mammals. At
Schöningen the remains are in organic muds and the age distribution indicates the
presence of at least one family group of horses. The fossil material is well preserved with
little variation in weathering [79] suggesting rapid accumulation. The French sites appear
to have been formed by a number of different hunting episodes, with the MIS 6 site
Coudoulous containing the remains of 232 bison. The evidence shows that Neandertals
were able to organize game drives using landscape features as natural traps, intercepting
 7

groups of animals at repeatedly used locations. At other times large and medium size
mammals were hunted individually. Scheduled hunting is not, as suggested by some [79],
the exclusive domain of modern humans.
Table S1. Sites with evidence of repeated kill episodes using topographic settings. Data
from [72, 79, 81] and references therein.
Site Age Main fauna MNI Topographic setting
Schöningen MIS 9 Equus 20- Muddy shore of a lake
13 II-4 mosbachensis 25
(Germany)

La Cotte St. Brelade, (Jersey) MIS 6 Mammuthus 7, 11 Base of a deep ravine

layers 3 and 6. primigenius,
Coelodonta 2, 3
antiquitatis
Coudoulous (SW France) MIS 6 Bison priscus 232 Karstic depression, open air
Layer 4
La Borde (SW France) single MIS 7 Bos primigenius 40 Karstic depression, open air
layer or 5
Mauran (SW France) MIS 3 Bison priscus 137 Base of an escarpment with
rocky barrier

Internal site structure and group size. Spatially organized activities and well delimited
activity areas have been documented by refitting at a number of Middle Paleolithic open-
air sites, such as Maastricht-Belvédère site K (Netherlands, MIS 7) [82], La Folie, SW
France, estimated age MIS 5e [83], Ksiecia Jozefa layer III (MIS 3) [84] and at rock
shelters such as Abric Romani (Spain, MIS 3) [85, 86] and Tor Faraj (Jordan, 55 ka) [87].
Intra-site spatial analysis combined with refitting and hearth distributions documents two
living floors at Tor Faraj, with an estimated range of 15 to 20 persons as hearth-side
occupants in each living floor, an area of 136 m2 and multiple, spatially segregated task-
specific areas. These estimates correspond to the number of people commonly seen in
modern foraging bands.
Evidence of complex internal site structure is also indicated at Kebara [88, 89], at the
open air site of Quneitra [90] and at Abric Romani [91]. At the latter site the spatial
distribution of wood remains in different areas of the shelter has been interpreted as
revealing systematic collecting and even storing of fuel sources (twigs and small
branches for lighting, large branches and trunks) to supply the 37 hearths of layer M,
dated to ca. 50 ka.
Hunting efficiency. It may be worth noting that hunting efficiency is hardly a measure of
modernity. The descriptions of San hunting techniques and recovery rates indicate clearly
that they invest an inordinate amount of time tracking wounded animals across the
landscape (because their bows are weak and they use a slow-acting poison) and their
recovery rates are relatively low, because they often lose their animals to other predators.
Return rates are also low for the Hadza, who get higher caloric returns from gathering
[92, 93].
 8

4. Neandertal weaponry was inferior to AMH projectile technology

The hunting weapons of Neandertals involve a number of arguments based on anatomical
and archeological evidence.

4.1 Hypothesis. Neandertals only had thrusting spears

Supporting evidence. Patterns of bone frequencies and breakage (see above “Traumas”)
and the upper limb morphology of Neandertals indicate close combat with spear
thrusting. Upper limb morphology associated with projectile throwing occurs in Upper
Paleolithic humans but is absent in Neandertals [73, 74].
Refuting evidence. In contrast to the Lehringen spear, interpreted as a thrusting spear,
experiments and measurements show that the Schöningen spears had ballistic qualities
indicating that they were thrown as javelins [94, 95] (and refs therein). The humerus and
shoulder morphology associated with overarm throwing were already present in Homo
erectus [96]. The particular characteristics of Neandertal humeri may reflect adaptation to
regular scraping activities and not spear thrusting [97].

4.2 Hypothesis. Complex projectile technology gave early modern humans a competitive
advantage over Neandertals.
Modern humans used spear throwers and/or bow and arrow to launch their projectiles.
This would have given them an advantage in their competition with Neandertals [98, 99].
Supporting evidence. Of the variety of measurements proposed to identify prehistoric
stone points as projectile points, the tip cross-sectional area (TCSA) is a widely applied
one [98, 100-101]. Based on a series of North American arrowhead and spear-thrower
launched dart tips as well as on experimental studies, Shea has suggested that early stone
projectile points should not present higher TCSA values than the more recent ones in his
sample. TCSA values of a series of Middle Paleolithic Levallois points were significantly
larger than the ones from his control sample, whereas the values from EUP points did not
differ from the control sample points. Values of TCSA from Blombos and other South
African sites also suggested to him that some of the points were used as spear-thrower
darts [103]. These results were interpreted as implying that Neandertals did not have a
complex projectile technology, in contrast to late MSA and EUP humans.
Refuting evidence. The TCSA values from Blombos used by Shea are incorrect, because
he used a data base which contained preforms and unfinished pieces (thus obtaining small
values from broken pieces) and because he excised large values using an arbitrary cut-off
point [94].
A recent study of Australian hafted dart projectile points [104] shows that TCSA values
of Australian macro-blade darts are significantly larger than the ones from the North
American dart points; in fact, their mean is significantly larger than the values Shea
obtained for (early) Middle Paleolithic Levallois points (compare Fig 4 in ref [104] with
Fig. 6 of ref. [101]). In other words the TCSA statistics have a limited value and cannot
be seen as documenting a particular delivery system.
For instance, Mousterian points from Italian and Spanish sites have significantly lower
TCSA values than those used by Shea for thrusting spears and are comparable to those
for darts [95]. We are not suggesting that Neandertals had spear-throwers. In fact, the
key issue illustrated by the Australian dart points is the importance of variability, both
within prehistoric and extant human hunter-gatherer populations [105]. Interestingly,
 9

hunter-gatherers in Australia are not known to have used bow-and-arrow. Tasmanian

aboriginals did not even use spear throwers, and only had hand delivered spears [106,
107].

4.3 Hypothesis. AMH had bow and arrow, Neandertals only hand-cast spears
This idea is based on two separate claims: a) small quartz pieces used to tip arrows occur
in the Howiesons Poort (HP), and b) backed bladelets made of heat-treated silcrete and
used as arrow-heads represent a weapon technology originating 71,000 years ago in a
pre-HP industry in South Africa.

a) Quartz-tipped arrows in the Howiesons Poort

Supporting evidence. A hypothesis of quartz-tipped arrows in the HP of South Africa,
first proposed in 1982 for the Klasies River Main Site [108] has been suggested for the
Sibudu HP by Wadley and Mohapi [109] and supported by Lombard [110] based on the
presence of very small quartz segments (<2.5 cm) interpreted as transverse arrowheads.
They conclude that projectile technology was developed in the MSA of South Africa
before 60 ka.
Refuting evidence. This claim is poorly supported owing to (i) the very small number of
possible arrowheads compared with the total of backed pieces at both sites, cf. below, (ii)
the fact that equally small or smaller pieces were used as barbs on spears in the Upper
Paleolithic, and (iii) the total lack of evidence for arrowheads in post-HP assemblages of
the following 15-20,000 years, including Sibudu [111]. Bow and arrow are generally
recognized as a successful weapon, widely adopted on most continents, used for hunting
a wide range of game in forested as well as in grassland settings. The apparent
subsequent loss of this successful technology is explained as a historical contingency
[111], that is, a chance historical event, due to undetermined or random factors. A simpler
alternative might be that there was no bow and arrow technology to loose.
It is only at about 44-42 ka that there is good evidence for the use of (poisoned bone)
arrows at Border Cave in South Africa [112], well after the hypothesized migrations of
modern humans into Eurasia [113]. At the White Paintings rock shelter in Botswana bone
points that appear to have been parts of reversible arrowheads are dated to 37-35 ka
[114].
As indicated above, the number of quartz pieces is very small. At Klasies there are 14
pieces < 2.5 cm of quartz and silcrete of a total of 109 backed pieces of all raw materials
in layer 20 [95]. At Sibudu the total number of quartz backed pieces from the HP layers
interpreted as arrowheads is 14 of a total of 281 backed pieces of all raw materials [95,
115] .
Impact notching on thin edges has been observed in experiments with replicated backed
pieces hafted transversely [116] and used to support an interpretation of transverse
arrowheads for 9 small quartz segments from HP layers of Sibudu [115]. However the
impact notches so described do not have a negative bulb and cannot be distinguished
from micro-fractures and pseudo-retouches produced by trampling or post-depositional
damage in the sediments [117, 118]. We do not consider this kind of notching diagnostic
of impact.
Experimental studies using 265 replicated Kebaran and Natufian microliths hafted on
arrows and shot with a bow [119] report bending fractures (type b3 in fig. 3 and Table 2
 10

in [119] and step-terminating scars (type b3 and d2m in figure 3 and Tables 2-3) for
transversely hafted microliths. The scars are unlike the notches described in Lombard and
Pargeter [116]. The latter type (d2m) also occurs on microliths used as barbs. For this and
other reasons, especially the absence of evidence for bow and arrow technology in post-
HP times [111], we consider the hypothesis of quartz-tipped arrows in the HP of Sibudu
poorly supported. Even if one should consider the bow and arrow hypothesis a
possibility, the record shows it was a short-lived experiment.

b) Backed bladelets used as arrow tips represent a weapon technology originating

71,000 years ago in South Africa.
Supporting evidence. Small backed bladelets, made on heat-treated silcrete, occur at
Pinnacle Point Site 5-6 and are found in layers dated between 71 and 60 ka [99]. The
Pinnacle Point evidence supposedly indicates that the production of backed bladelets is
even earlier than the HP and represents a long-lasting technology that “conferred
substantive advantage on modern humans as they left Africa and encountered
Neandertals equipped with only hand-cast spears” [99].
Refuting evidence. The proposed date of 71 ka for the origin of backed bladelet
technology has been shown to be incorrect since backed bladelets occur much earlier at
Diepkloof in the Early HP, dated by OSL to 109±10 ka and by TL to 105±10 ka [20]. We
will not deal with this chronological problem here, only with the claim of a technological
supremacy of early moderns.
The idea that backed bladelets provided modern humans with projectile weaponry that
gave them a huge advantage on Neandertals, once modern humans moved out of Africa is
difficult to prove with the existing data. Intentional bladelet production appears at Umm-
el Tlel (Syria) at about 40,000 BP (based on uncalibrated 14C dates) in the Early
Ahmarian at about 41.6 - 40.3 ka cal BP at Ksar Akil and Üçağikli [120-121] and in the
Proto-Aurignacian at c. 41.5-39.9 ka cal BP [122]. The Emiran or Initial Upper
Paleolithic industry (IUP) of Boker Tachtit (Israel) levels 1 and 2, dated to about 47,280
± 9050 and 46,930 ± 2400 (these are dates for Level 1 but note the very large standard
deviation) lacks backed bladelets [123], while the similar IUP assemblages of Tor Sadaf
A and B (Jordan) have only a very small number of retouched bladelets [124]. In Emiran,
Bohunician or Bachokirian industries points predominate and were probably used to arm
spears [125]. The success of bladelet production begins with the Ahmarian in the Levant
and in the Proto-Aurignacian in Europe. According to recent OSL dates, microblade
technology appeared in India at 44 ± 2 ka (weighted average of four dates) at the open-air
site of Mehtakeri [126] roughly comparable in age to similar dates for the Ahmarian.

If the age of migration of modern humans into Southwest Asia and India is set prior to 60
ka, there is a difference of more than 15,000 years between the hypothesized migration
and the occurrence of bladelet production in the Near East and India.
If the migration is set at c. 60-50 ka [113] this would rule out South African populations,
since by 60 ka the HP tradition of backed tools made on blades and bladelets produced by
soft stone hammer had given way to the post-HP assemblages characterized by a variety
of flake tools and blades produced by hard hammer percussion, but without backed
blades [127].
 11

If modern humans migrated from East Africa shortly after 100 ka [128] the idea of an
intergenerational transmission of a long-lasting technology of backed pieces helping the
spread of humans out of Africa is again ruled out by the difference of c 60,000 years
between the hypothesized migration, the Neandertal demise and the systematic
production of bladelets in the Near East and India. The latest Neandertal currently known
from the Levant is the adult male skeleton from Amud Cave (Israel) with an ESR date of
53 ± 8 ka on tooth enamel [129] corresponding well with TL dates on heated flints from
the same deposits [130].

Finally, if the new technology spread from Africa to the rest of the world accompanying
the dispersal and supremacy of modern humans, why is it that the hunting arsenal of
Australian Aboriginals does not include bow and arrow? In North America the spear-
thrower was probably introduced by the first settlers but the advent of the bow is much
later, roughly around 2300-1300 BP in western North America, even later in the eastern
part of the country [131, 132].
In conclusion, the idea that bow and arrow technology was a factor in the demise of
Neandertals remains as yet unsupported.

5. Neandertals had a narrow diet that proved unsuccessful in competition with

modern humans that had a more diversified diet.
Hypothesis. Neandertal diet was restricted to large and medium size herbivores. Early
modern humans had a broader diet, including significant quantities of aquatic foods.
Neandertals were unable to acquire small fast game (birds, wild rabbits) and did not
exploit plant resources [74, 75].
Supporting evidence. Carbon and nitrogen isotope values of Neandertal bones from
northern regions or cold environments such as Belgium, Germany, Southwest France, and
Croatia, indicate that Neandertal were top-level carnivores obtaining most of their
proteins from large herbivores such as bovids and horses [133]. More recently isotope
analysis has been applied to Neandertal remains from 2 Spanish sites, Cova Negra and El
Salt. The results also show that Neandertals consumed large amounts of meat [134].
Refuting evidence. Isotope analyses have a low resolution detecting plant protein when
the diet is dominated by animal protein. Nutritional ecology studies have stressed that
Neandertals would have needed more than just large mammal protein, including a range
of micronutrients not readily available through terrestrial mammals [135]. A broader
picture is provided by zooarchaeology and studies of plant remains.
The faunal remains in Mediterranean regions indicate an abundance of small ungulates
i.e. smaller than red deer (gazelle, roe deer, fallow deer, caprids) at cave sites such as
Kebara, Misliya, Amud and Qesem [136]. Small prey, such as wild rabbits, is well
documented by cutmarks and anthropic fractures at Bolomor Cave (Spain) in layers
XVIIc (dated to MIS 9, NISP = 457) layer XI (MIS 6, NISP = 262) and layer IV (MIS 5e,
NISP = 789) , which also yielded abundant evidence for tortoise consumption [137-139]
at Les Canalettes layer 4 (France, c 70 ka; NISP = 1209) [140] and at Abri du Maras
(Ardèche Valley, France, beginning of MIS 4, only a few remains but with two tools also
showing leporid hair fragments on their working edges [141].
Layer XI of Bolomor Cave has also yielded 202 NISP of the genus Aythya (diving duck)
corresponding to a MNI of 8. This species which lives in marshes and lakes is
 12

represented by various elements, especially leg bones and the coracoids. Cutmarks and
bone breakage indicate human consumption [142]. At the late Middle Paleolithic open air
site Salzgitter-Lebenstedt (Germany), cut-marked bones of duck and swan were
identified [71]. The capture of raptors, corvids and wood pigeons for removal of wing
feathers is documented by cutmarks and bone breakage at Fumane in northern Italy [37]
and at Gorham and Vanguard Caves in Gibraltar [38]. At the latter sites the capture of
birds may perhaps have been related to use of feathers as ornaments (see Hypothesis 1).
The capture of hare, beaver, marmot and some carnivore (bear, fox and mustelids,
documented by cutmarks and possibly related to the use of pelts) is documented again at
Fumane; cutmarks also occur on beaver bones at Grotta Maggiore di San Bernardino in
northern Italy [39]. Cutmarks and bone breakage patterns document the capture of rabbits
and birds at an even older site in Spain, Gran Dolina level TD 10-1, TL dated to MIS 9
[143].
Evidence of plant consumption by Neandertals is provided in Table S2.

Table S2. Plant consumption by Neandertals. Data based on [134, 144-149]

Site Archeological evidence Kind of food

Kebara (Israel) 4205 charred seeds and Legumes (wild peas),
fruit, phytoliths acorns, pistachios
Amud (Israel) Phytolith analysis Grass seeds
Gorham’s Cave Charred nuts Pine nuts
(Gibraltar)
Spy (Belgium) Microfossils in dental Water lilies? grass seeds
calculus
Shanidar (Iraq) Microfossils in dental Date palms, grass seeds
calculus
El Sidrón (Spain) Microfossils in dental Cooked carbohydrates
calculus
Sima de las Palomas Microfossils in dental Grass seeds and other plant
(Spain) calculus types

Fishing. Remains of freshwater fish (trout, chub, eel, NISP =21) are documented in
various Mousterian levels of Castelcivita Cave (Italy) older than 40 ka [150]. The
Mousterian layers there underlie the Uluzzian, Protoaurignacian and the Campanian
ignimbrite ashes. Additional evidence of freshwater fish consumption is provided by the
sites of Payre in France [151], Kudaro in the Caucasus [152]] and Abri du Maras (MIS 4;
mostly cyprinids and percid, NISP = 167) [141]. At Abri du Maras the contribution of
non-human predators to the accumulation of fish bones and scales is excluded by the
absence of any signs of chewing and digestion. More taphonomic studies are needed to
ascertain the role of fishing activities in the diet of Neandertals. It needs to be stressed
that in Europe high numbers of fish remains are documented only in the late Upper
Paleolithic [132].

In conclusion, recent research by different scholars using different analytical methods

shows that Neandertal diet was more diverse than generally acknowledged, and varied
 13

according to different environments. A broad-based subsistence was already in place by

the second half of the Middle Pleistocene and was not limited to Mediterranean regions,
as indicated by sites such as Spy, Kudaro, Shanidar, El Sidrón, Payre and Abri du Maras,
going from Belgium to Iraq and from the Atlantic sea shore to the Caucasus.
6. The use of traps and snares to capture animals is the exclusive domain of modern
humans.
Hypothesis. It is often suggested that the capture of fast game such as birds and wild
rabbits implies the use of traps and snares. For some authors [80] the use of snares and
traps is an indication of advanced planning abilities and complex cognition which are
attributes of the modern mind and hence relatively recent developments. Use of plant
fibers for binding, weaving, basketry and possibly net-making is documented in the
Upper Paleolithic of Dzudzuana Cave in Georgia at 30 ka [153] and at the roughly
contemporary Gravettian sites of Dolní Vestonice and Pavlov 1 in the Czech Republic
[154]. High numbers of small fur-bearers like hares and foxes have been recovered from
the Gravettian sites.
Supporting evidence. The supposedly rare presence of bird and lagomorphs remains in
the Middle Paleolithic has been interpreted as due to absence of trap-related technology
and related to low population densities [155]. Indirect evidence of using traps in the MSA
has been argued by Wadley [156] based on the abundance of tiny blue duiker remains,
easily captured with nets or snares, in the Sibudu Howiesons Poort (c. 65-61 ka) faunas.
Refuting evidence. Our data on the capture of small fast game in the Middle Paleolithic
(Hypothesis 5) suggest that catching smaller and fast moving game, thought to indicate
the “enhancement of working memory capacity” [80], was already within the realm of
Neandertals, as early as Middle Pleistocene times. This is supported by the presence of
twisted fibers (found as residue on 12 stone artifacts) at Abri du Maras (France, early
MIS 4]. Since fibers are not twisted in their natural state they provide tantalizing
evidence for the manufacture of strings or cordage which would facilitate the
manufacture of multi-component technology such as snares [141].

7. Modern humans had larger social networks

Hypothesis. Long-distance exchange networks can operate in times of scarcity to allow
access to distant areas where resources are plentiful, thus acting as a buffer against
environmental downturns and increasing the long term survival of groups [157, 158].
Modern humans established such far-reaching extended social networks, unlike those of
the Neandertals [159].
Supporting evidence. Long distance transfers of lithics and especially shells, inferred to
reflect the geographical size of social networks, do increase in the Upper Paleolithic of
Europe, both in terms of quantity of transported (lithics) items and the distances from
their sources, though the changes are not dramatic. A number of Upper Paleolithic
assemblages (n=23) documents maximum distances covered of > 300 km, from the
Aurignacian onward, with the largest distances documented for the Gravettian and the
Magdalenian, virtually all entailing movement of shells [159-165].
At Ortvale Klde (Georgia) the source of high-quality obsidian was about 100 km to the
southeast. A comparison of the late Middle Paleolithic and the early Upper Paleolithic
assemblages shows higher proportions of obsidian in the early Upper Paleolithic. In the
early Upper Paleolithic obsidian is represented by full reduction sequences while in the
 14

late Middle Paleolithic obsidian artifacts appear as small debitage and heavily reduced
tools. The conclusion is that Neandertals moved only infrequently between regions and
their social relations were structured into smaller territories than those of the Upper
Paleolithic modern humans [166].
However all these are comparisons between Neandertals and their younger successors.
We should compare Neandertals with their modern human contemporaries in Africa and
this is the evidence we consider here. McBrearty and Brooks [157] report distances of
transport of obsidian at MSA sites in East Africa up to 240 km at the Nasera Rock Shelter
and 320 km in Bed VI at Mumba (both sites in northern Tanzania). In the MSA of South
Africa fine-grained raw materials have been reported as non-local. Deacon [167] and
Wurz [168] suggest that the backed pieces of the HP had a symbolic value and may have
been used in a reciprocal exchange network.
Refuting evidence. The quantities of obsidian transport in the early MSA of East Africa
are small, indicating weak group interactions [169]. This is also true of Middle
Paleolithic assemblages where maximum transport distances in Western Europe are
around 110-120 km (contrary to the opinion of [170]).
It is only at the end of the MSA or during the MSA/LSA transition (older than 40 ka) in
assemblages like Prolonged Drift and Ntumot in Kenya, that the proportions of obsidian
from sources 45-70 km away increase, possibly indicating more intensive intergroup
exchange [169]. In the later Middle Paleolithic record of Central and Eastern Europe
maximum distances are much larger than in Western Europe (> 200 km), usually
interpreted as a result of ecological differences: in exceptional cases trans-Carpathian
transport moved materials up to 280 km [160, 162, 171, 172]. While these large distances
again mostly relate to isolated and heavily reduced pieces, the Last Interglacial (MIS5)
levels at Kulna cave in the Czech Republic [173], yielded exotic materials in the form of
blanks and rough cores, imported over a distance of 230 km [161], i.e. comparable to the
form of transport recently described for the 50-100 ka old deposits from the White
Painting Shelter site in Botswana [174].
The conventional view that HP was associated with long distances of raw material
transport is incorrect. At several sites raw materials were local or transported over short
distances. At Rose Cottage the main raw material (frequencies of cores and retouched
pieces > 90%) is opaline which occurs in the Caledon river gravels 8-10 km from the site
[58]. At Sibudu the main raw materials are hornfels, dolerite, quartz and quartzite, all of
which occur near the site or at a distance of about 15 km [175]. At Klasies River quartz
was interpreted as non-local by Wymer but it is very likely local [176] since it occurs as
seams and cobble inclusions within the Table Mountain Sandstone, the parent material of
the Klasies River Cave. The sources of silcrete are not known; however the presence of
rolled cortex on several silcrete cores and retouched pieces suggests beaches (which are
adjacent to the site) or river gravels as one of the sources [59]. At Blombos there seem to
have been two sources of silcrete: from deposits 30 km north of Blombos and from river
valleys which exit into the ocean 20 km east and west of the site [177]. At Diepkloof
high-quality silcrete comes from sources > 20 km from the site [178]. At Border Cave the
main raw material in the HP and younger periods is rhyolite which is the local bedrock.
Another, less abundant, raw material is chalcedony, which occur as vesicles in the
rhyolite and these seem to be most common 40 km from the site [111]. In sum, the MSA
 15

and Middle Paleolithic records for raw material transfer distances are not significantly
different.
Finally the idea that Neandertals groups were “limiting themselves to a single river valley
and only occasionally did they venture farther afield” [55] is refuted by abundant
evidence for transport of raw materials across major river valleys in the Middle
Paleolithic of western and central Europe [160-161, 163, 172].

8. Modern human populations were larger than Neandertal populations

Hypothesis. Modern human initial populations entering Neandertal territory were
significantly larger, in southwestern France an estimated ten times, than the regional
Neandertal populations: “…numerical supremacy alone must have been a powerful, if not
overwhelming, factor in direct demographic and territorial competition between modern
humans and Neandertals” [179].
Supporting evidence. Mellars and French have attempted to estimate population numbers
for late Neandertals (as reflected by Mousterian of Acheulian Tradition and
Châtelperronian sites) and the earliest modern humans (thought to be reflected in
Aurignacian sites) for southwestern France using archeological proxies: (i) data on total
numbers of occupied sites in the study region during the Neandertal–to–modern human
transition period, (ii) the overall intensity of occupation in these sites, as reflected by the
accumulation rates of stone artifacts and animal bones in these sites and (iii) data on the
overall spatial extent of the archeological occupation levels, as a potential reflection of
the sizes of the human groups who occupied the sites in the period at stake. With the
arrival of Aurignacian producing groups they observed a sharp increase in the total
number of occupied sites, much higher densities of occupation and larger areas of
occupation in the sites, thought to be proportional to the number of people occupying a
site. They conclude that earliest modern human populations penetrated the region in at
least ten times larger numbers than those of the local Neanderthal populations in the same
regions.
Refuting evidence. The Mellars and French population size hypothesis has recently been
reviewed in extenso by Dogandžić and McPherron [180], who analyzed each of the three
inferred proxies for population sizes, in terms of their assumptions, their significance and
their archeological visibility. Estimating population numbers using archeological proxies
is a challenging enterprise, as acknowledged by Mellars and French themselves. But
some of the problems were created by their own methods. For instance, the list of sites
inhabited by the last Neandertals does not include sites belonging to different
technocomplexes (Denticulate Mousterian and Quina Mousterian) and yet falling into
their selected time span (even more sites can be added to those listed by Dogandžić and
McPherron). Stone artifacts and faunal densities are generally calculated by cubic meters
of excavated deposits. Mellars and French instead calculated densities by square meters
per 1,000 years of estimated duration of each technocomplex, a debatable procedure since
it assumes that a local occurrence of a technocomplex spans the entire known
chronological range of that technocomplex. The proxy used to calculate the number of
people living at a site is also affected by various factors, such as a) the small size of some
caves which constrains the use of space, b) in the case of old excavations by inadequate
documentation of the site size, and c) changes in site configuration through erosion.
Dogandžić and McPherron show in detail that the Mellars and French analysis is severely
 16

flawed, concluding that “…Mellars and French [179] have biased their results in favor of
a conclusion that AMH populations were larger than late Neandertal populations. With a
more conservative set of assumptions and expected outcomes, their own data show
roughly equivalent values for the respective proxies meaning that they have not
demonstrated a tenfold increase in Western Europe at the Neandertal-to-modern human
transition”(p. 313). The Mellars and French rebuttal [181] fails to address their critique
adequately, and in fact even underlines it, by admitting that taking into consideration
“…the maximum allowance for factors of this kind…” (as discussed by Dogandžić and
McPherron) would lead “..to a more modest increase in our calculated demographic
estimates - say down to a five-fold as opposed to ten-fold population increase “.
Conard [182] uses a better controlled set of variables (size of excavation area and number
of lithic and organic artifacts by cubic meter) to reach an approximation of occupational
intensity and mobility of Neandertals versus Aurignacian groups in the Swabian Jura in
southern Germany, a hilly region rich in cave sites with Middle Paleolithic and
Aurignacian deposits. The study area is approximately 5000 km2, much smaller than the
study area of Mellars and French (75,000 km2). He concludes that Neandertal population
densities in the region appear to have been uniformly low, lower than in the subsequent
Aurignacian period. There are however significant gaps and uncertainties in our
knowledge, as noted by the author himself. Open-air sites had to be excluded from
analysis due to lack of preservation of organic artifacts, so only ten multi-component
cave sites could be analyzed The high residential mobility typical of Neandertals [183]
probably resulted in brief periods of occupation and low densities of artifacts. In sum,
archeological data suggest -but do not provide decisive evidence- that small population
size may have played a role in the replacement of Neandertals by AMH in that region.

9. Hafting, heat treatment and modern cognition

Complex hafting procedures which required use of fire and the use of fire for pitch
production are discussed in the text. In fact, heat treatment was a component of
Neandertal technology. Neandertals used fire to heat-treat existing natural materials, such
as bitumen for hafting purposes. This is clear from 70,000-y-old tools with traces of
bitumen on their surfaces from the site Umm El Tlel (Syria). The bitumen was collected
at 40 km from the site [184]. Bitumen was also recovered from a Mousterian tool at the
site of Gura Cheii-Râşnov Cave in Romania [36].
The idea that Neandertals had unsubstantial hearths, less effective than those of modern
humans in protecting from the cold [55, 185, 186] tends to persist but there is ample
published evidence that it is incorrect [187].

10. Climate as a factor in the extinction of Neandertals

Hypothesis. The Campanian Ignimbrite (CI), the largest volcanic eruption in Europe,
from the Phlegrean Fields north of Naples, spread ashes over an area of 5,000,000 km2,
eastward to Greece, Bulgaria and into Russia. As it occurred right after the onset of the
Heinrich Event 4, dated to c. 40 ka, and characterized by very cold and dry climate, it is
likely to have caused a volcanic winter [188].
The climatic deterioration and resulting population redistribution may have triggered the
major cultural changes of the Middle to Upper Paleolithic transition, ultimately leading to
the decline of Neandertals [189].
 17

Supporting evidence. The chemical composition, age and spatial distribution of the CI are
well documented. At some sites (Crvena Stijena in Montenegro, Franchthi Cave in
Greece and Temnata (Bulgaria) the tephra occurs above the last Mousterian layer and
below the Upper Paleolithic [190]. However, no strong evidence in support of the idea of
a demographic crisis of Neandertals is provided by Fedele et al. [189]. Giaccio et al.
[191] consider the climatic instability of MIS as a potential factor involved in the cultural
changes of the Middle to Upper Paleolithic transition.
Refuting evidence. The discovery of cryptotephra ash deposits (layers not visible to the
naked eye) above the Uluzzian, Proto-Aurignacian, Initial Upper Paleolithic and Dabban
layers at sites such Haua Fteah (Libya) and four sites in Central Europe (Klissoura,
Golema Pesht, Kozarnika and Tabula Traiana) combined with previously known
occurrences of the tephra above Proto-Aurignacian deposits (at Serino and Castelcivita,
in Italy) and the Initial Upper Paleolithic (at Kostienki-Borschevo, Russia) show that
neither Neandertals nor modern humans were adversely affected by the climatic cooling
[192]. We conclude that the decline of Neandertals was not associated with the Heinrich
Event 4 and the CI eruption.

11. The Toba eruption as an indirect factor in the extinction of Neandertals

Hypothesis. The super-eruption of Mount Toba (Sumatra) at about 75 ka caused a
“volcanic winter” of such a catastrophic scale that it led to a human genetic bottleneck,
reducing the number of AMH in Africa to near extinction. The modern human population
expanded again later and migrated out of Africa [193, 194] eventually causing the demise
of Neandertals.
Supporting evidence. The bottleneck would explain the limited modern-day genetic
diversity. Release from the bottleneck with a significant increase in population could
have coincided with a climatic amelioration of the Dansgaard-Oeschger event 19. The
chronology of the earliest LSA in East Africa at about 40-50 ka for the Nasampolai
industry at Enkapune Ya Muto [195] and the Middle to Upper Paleolithic transition fits
the genetic evidence.
Refuting evidence. There is no associated evidence of mammal decline or extinction even
in environmentally-sensitive species. There is no evidence of habitat reduction at that
time in Africa [196]. More important, analysis of crypto-tephra (ash) from the Toba
eruption in a layer of Lake Malawi (East Africa) and accompanying sediments shows that
there is no major change in temperature or sediment composition. The Toba eruption did
not have a significant effect on the climate of East Africa and was not the cause of a
human bottleneck in Africa at around 75 ka [197]. Likewise, there is archeological
evidence that the Toba eruption did not affect the behavior of populations inhabiting
peninsular India [198].
It has been claimed that the response of modern humans to the unpredictable environment
of the period was to extend inter-group cooperative networks to reduce risks while
Neandertals continued to live in small territories with limited intergroup interaction, often
involving violence and cannibalism. These differences may have been responsible for the
replacement of Neandertals by African modern humans [199]. Since there was no
“volcanic winter” in Africa, such inferences must also be rejected.
 18

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 1

Neandertal Demise: An Archaeological Analysis of the Modern Human

Superiority Complex

Paola Villa, Wil Roebroeks

Supporting Information

Text S2. A single package of modern behavior?

The Upper Paleolithic record of Europe has often been treated as a yardstick for modern
human behavior, with its characteristics translated into a trait list used to identify the
presence of modern human behavioral capacities [1]. In actual fact contemporary modern
humans outside of Europe created records which were often very different from the
European ones. The limitations of an “Eurocentric (or rather Western European) outlook
on cultural evolution” [2] have been stressed many times and for various regions,
including the Levant [2], China [3] and North America [4], and especially by students of
the wider Australian record [5]. As detailed by O’Connell and Allen [6, 7], the first
occupants of Sahul were modern humans, seafaring populations, who had crossed
Wallacea and successfully colonized New Guinea, the Bismarck Archipelago and
Australia in probably less than five thousand years. This colonization entailed the
production of boats and the development of sophisticated seafaring capacities, generally
seen as an unambiguous reflection of “modern” behavior [8]. Nevertheless, over the next
twenty thousand years after first landfall in Australia, the descendants of these colonists
produced a record with very few indices of ‘modernity’, very different from their
contemporaries in western Europe, with for instance almost no evidence of complex lithic
and organic technology [6]. The Tasmanian record is particularly relevant in this context
[9, 10]. Tasmanian Aboriginal stone technology is similar to the European Middle
Paleolithic, but lacks its stone points and Levallois technology, and is mainly composed
of multi- and single platform cores, primary flakes, retouched flakes and various sized
scrapers [11]. In contrast to the Neandertal Middle Paleolithic record, there is also no
direct archeological evidence of hafting or the use of resins or mastics for gluing. The
Tasmanian Pleistocene record does not have blades, microliths, hafted bone tools, and
carved bone ornaments, no stone lined-hearths and strict spatial organization of activities,
and there is little if any technological change occurring for at least 25,000 years.
Nonetheless, the first Tasmanians were AMH, as were all occupants of Sahul. What these
records clearly show is that the models used to describe human groups as either ‘archaic’
or ‘modern’ are flawed. There is no single package of modern human behavior [5], with
the variability of the archeological record of AMH being larger than usually
acknowledged. The factors possibly underlying that variability are debated, but
demography is often seen as a contributing one [6].

References

1. McBrearty S, Brooks A (2000) The revolution that wasn't: a new interpretation of

the origin of modern human behavior. J Hum Evol 39:453-563.
 2

2. Belfer-Cohen A, Hovers E (2010) Modernity, Enhanced Working Memory, and

the Middle to Upper Paleolithic Record in the Levant. Curr Anthropol
51(S1):S167-S175.
3. Bar-Yosef O,Wang Y (2012) Paleolithic Archaeology in China. Annu Rev
Anthropol 41: 319-335.
4. Speth JD (2004) News Flash: Negative Evidence Convicts Neanderthals of Gross
Mental Incompetence. World Archaeology 36: 519-526.
5. Habgood PJ, Franklin NR (2008) The revolution that didn't arrive: A review of
Pleistocene Sahul. J Hum Evol 55:187-222.
6. O'Connell JF, Allen J (2007) Pre-LGM Sahul (Pleistocene Australia-New Guinea)
and the archaeology of early modern humans. In: Mellars P, Boyle K, Bar-Yosef
O, Stringer CB editors. Rethinking the Human Revolution. McDonald Institute
for Archaeological Research, Cambridge) pp 395-410.
7. O'Connell JF, Allen J (2012) The Restaurant at the End of the Universe:
Modelling the Colonisation of Sahul Australian Archaeology 74:5-16.
8. Noble W, Davidson I (1996) Human Evolution, Language and Mind.Cambridge
University Press, Cambridge.
9. Cosgrove R, Pike-Tay A, Roebroeks W (2014) Tasmanian archaeology and
reflections on modern human behaviour. In: Dennel R, Porr M editors. Southern
Asia, Australia and the Search for Human Origins. Cambridge University Press,
Cambridge, pp 175-188.
10. Holdaway S, Cosgrove R (1997) The archaeological attributes of behaviour:
difference or variability? Endeavour 21(2):66-71.
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markers: change, evolution and complexity. A.I.A.S., Canberra, pp 189-204.
 1

Neandertal Demise: An Archaeological Analysis of the Modern Human

Superiority Complex

Paola Villa, Wil Roebroeks

Supporting Information
Text S3. Lissoirs

Archaeologists distinguish bone tools made by percussion flaking, i.e. the same
techniques used in knapping stone artifacts, and so-called “formal” bone tools made with
techniques specifically conceived for bone and ivory materials, such as splitting and
wedging, scraping, grinding and polishing. Bifaces made on elephant bone, such as those
found at the Italian Lower Paleolithic sites Castel di Guido and Fontana Ranuccio [1], are
examples of flaked bone tools. Bone and antler projectile points common in Upper
Paleolithic sites are examples of formal bone tools.
Lissoirs (a French term meaning “smoothers”) are tools made on ungulate ribs,
longitudinally split to produce two thin half ribs. These half ribs are then shaped by
grinding and scraping, with a rounded end polished by use, showing wear facets and
striations. By their similarity to ethnographic bone tools used by the Sami people (Lapps,
indigenous people of northern Europe) they are interpreted as tools for smoothing dry
hides [2]. They are common in Aurignacian assemblages; for instance, 117 lissoirs are
reported from three Aurignacian sites in Southwest France (Castanet, Brassempouy and
Gatzarria [2]). A few have also been reported in the Protoaurignacian (five in level VII)
of Grotte du Renne at Arcy [3]. Twelve “burnishing tools” have been reported in the
Châtelperronian level X of Grotte du Renne [4] although by their description only few
fall into the category of lissoirs [5]. Bone spatulas (with a wider end, common in the
Magdalenian) and thicker bone tools made on reindeer antler are often lumped together
with lissoirs but the bone tools discussed here are all lissoirs in a strict sense.
The four lissoirs described by Soressi et al. [6] come from two Mousterian of Acheulian
Tradition sites, Pech de l’Azė 1 and Abri Peyrony (Southwest France). Three are exactly
like Aurignacian lissoirs; the fourth (from Abri Peyrony) was on an unsplit rib but its
rounded tip was heavily modified and shows spongy bone. All exhibit striations and
polish on their rounded end. The Pech de l’Azė 1 tool comes from layer 4, underlying 3
m of undisturbed Middle Paleolithic deposits and is dated by OSL to 51.4 ± 2.0 ka. The
three Abri Peyrony tools from two superimposed levels are dated by 14C AMS to between
74,710-41,130 cal BP. There are no Upper Paleolithic deposits at both sites. The
interpretation of these tools as used for a specific purpose (smoothing of dry hides) is
based on their wear and on experimental data. Cultural diffusion from Neandertals to
modern humans is a plausible interpretation and supports the idea discussed in the text
that some of the innovative technologies of the Protoaurignacian and Aurignacian may
have developed out of a Middle Paleolithic base [7].
 2

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