Quaternary International 247 (2012) 230e235
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Quaternary International
journal homepage: www.elsevier.com/locate/quaint
Spatial abilities, cognition and the pattern of Neanderthal and modern human
dispersals
Ariane Burke
Universite de Montreal, Anthropologie, CP 6128, Centre-Ville, Montreal, Quebec, Canada H3C 3J7
a r t i c l e i n f o
a b s t r a c t
Article history:
Available online 9 November 2010
Neanderthals and anatomically modern humans (AMHs) are very closely related, although their skeletal
morphology is distinct enough for them to be classified as separate species. Their physical differences,
including the architecture of the skull and post-cranial skeletal proportions, do not explain the extinction
of the Neanderthals as a species, however. The post-cranial morphology of the Neanderthals, which is
generally believed to reflect an active lifestyle under cold conditions, falls within the range of modern
human variation. Their cranial capacities are essentially identical and skull morphology does not convey
any obvious adaptive advantage to either species. The existence of other, less easily discernible biological
traits that might have contributed to the extinction of the Neanderthals e such as differences in
reproductive rates or developmental histories e cannot be ruled out, but are not supported by current
data. Behaviourally (culturally) based explanations for the process of replacement of Neanderthal populations by AMHs center on archaeological evidence for more complex patterns of social organisation,
associated with the use of symbols and the development of new technologies. This paper explores the
link between social complexity, spatial distribution and cognition during the Late Pleistocene, suggesting
a mechanism whereby cognitive differences between hominid species may have arisen.
Ó 2010 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
Modern human dispersal into Eurasia and the subsequent
extinction of Neanderthals during the Late Pleistocene are two of the
most fascinating and enduring problems in prehistory. Various
explanations have been suggested over the years, mostly centered on
attempts to identify an adaptive advantage for modern humans,
whether biological or cultural, that would link the two processes.
Paleontological and archaeological data, including the spatial distribution of archaeological sites, evidence for the production of material
culture and subsistence strategies all come into play in attempts to
resolve this puzzle. This paper focuses on the development of
spatially extensive social networks and heightened mobility, associated with the adoption of a modern huntingegathering way of life,
as a factor stimulating the development of spatial abilities in modern
humans that may have resulted in the emergence of differences in
spatial cognition.
2. Biological explanations
Skeletal indicators of an adaptation to cold climates (Howell,
1952; Trinkaus, 1981; Trinkaus et al., 1999) were originally considered significant insofar as they corroborated the hypothesis that
E-mail address: [email protected].
1040-6182/$ e see front matter Ó 2010 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2010.10.029
Neanderthals evolved from a geographically isolated, Western
European hominin population during the course of the Late Pleistocene. The brachial and crural indices of Neanderthals are considered an adaptation to cold temperatures following Allen’s rule
(Howell, 1952) e in contradistinction to the more “tropical” body
proportions of AMHs e and their capacious nasal cavities are also
thought to represent an adaptation to the cold, although in both
cases the observed traits fall within modern human variation
(Trinkaus, 1981). Their post-cranial morphology does not explain the
disappearance of Neanderthals at the end of the last Glacial e a time
when a physical adaptation to cold conditions should have been
advantageous, however.
Another biological explanation for adaptive differences between
AMHs and Neanderthals is genetic. The role of genetic difference in
this debate is difficult to assess, however. The FOXP2 gene, which
makes human speech possible, was once thought to differentiate
between AMHs and Neanderthals (Klein, 2000; Enard et al., 2002),
but is now known to exist in the Neanderthal genome (Krause et al.,
2007a; Trinkaus, 2007).
Cognitive differences between AMHs and Neanderthals have also
been invoked, despite the difficulties involved in constructing
bridging arguments between cognition and the archaeological record
(Wynn, 2003). The distribution of archaeological sites and the organisation of their settlement systems led Binford (1989) to suggest that
AMHs were able to plan logistically, whereas Neanderthals lacked
A. Burke / Quaternary International 247 (2012) 230e235
tactical depth, moving from site to site as required, pursuing an
opportunistic strategy of resource procurement. Archaeozoological
evidence suggests that Neanderthals were not limited to opportunistic resource procurement strategies, however. On the contrary,
Neanderthals were clearly capable of targeting gregarious prey
species, anticipating their movements and making efficient use of the
landscape to hunt them e a strategy that requires planning depth
(Farizy and David, 1989; Farizy et al., 1994; David & Enloe 1993;
Gaudzinski, 1996; Gaudzinski & Roebroeks 2000; Burke, 2000;
Patou-Mathis and Chabai, 2003). Based on the faunal evidence,
therefore, there is little reason to suggest that Neanderthals lacked an
innate (cognitive) inability to plan ahead and think tactically.
3. Archaeological explanations
Attempts to identify a culturally based adaptive advantage for
AMHs in the archaeological record are reviewed in full elsewhere
(Bar-Yosef, 2002) and are only briefly summarized here. Most of the
technological advantages once thought to distinguish the Upper
Palaeolithic (UP) are now known to have appeared earlier in the
archaeological record. The production of blades, for example, is now
recognized as a feature of both AMH and Neanderthal lithic industries at different times during the MP/MSA (Bar Yosef and Kuhn,
1999; Meignen, 1994). The invention of hafting, thought to signal
the relative sophistication of UP tool making abilities, is now
recognized in both the MP and the MSA (Boeda et al., 1996; Kay,
1998; Hardy et al., 2001; Lombard, 2005; Wadley, 2005; Wadley
et al., 2009) and a developed bone tool industry is incontrovertibly part of the MSA (e.g. Henshilwood et al. 2002; d’Errico and
Henshilwood, 2007; Backwell et al., 2008). If relative technological
sophistication is an indicator of cognitive abilities, therefore, there is
not much to distinguish between AMHs and Neanderthals e at least
during the timeframe of interest.
In terms of subsistence strategies, Neanderthals scavenged or
hunted large game when and where appropriate (Speth and
Tchernov, 2001) and exploited small game and shellfish (Stiner,
1994, 2001; Barton et al., 1999; Stringer et al., 2008) as well as
plants and nuts (Barton et al., 1999; Albert et al., 2000). Examples of
specialised hunting, once thought to distinguish UP subsistence
strategies, have been demonstrated in a number of Middle Palaeolithic contexts (David and Enloe, 1993; Gaudzinski, 1996; Grayson
and Delpech, 2002; Costamagno et al., 2006). The distinction
between AMHs and Neanderthals is once again lost as it appears that
both species shared a range of subsistence strategies and exploited
a wide range of resources, adjusting to local conditions (Bar-Yosef,
2004). Neither the paleontological nor the archaeological records,
therefore, yield convincing evidence for the existence of a clear
adaptive advantage that would explain the successful dispersal of
AMHs and the eventual extinction of the Neanderthals.
4. Spatial patterning
Spatial patterning, on the other hand, hints at significant differences between Neanderthals and AMHs. The spatial distribution of
a species provides information about its ecological requirements,
environmental limitations and adaptive flexibility; the core range
reflects the environment within which a species initially evolved
through a process of adaptation, while the effective range reflects
both innate adaptive flexibility and the outcome of interactions with
other species (prey and competitors). Lamentably, the total extent of
hominin ranges during the Late Pleistocene cannot be assessed,
because ancient coastlines, now submerged, cannot be surveyed
without considerable effort (e.g., Bailey et al., 2008). In addition, the
reliability of Late Pleistocene time series for the critical time period
between ca. 32 and 35 ka 14C is questionable, despite recent
231
advances in radiocarbon calibration (Weninger and Joris, 2008).
These and other confounding factors make reconstructing the
demographic patterns of humans (AMHs) and Neanderthals a bit of
a black art. Nevertheless, it is clear that the initial ranges of these
species differed and that spatial separation was maintained for
a relatively long time.
Western Europe is considered to be the “classic” (or “core”)
Neanderthal range based on the distribution and relative age of
known archaeological sites (Bocquet-Appel and Demars, 2000) and
the distribution of dated skeletal remains (Serangeli and Bolus,
2008). The total geographic range of Neanderthals is known to
extend as far south as the Levant, and as far east as the Altai (Krause
et al., 2007b). Neanderthals, therefore, were capable of living in
a variety of environments under different climatic conditions e
whether they thrived in all of them is questionable, however, and the
implications of this observation will be discussed below.
On a regional scale, Neanderthals are generally considered to
have moved frequently within relatively small territories, intensively using local resources (Mellars, 1996), although they were
capable of exercising variable degrees of mobility according to the
structure of the environment (Davies, 2007). The observation that
raw material transfer distances in Europe during the MP are
regionally variable and correlate broadly with landscape variability
supports this suggestion (Féblot-Augustins, 1993). Nevertheless,
Neanderthals generally appear to have mapped onto relatively
smaller territories than AMHs, e.g. (Meignen and Brugal, 2001; Adler
et al., 2006; Conard et al., 2006).
On a continental scale, MP sites are relatively under-represented
in the archaeological record of the East European Plain (Vishnyatsky
and Nehoroshev, 2004) where they are distributed almost exclusively in the south (Hoffecker, 1999; Pavlov et al., 2004). In contrast,
not only is the Upper Palaeolithic record of the East European Plain
relatively richer, it also suggests a pattern of rapid colonization
(Dolukhanov, 1999). Accepting that Neanderthals were morphologically adapted to cold conditions, capable of existing in a range of
distinct landscapes, technologically advanced and able to prey upon
large-bodied, gregarious herbivores, then the northern plains of
Europe should not have represented an insurmountable geographical challenge. The fact that AMHs were apparently better equipped
for life in the Eastern European plains than Neanderthals indicates
that other aspects of their adaptation have to be considered.
One explanation for the observed spatial patterns is the new
pattern of social organisation that emerged during the MSA
(Gamble, 1991; McBrearty and Brooks, 2000; Henshilwood et al.,
2002; Wadley et al., 2009). The development of a “modern” way of
life is indicated by the presence of tools for the communication of
social identity and the expression and maintenance of personal
relationships in absentia and over long distances (Gamble, 1998).
The archaeological and ethnographic records suggest that social
identities are constructed and maintained through the production
of material culture accompanied by stylistic variation (Weissner,
1984) and symbolic expression. Early manifestations of symbolic
expression through the production of material culture are documented for the MSA by McBrearty and Brooks (2000) and others
(d’Errico et al., 2005; Bouzouggar et al., 2007; d’Errico and
Henshilwood, 2007) as well as for the MP of the Levant (Hovers
et al., 2003; Bar-Yosef Mayer et al., 2009) associated with AMHs.
Symbolic behavior is not absent in Neanderthal contexts (Langley
et al., 2008; Zilhão et al., 2010) but is a rare occurrence before the
arrival of UP industries in Europe. The more “modern” aspects of the
material culture of Homo sapiens identified by McBrearty and Brooks
(2000), interpreted as signs of a gradual process of cultural intensification, are not uniformly present during the MSA which leads
the authors to suggest a stepwise, sometimes intermittent process;
this process may be linked to population pressure in regional
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A. Burke / Quaternary International 247 (2012) 230e235
populations (Henshilwood and Marean, 2003). The fact that relatively “late” Neanderthal cultures, such as the Chatelperonian, hint
at the emergence of similar cultural developments in Western
Europe could be seen as validating this hypothesis, since the
contraction of Neanderthal populations into their core ranges
towards the end of MIS 3 (Bocquet-Appel and Demars, 2000;
Serangeli and Bolus, 2008) could have resulted in population pressure through demographic packing. This implies the existence of
similar adaptations to social and demographic pressures, and social
cognition, in both species but different timing for the triggering of
the process (earlier for AMHs, later e possibly too late e for
Neanderthals).
The cultural manifestations identified in the MSA (and the Levant)
therefore, are harbingers of a modern pattern of social organisation
that would fully emerge during the Upper Palaeolithic, enabling the
emergence of socially integrated regional groups (Conkey, 1980,
1984; Gamble, 2000). Another result of this development is that
local groups could counter resource stress by redistributing members
within a wider regional population (Weissner, 1982; Cashdan, 1985).
Information (crucial to fine-tuning movements to coincide with
resource availability) would also have flowed more efficiently as
individuals moved between groups (Whallon, 2006). The flow of
information and people on a regional scale would have maintained
equilibrium between resource availability and local group size within
the population, facilitating adaptation to challenging environments.
The relative success of AMH dispersals, particularly in difficult
environments such as the Northern Plains of Eurasia (Hoffecker and
Cleghorn, 2000), may therefore be explained. While AMHs would
develop and maintain spatially extensive, well-integrated social
networks during the course of the MSA, culminating in wide-spread
dispersals, Neanderthals, by and large, retained a more local pattern
of social interaction.
5. Spatial abilities and spatial cognition
The development of a modern hunting and gathering way of life
could have conferred an adaptive advantage on dispersing AMH
populations for the reasons explained above. It may also have
fostered the development of cognitive differences between AMHs
and Neanderthals. The challenge of maintaining complex, spatially
extensive social networks, implying heightened mobility and
spatial awareness, requires the deployment of specific cognitive
skills related to wayfinding, stimulating the development of spatial
abilities that could eventually lead to changes in spatial cognition.
Golledge (1999:47) defines wayfinding as: “purposeful movement to a specific destination that is distal and thus cannot be
perceived by the traveller”. Two distinct wayfinding strategies are
used by humans. The first strategy (an “allocentric” or “survey
strategy”) uses Euclidian metrics (direction and distance estimates)
and an “enduring [.] objecteobject or environmenteobject association” (also known as a “cognitive map” or “cognitive representation” (Golledge, 1999, 2003; Kitchin, 1994)). The second strategy
is an “egocentric” or “landmark strategy” that relies on landmark
recognition (or “transient action-oriented egocentric self-object
associations” (Burgess, 2006). The process of linking the two
representations (aka “spatial updating”) ensures that humans can
recognise their current position and translate it onto the enduring
representation. The internal representations (cognitive maps) are
built up from direct experience of the world through a process of
path integration (Golledge, 1999; Loomis et al., 1999) and landmark
recognition (Foo et al., 2005), implying the motivated and deliberate encoding of environmental information (Golledge, 2003:30).
Allocentric and egocentric wayfinding strategies make use of
distinct spatial skills and correlate with the development of different
spatial abilities in humans (Shelton and Gabrieli, 2004). For
example, the link between preference for a “survey” strategy and
good mental rotation skills is thought to result from differential
stimulation of the hippocampus (part of the brain implicated in
spatial cognition and in the performance of mental rotation) during
allocentric encoding (Burgess et al., 2006; Parslow et al., 2004). This
is supported by the observation that people who have a high
dependence on their wayfinding skills and habitually use maps
develop relatively enlarged posterior hippocampi (Maguire et al.,
2000). Preferential use of a particular wayfinding strategy, therefore, leads to the development of, or alternatively reinforces existing
differences in spatial ability and may result in concomitant changes
to the brain.
Ranging patterns (mobility) are linked to sex-based preferences
for one or the other wayfinding strategy in some species of
mammal and correlate with differences in spatial abilities (Gaulin
and Fitzgerald, 1989). In humans, experimental psychologists
have noted differences between the sexes in the performance of
tests designed to measure spatial abilities that could reflect preferences in the selection of a wayfinding strategy. For example, men
typically out-perform women in tests of spatial perception and
mental rotation (Montello et al., 1999; Voyer et al., 1995), whilst
women are better than men at memorizing spatial configurations
(Barnfield, 1999; Tottenham et al., 2003; Levy et al., 2005). These
differences are thought to correlate with use of survey strategies in
men and landmark-based strategies in women.
Evolutionary psychologists suggest that differential spatial abilities of men and women reflect differences in spatial cognition that
evolved through time as a result of the adoption of a gendered
division of labour in hunting and gathering societies and with it,
gendered patterns of mobility (Eals and Silverman, 1994; Silverman
et al., 2000, 2007; Ecuyer-Dab and Robert, 2004). Men are generally
considered to range further than women in hunting and gathering
populations, and since allocentric representations of space work best
when distances are great (Burgess, 2006) proponents of the “Hunteregatherer theory of spatial sex differences” (HGT) suggest that
hunters develop a preference for a “survey based” or “allocentric”
strategy. This preference would explain a male advantage in mental
rotation tests. Conversely, the smaller-scale mobility assumed to be
correlated with foraging activities is proposed as the context within
which spatial memory is selected for in women, explaining the
female advantage in object-location memory tests (Ecuyer-Dab and
Robert, 2007).
The HGT, therefore, suggests that differences between the
spatial abilities of men and women reflect sex-based differences in
spatial cognition that evolved as a result of the adoption of
gendered activity patterns. It is debatable whether observed
differences in spatial abilities in men and women reflect sex-based
differences in spatial cognition, or whether they are the result of
a gender-based training effect promoting the differential development of spatial abilities. Feng et al. (2007) show that individuals can
be trained to perform equally well in mental rotation tests, irrespective of their sex. Similarly, a recent study of orienteering
suggests that trained participants of both sexes perform navigation
tasks with the same efficiency (Burke et al., nd). The existence of
a training effect is supported by evidence that length of exposure to
complex wayfinding tasks requiring the use of an allocentric
perspective is positively correlated with physical changes to the
brain (Maguire et al., 2003). In other words, the plasticity of the
human brain allows significant differences in spatial abilities (and
neuroanatomical changes) to develop when a subject is provided
with training opportunities. Regardless of whether or not sexbased differences in spatial abilities exist as a result of training or
innate, cognitive differences, there is clear evidence for a link
between their development and habitual patterns of activity in
humans.
A. Burke / Quaternary International 247 (2012) 230e235
The link between patterns of mobility and the development of
spatial abilities allows for the suggestion that when AMHs developed the ability to maintain spatially extensive social networks and
adopted a more mobile lifestyle (i.e., during the course of the MSA)
they created conditions under which specific spatial abilities were
selected for, thus placing new demands upon the neural substrate
(Maguire et al., 2003; Hartley et al., 2007). Furthermore, exploratory travel, such as would have occurred during the initial phases
of dispersal, makes very different demands upon the brain than
travelling a known route (Hartley et al., 2003). The effort required
to maintain relatedness over greater distances and the skills
deployed during exploratory travel together would have promoted
the use of allocentric wayfinding strategies, which work best in
situations where distances are great and more locations have to be
memorized (Burgess, 2006). Put another way, modern human
dispersals were facilitated by the development of a more generalised, transferable system for learning a landscape, rather than
a reliance on detailed local knowledge (Kelly, 2003). Dispersing
populations of AMHs would have been drawing upon different
cognitive resources than Neanderthals, therefore, given the spatial
patterning in the archaeological record discussed above; eventually, this could have resulted in physical changes to the brain
(Maguire et al., 2003). This process would have cycled through
generations, reinforcing itself over time and widening the cognitive
gap between AMHs and Neanderthals. The pace of change would
have been similar to that proposed in the accretion model of the
development of fully modern culture during the MSA proposed by
several authors (e.g., McBrearty and Brooks, 2000; Clark, 2002).
What is proposed here, therefore, is a cultural mechanism for the
emergence of differences in spatial abilities, possibly accompanied
by structural changes to the brain, which would have enabled the
emergence of differences in spatial cognition between AMHs and
Neanderthals. It is possible to make predictions about the archaeological record on the basis of this hypothesis. The development of
a “modern” hunting and gathering adaptation including spatially
extensive social networks and more mobility will be correlated with
the adoption of allocentric wayfinding strategies which enable rapid
movement and exploratory travel over greater distances and in turn,
will have facilitated rapid population dispersal. Information about
novel landscapes will have been more efficiently relayed through
spatially extensive social networks and would also have facilitated
dispersal. These developments would have made AMHs better able
to colonise vast, relatively undifferentiated spaces which present
a challenge to wayfinding strategies requiring detailed local
knowledge and landmarks being less “legible”, sensu Golledge
(1999). The pace of hominid dispersals is difficult to measure
given the relative paucity of sites attributable to the early UP, their
spatial distribution and the error associated with dating methods.
Nevertheless, looking at the distribution of archaeological sites in
the East European plain suggests that AMHs were the first to really
penetrate the plains (Hoffecker, 2002; Pavlov et al., 2004) and that
colonization was probably a relatively rapid event (Dolukhanov,
1999). In contrast, the Neanderthals thrived on the fringes of the
plains in more topographically complex regions, better-suited to
landmark-based wayfinding strategies, such as the Crimean Peninsula (Burke, 2006). The pattern of site distribution in the East
European Plain suggests that the prediction made above is supported by the archaeological record.
It is also possible to predict that along with an ability to maintain
connectedness over greater distances, humans developed a more
efficient means of communicating information about the landscape.
One of the most efficient ways of communicating about the landscape is via schematic representations, i.e., maps. Mapping abilities
are a universal aspect of modern human spatial cognition (Blaut
et al., 2003). It is hard to say when schematic representations of
233
the landscape first developed but the archaeological record suggest
that the first such representations of the landscape appear during the
Upper Palaeolithic. The existence of symbolic expressions interpretable as attempts to schematize the landscape all date to the UP
(Kozlowski, 1992; Svoboda, 2007; Utrilla et al., 2009). The existence
of landscape representations, therefore, could signal the culmination
of the process of evolution of modern human spatial cognition
during the Upper Palaeolithic.
6. Conclusion
It is likely that the extinction of the Neanderthals was a complex
process involving climatological, biological, cognitive and social
factors. Here, we have concentrated on the link between spatial
abilities and changes in social organisation and consequently,
mobility, that emerged during the MSA. The development of tools
for the symbolic expression of social identity, which in turn allowed
humans to create large social networks and maintain relatedness
over great distances, is linked to the emergence of heightened
mobility just prior to the appearance of Upper Palaeolithic cultures
in the archaeological record. These factors combine to create
a context for the differential development of enhanced spatial
abilities in AMHs. In contrast, the less spatially extensive social
systems of the Neanderthals did not make the same demands upon
their cognitive systems. Enhanced spatial abilities in AMHs set the
scene for the evolution of differences in spatial cognition between
AMHs and Neanderthals, The emergence of a preference for allocentric wayfinding strategies which accompanied this process
would explain evidence for rapid human dispersals across new
landscapes, including the northern plains of Eurasia. Differences in
spatial cognition leading to the successful colonization of new
regions could have played a role in the relative evolutionary success
of AMHs at the expense of the Neanderthals.
Acknowledgements
The author would like to thank the Institut Català de Paleoecologia Humana I Evolució Social and the organisers of the
workshop “Neanderthal home: spatial and social behaviours” for
a stimulating series of discussions and the many interesting questions that greatly assisted her in the preparation of this manuscript.
Discussions with Dr. J. Steele and colleagues at the CECD (Institute
of Archaeology, University College London) formed the basis for
initial research on this topic.
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