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the mid latitude belt from Portugal and the Maghreb to the Caucasus and the
Altai would have presented a large mass of heterogeneous landscape.
The plains of Eurasia would have been, always in the east and during cold/arid
events in the centre and west too, homogeneous in human terms. In the west
the development of forest, forest margins and the presence of lakes during mild
160 Neanderthals and Modern Humans

oceanic phases would have opened up opportunities for ecotonal human eco-
types and reduced them for plains human ecotypes. Morphology and behaviour
would have been major components permitting different human populations
to successfully exploit plains or ecotones. In this book I have suggested that
the long-limbed, gracile, morphology of Moderns, coupled with an appropriate
social and behavioural lifestyle, suited them particularly well to the long-range,
highly mobile, system of the plains (chapters 4 and 5; Finlayson et al., 2000a).
Similarly, the more robust morphology of the Neanderthals would have been
less suited for an open plains existence and the evidence of severe limb wear
would appear to con¬rm this view. Thus, Neanderthals living in the ecotonal
conditions of the heterogeneous landscapes of southern Eurasia and used to ex-
ploiting a range of resources over a small area, would have extended northwards
into the Eurasian Plain when mild conditions induced the spread of the forests
and generated an extension of the ecotonal conditions. It is not surprising, there-
fore, that Neanderthals never colonised the steppe environments of the eastern
European Plain even though they lived close by in the hills and mountains of
Crimea, the Caucasus and the Altai (Soffer, 1994). Similarly, when cold and
arid conditions took hold the range of the Neanderthals receded as the wood-
land of western and central Europe became steppe. It is in edge areas that we
would expect the greatest stress as populations attempted to adapt to the rapidly
changing landscape. These areas would have included south-western France,
the Italian Peninsula, the northern and central Balkans, hilly landscapes in cen-
tral and eastern Europe and sites along the edge of the Russian Plain. It is in
these areas that we would predict the presence of ˜Upper Palaeolithic™ tech-
nologies among Neanderthal groups as they attempted to adapt their tool kit to
the changing circumstances and in the direction of the plains dwellers that were
used to exploiting such environments (Figure 7.3). In such a scenario I would
also predict that the last Neanderthals would have lived close to the topograph-
ically heterogeneous zones. Within these, the populations in edge zones would
have attempted to adapt technologically whereas those in core areas (such as
Iberia) would have maintained a traditional technology to the end.
If my interpretation is correct, then the Neanderthals were a people of mid-
latitude Europe that were able to extend their geographical range northwards
during mild events. They evolved in the rich heterogeneous landscapes of mid-
latitude Europe and their morphology was best suited for the kind of rugged ter-
rain and close-quarter hunting that the landscape demanded (chapters 4 and 5).
As with many other animals, attributes of exploitation of such landscapes would
have included small home ranges, diverse diet at the annual scale as different
resources were seasonally cropped, small population units that were in constant
contact as they moved across the home range, precocious children that would
be able to move with the adults at an early age and an intimate knowledge of the
Modern Human colonisation and Neanderthal extinction 161

home range and the seasons (Chapter 5). The down side of such a strategy would
have been increased likelihood of fragmentation and isolation with consequent
genetic effects.
The Moderns most probably entered the Eurasian Plain somewhere in the
region between the Black and the Caspian Seas. Whichever way it was, by 40 kyr
we see the expansion of the geographical range of this form. The rapid expansion
shows the characteristics of an ecological release and the ¬‚at landscapes of the
Eurasian Plain undoubtedly played a catalytic role as they would persistently
throughout history (for example for the huns or the avars). The nature and
distribution of resources determines home-range size. The exploitation of the
plains required large home ranges and a distinct social system and probably a
greater within-group division of labour, centralised base camps and systems of
storage that would only be possible if such bases existed (Chapter 5). Life in
the plains would have been demanding, not least being the reduction in winter
daylength and the great reduction in resource range compared to tropical and
mid-latitude areas. As group components were separated for periods of time,
there would have been a greater pressure for the development of sophisticated
communication and social-binding systems so it is perhaps not too surprising
to ¬nd so much art and ornamentation in these groups.
In the ¬nal analysis there is therefore very little difference between Nean-
derthals and Moderns. They exploited the same range of food resources and had
similar technological abilities. Observed differences re¬‚ect population adapta-
tion and there are no linear, directional, trends. There is no clear Modern“
Neanderthal boundary that cannot be explained by differences in ecological
setting. Moderns differed from Neanderthals in adaptations (morphological
and behavioural) that enabled them to operate at larger spatial scales. The high
frequency of climatic oscillations and the trend towards cooling towards the
end of OIS 3 introduced environmental instability (Chapter 6). The exploita-
tion of heterogeneous landscapes, as we have seen, was the Neanderthal way of
dealing with short-term instability. Prolonged instability meant that the scale of
Neanderthal response did not match the scale of the perturbations. The Mod-
erns, on the other hand, could deal with such large-scale instability because
they operated on larger scales. The expansion of favoured open, homogeneous,
landscapes and their associated faunas, further enhanced their probability of

Glacial refugia

The inescapable consequence of the climatic ¬‚uctuations of the Pleistocene for
many animals and plants were the huge changes in geographical distribution
162 Neanderthals and Modern Humans

(Hewitt, 1996, 1999, 2000). Some species were able to maintain themselves
in southern European refugia for a number of glaciations while others have
arrived more recently. Extinction would have been a feature of the glaciations
even in southern refugia (Hewitt, 2000; O™Regan et al., 2002). O™Reagan et al.
(2002) have highlighted the importance of southern refugia in the extinction
process of large carnivores, with chance playing a major role in the survival of
the reduced and isolated populations. Such was the case of the Neanderthals,
probably originating from a recent European arrival (c. 500 kyr) and managing
to survive several glaciations in southern refugia before ¬nally becoming extinct
just before the Last Glacial Maximum (LGM).
An increasing number of studies are clarifying the generalised responses of
European populations of many organisms to these climatic ¬‚uctuations. Taberlet
et al. (1998) and Hewitt (1999, 2000) have summarised the patterns. The Balkan
Peninsula was a refuge that acted as the source for recolonisation by all species in
the east and also many in the west. Turkey and the Black Sea“Caspian Sea region
also appear as refugia. Italian populations, on the other hand, rarely repopulated
Europe, the Alps apparently acting as a signi¬cant barrier. The Pyrenees were
also a barrier to populations dispersing from Iberia but it seems that they were
more porous than the Alps. Finally, there is evidence of isolated patches further
north, along the southern edge of the steppe“tundra zone, which acted as local
refugia (Willis et al., 2000, 2001; Carcaillet & Vernet, 2001; Stewart & Lister,
2001). I suggest that the Balkans refugium, always being more continental in
characteristics than the oceanic Iberian refugium, may have additionally held
populations that were physiologically better able to expand into temperate areas
in the initial stages of a deglaciation. This may explain the importance of this
refugium for temperate trees (Chapter 6; Bennett et al., 1991).

The Iberian refugium

In this section and the next I use the Iberian Peninsula as a model for the study of
human dispersion and dispersal during the Quaternary. Iberia is diverse and large
enough to act as a natural laboratory for the study of human interactions in the
Pleistocene. The southern Iberian Peninsula has been occupied by humans since
at least 500 kyr but probably signi¬cantly earlier. I proposed in Chapter 3 that
the hominids that were the ancestors of those inhabiting Atapuerca over 780 kyr
(Bermudez de Castro et al., 1997) may have reached Europe across the Strait
of Gibraltar and there is also a claim of hominid occupation in Orce (Granada)
at 1.2 Myr (Oms et al., 2000) which must await further evidence. The southern
Iberian Peninsula has been a crucial region throughout this period, acting as a
refugium for human populations during glaciations (Finlayson, 1999; Finlayson
Modern Human colonisation and Neanderthal extinction 163

et al., 2000a; Straus, 2000), being one of the areas of late Neanderthal survival
(Vega-Toscano, 1990; Finlayson, 1999). Giles Pacheco et al. (2003) examined
the distribution of humans in southern Iberia (Andaluc´a and Gibraltar) after
500 kyr based on an inventory of archaeological and palaeontological sites.
They analysed these data against climate for the period 90“0 kyr (GRIP, 1993)
at the scale of 0.5 kyr to test the relationships between climate parameters and
human distribution.
Giles Pacheco et al. (2003) surveyed the literature to identify sites that be-
longed to distinct archaeological periods (hereafter referred to as cultures) in
southern Iberia. The following divisions were established.

Late Acheulian (Mode 2/3)
The Acheulian was established in Europe by 500 kyr (Foley & Lahr, 1997).
The data used by Giles Pacheco et al. (2003) represented the late Acheulian
which is characterised by the standardisation of the use of ¬‚int and a generalised
introduction of Mode 3 (Giles Pacheco et al., 1993, 2003), and was represented
by sites leading up to the last interglacial.

Mousterian (Mode 3)
The Mousterian appeared in Europe by 250 kyr (Foley & Lahr, 1997). It is
characterised by the use of the Levallois method of extraction (Klein, 1999)
and by a homogenisation of the use of ¬‚int and the standardisation of types.
It was represented by sites that date from before the last interglacial to 31 kyr
(Finlayson & Giles Pacheco, 2000).

Aurignacian (Mode 4)
The Aurignacian, generally associated with Modern Humans, appeared in
Europe around 45 kyr (Bar-Yosef, 2000). It reached northern Spain by 40 kyr
(Straus & Winegardner, 2000) and is very rare in the south to the point that
Finlayson et al. (2000b) have questioned its signi¬cance there. The use of bone,
the manufacture of blades and the appearance of parietal art are characteristics
of the Aurignacian (Klein, 1999).

Gravettian (Mode 4)
The Gravettian is found in Iberia from 29 kyr (Marks, 2000). It is charac-
terised by the presence of backed elements, abundant burins and the absence of
Aurignacian-type thick endscrapers, Dufour bladelets or bone points (Straus &
Winegardner, 2000).
164 Neanderthals and Modern Humans

Solutrean (Mode 4)
The Solutrean in Iberia spans the period 20.5“16.5 kyr (Straus & Winegardner,
2000). The technology is distinctive with bifacial techniques with concave base
and rhomboidal forms, the appearance of peduncular points, an increase in bone
technology (Aura Tortosa, 1989; Ripoll L´ pez & Cacho Quesada, 1990; Villa-
verde & Fullola, 1990) and an explosion of parietal art (Fortea P©rez, 1978).

Magdalenian (Mode 4)
The Magdalenian, which spans the period 16.5“11 kyr, is highly diverse and in-
cludes bone implements, a reduction in tool size and the appearance of portable
art. Parietal art reaches its peak (Aura, 1989; Straus & Winegardner, 2000).

Epipalaeolithic (Mode 5)
The Epipalaeolithic commences around 11“9 kyr (Straus & Winegardner, 2000)
and the last populations are indenti¬able to around 6.3 kyr (Oliver & Juan-
Cabanilles, 2000). The characteristic innovation is the geometric microlith
(Fortea, 1973).

Early and Middle Neolithic
The ¬rst two Neolithic divisions were considered by Giles Pacheco et al. (2003).
The Neolithic reached Iberia around 6.5 kyr or 5.4 Cal bc (Zilhao, 2001). It
marked the ¬rst presence of ceramic with cardial patterning. The Middle Ne-
olithic, with epicardial ceramics, commenced around 5.7 kyr or 4.5 Cal bc
(Oliver & Juan-Cabanilles, 2000).
The number of sites within each technological period and time frame was
converted to site density by dividing the number of sites by the time span of the
technology and multiplying by 1000, thus representing them as sites/millennium
(Straus & Winegardner, 2000). Climate data used were for the period 90“0 kyr
(GRIP, 1993) at intervals of 0.5 kyr. The parameters used were: mean δ 18 O (0 /oo )
that is an indicator of temperature; and the coef¬cient of variation (Sokal &
Rohlf, 1981) of δ 18 O. For the analysis of mean δ 18 O and coef¬cient of variation
of δ 18 O, 100 randomly selected samples of n = 5 from each period were iterated.
By bootstrapping Giles Pacheco et al. (2003) attempted to remove sampling
effects related to the difference in duration of each cultural period.
Site density increased from the Acheulian to the Neolithic, with the main
increase in the Holocene, peaking in the Middle Neolithic, but with a notable
increase also during the Solutrean which was signi¬cantly higher than predicted
by the model (Figure 7.4). The duration of each culture decreased through time
and was especially evident in the Upper Palaeolithic (Figure 7.5). Temperature
(mean δ 18 O/0.5 kyr) decreased gradually prior to 20 kyr and then increased after
the LGM (Figure 7.6a). Climate variability (coef¬cient of variation) decreased
Modern Human colonisation and Neanderthal extinction 165


In Sites/Millennium




’375 ’125 ’33.5 ’23.75 ’18.5 ’13.5 ’8.5 ’6.1 ’5.4
Time Mode (kyr)

Figure 7.4. Change in density of sites (log n sites/millennium) through time (bars).
Curve shows best model ¬t. The relationship is highly statistically signi¬cant (R2 =
0.825; P<0.001) and is best described by an S regression model (ln(y) = ’2.0279 +
(’37.975/x)). White bars, Middle Palaeolithic (Acheulian, Mousterian); grey bars,
Upper Palaeolithic (Aurignacian, Gravettian, Solutrean, Magdalenian, Epipalaeolithic,
early Neolithic, advanced Neolithic). After Giles Pacheco et al. (2003).

through time, especially after the LGM (Figure 7.6b). There was a signi¬cant
increase in site density with an increase in temperature (Figure 7.7a) and an
even stronger relationship with climate stability (Figure 7.7b).
Several patterns emerged from the results of Giles Pacheco et al.™s (2003)
analysis. There was a trend for site density to increase from the Acheulian to
the Neolithic and the rate of increase was greater in the Holocene starting in the
Epipalaeolithic (Figure 7.4). There was also a peak during the Solutrean, that
had previously been recorded regionally (Finlayson & Giles Pacheco, 2000)
and in other parts of Iberia (Straus & Winegardener, 2000). Giles Pacheco et al.
(2003) interpreted these results as follows: conditions in southern Iberia during
the Acheulian and Mousterian and the methods that humans used for exploiting
the landscape were such that, at the scale observed, there was very little change
during this period. Assuming that site density was in some way proportional to


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