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at intermediate and high levels of availability respectively and both actively
selected. Both probably occurred at low availability in the plains, wild boar
being selected at the rate of encounter and chamois being actively selected.



Intermediate species

These were rhinoceros and aurochs. Gamble (1995) does not differentiate be-
tween rhinoceros species. If he had, differences between Plains species and
those of more vegetated habitats may have emerged. Rhinoceros and aurochs
occurred at intermediate levels of availability on the plains and the MLB.
Rhinoceros were actively selected on the Plains and selected as encountered in
the MLB. The pattern was reversed for aurochs. The dataset is incomplete for
the roe deer but we may tentatively place it as an intermediate species occurring
at intermediate levels of availability on the plains and the MLB and selected as
encountered in the two regions.
Humans therefore appear to follow particular prey selection strategies that
we may summarise as follows:

(1) There appears to be a greater specialisation in the plains where most species
are actively selected. In the MLB many more species appear to be se-
lected as they are encountered. This difference may explain claims that
Neanderthals hunted prey as it was encountered and Moderns by planned
searching of particular prey species. We can see how the predominance of
Neanderthal sites in the MLB and Moderns sites on the plains can lead to
this apparent pattern.
(2) A number of prey species are actively selected in situations in which they
occur at high density. On the plains we have the three main herding species:
mammoth, horse and reindeer. On the MLB we have the two rocky habitat
herding species: ibex and chamois. There are no cases of prey that occupy
intermediate or closed vegetation in this category.
Comparative behaviour and ecology 103

(3) A number of prey species are actively selected even though they occur at
low levels of availability. On the plains these are typical MLB species:
ibex, chamois and red deer. On the MLB they are typical Plains species:
horse, reindeer and giant deer. Four are herding species: ibex, chamois,
horse and reindeer, that were also entered in (2). Red deer is also a herding
species that would be accessible in open country as would be the case in
the plains. Giant deer may be a similar case.
(4) There are several species that are actively selected but occur at interme-
diate levels of availability. The reasons for their selection may lie in a
combination of the factors described in (2) and (3). On the plains these
species are giant deer, elk and rhinoceros. On the MLB they are aurochs
and wild boar.
(5) The species that are selected at the rate of encounter are species that are
either: (a) rare in marginal geographical areas “ mammoth in MLB and
wild boar on the plains; or (b) species that are dispersed in vegetation and
rarely venture into open vegetation “ roe deer everywhere, aurochs on the
plains and rhinoceros and red deer on the MLB.
(6) The saiga appears anomalous. It is an open plains species that can ag-
gregate and would have occurred at times at intermediate or even high
levels of availability. Two reasons may explain the anomaly. The species
was sporadic in Europe or its small size reduced its appeal to human
hunters.

These results support the view that mammalian herbivore exploitation by
Pleistocene humans was related to ecology and not to the human type. There
are very few obvious shifts in prey exploitation between the Middle and early
Upper Palaeolithic and when they occur, as with the horse, they appear related
to shifting ecological boundaries.
While there may be a case for using ˜overkill™ hypotheses in the case of
colonising human populations, such as those arriving in the plains of Eurasia at
the end of the Pleistocene (and I am not totally convinced), such an argument
would seem to have little value when examining well-established populations of
hunter“gatherers such as the Mediterranean Neanderthals. The most probable
relationship between Neanderthals and their resources would have been one
of density-dependent population regulation and not over-exploitation. In the
absence of ¬ne-grained data showing the contrary this must remain the most
ecologically plausible and parsimonious explanation. Furthermore, the often
rapid climatic oscillations of the Pleistocene in Europe would have generated
continuous range and density shifts in many species that were consumed by
Neanderthals. In such situations of instability abiotic factors would have been
the key to continuously alter prey densities.
104 Neanderthals and Modern Humans

I therefore conclude that there were signi¬cant dietary differences between
peoples (modern or archaic) inhabiting the northern plains (largely mammal-
meat consumers) and those in the heterogenous landscapes to the south (where
they had broader diets) (Table 5.2). Those in the tropics would have had, as they
do today, the greatest available range of foods. It would have been the plains
dwellers that evolved the most sophisticated behavioural and physiological risk
reduction tactics.


Habitat, landscape and geographical range

The only quantitative study of Neanderthal habitat that looked at vegetation
structure as well as species composition was the one that examined the Gibral-
tar Neanderthals (Finlayson & Giles, 2000). In that study I demonstrated that
Neanderthals in Gibraltar hunted in what I described as a Mediterranean wooded
savannah, that is a fairly open vegetation with a mix of shrub and light tree cover
(Figure 5.1). In other words, Neanderthals were exploiting situations that were
of an intermediate structural nature, that is neither dense forest nor fully open
plains. Such ecotones or areas of high habitat heterogeneity are expected to be
high in mammal species richness (Kerr & Packer, 1997). I also showed how a
change towards dense montane forest at the end of OIS 3 corresponded to the
disappearance of the Neanderthals from the area (Figure 5.2). Neanderthals also
exploited other habitats, speci¬cally cliffs and similar rocky areas, estuaries and
coastal habitats. The evidence from other regions shows that, as in Gibraltar,
Neanderthals exploited intermediate habitats between closed forest and open
plains (Soffer, 1994; Mellars, 1996). These habitats would have suited them
well as they would have had a rich grass layer that would have been attrac-
tive to grazers (Finlayson & Giles, 2000). There would have been some cover
for prey to be stalked and the cover would not have been too dense to restrict
hunting and herbivore activity.
Another link between Neanderthals and habitats comes through fresh wa-
ter. The Gibraltar Neanderthals would have had ample supplies of freshwater
close by (Finlayson & Giles, 2000). In the Perigord, south-west France, the
distribution of Neanderthals is close to rivers (Mellars, 1996) and the associ-
ation between Neanderthals and other contemporary humans with lacustrine
and other freshwater habitats seems to be a widespread and trans-continental
phenomenon (Nicholas, 1998). The association would seem to have a dual
advantage: the availability of drinking water and the attraction such habitats
have for other animals and therefore as a source of prey.
The distribution of Moderns in the early stages in Eurasia is associated with
open plains habitats (Soffer, 1985; Finlayson, 1999; Finlayson et al., 2000a).
Table 5.2. (a) Summary of predicted utilisation of food resources by late Pleistocene humans on the Eurasian Plain and the
Eurasian mid-latitude belt. Main resources are in dark grey cells. Important resources are in pale grey. (b) Summary of
predicted habitat use by Late Pleistocene humans
(a)
Mammalian herbivores
1000“500 500“100 Small Marine Marine
>1000 <100
kg kg kg kg mammals mammals Birds Tortoise Fish molluscs Fruit

Eurasian Plain +++ +++ +++ + + ’ ++ ’ ++ ’ +
Mid-latitude Belt + ++ ++ ++ ++ ++ +++ +++ ++ +++ ++

(b)
Closed Intermediate Open Rocky Wetland Coast

Eurasian Plain ’ + +++ + ++ +
Mid-latitude Belt +++ +++ + +++ ++ +++
106 Neanderthals and Modern Humans

This immediately suggests a difference in habitat use between Moderns and
Neanderthals. There is a range of habitats utilised by Moderns that includes
the types used by Neanderthals and all we can conclude, on present evidence,
is that Moderns included open plains as habitats that could be exploited much
more intensely and frequently than did Neanderthals.
The preference for intermediate structural habitats by Neanderthals is also
detectable at the landscape level. At this level, a number of studies from such
diverse geographical regions as Iberia (Finlayson & Giles, 2000), south-west
France (Mellars, 1996), the Middle East (Shea, 1998) and the edge of the
Russian Plain (Soffer, 1994) show beyond doubt that Neanderthals occupied
landscapes that were ecotonally rich “ that is landscapes that included a di-
versity of habitats over a small area. Topographically heterogeneous regions
are especially diverse. Other important ecotonal landscapes that appear to have
been repeatedly used by Neanderthals are wetlands, coastal landscapes, lake
mosaics and linear riverine stretches. The advantage of such areas is that they


50
(a)


40



30



20



10
Probability (%)




0 Max
Min
Bare Ground
-10
BARE0 BARE50 BARE100
BARE25 BARE75
Bare Ground (%)

Figure 5.1. Predicted patterns of vegetation structure in the Neanderthal Oxygen
Isotope Stage 3 site of Gibraltar (after Finlayson & Giles, 2000). (a) Distribution of
bare ground; (b) distribution of tree heights; (c) tree density (trees/ha.); (d) distribution
of shrub heights; (e) distribution of grass heights; (f ) cover of stone pine Pinus pinea;
(g) cover of juniper Juniperus phoenicea; (h) distribution of trees by trunk
circumference.
70 (b)

60


50

Max
40
Min
Tall Trees
30

Max
20
Min
Probability (%)




Medium Trees
10

Max
0
Min
Low Trees
-10
0 25 50 75 100
Cover (%)



60 (c)

50


40


30


20
Probability (%)




10


Max
0
Min
Number of Trees
-10
0-50 100-150 >200
50-100 150-200
Tree Number/Ha

Figure 5.1. (cont.)
60 (d)

50


40
Max
Min
30
Tall Shrubs

20
Max
Min
Probability (%)

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