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10 Medium Shrubs


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


80 (e)



60


Max
40 Min
Tall Grasses


Max
20
Min
Probability (%)




Medium Grasses
0
Max
Min
Low Grasses
-20
0 25 50 75 100
Cover (%)

Figure 5.1. (cont.)
50 (f)



40




30




20
Probability (%)




10
Max
Min
Stone Pine
0
0 25 50 75 100
Pinus pinea (%)


70 (g)


60


50


40


30
Probability (%)




20


Max
10
Min
Juniper
0
0 25 50 75 100
Juniperus phoenicea (%)

Figure 5.1. (cont.)
110 Neanderthals and Modern Humans

80 (h)



60


Max
40 Min
Small Circumf. Trees


Max
20
Min
Probability (%)




Med. Circumf. Trees
0
Max
Min
Large Circumf. Trees
-20
0 25 50 75 100
Presence (%)

Figure 5.1. (cont.)



contain a rich diversity of potential prey over short distances. These landscapes,
by providing a range of options, are also more buffered than more homogeneous
landscapes against environmental instability.
Because open plains habitats stretch across great areas over topographically
homogeneous land, Moderns utilising open plains de facto occupied homoge-
neous landscapes. There are several requirements for successfully exploiting
homogeneous open landscapes: (a) because a small range of highly mobile prey,
that may occur in localised areas at high density, are available, predators will
need to have behavioural tactics that allow for such mobility; (b) because of
the lack of cover predators will require speci¬c and specialised hunting tech-
niques; (c) because of the wide ranging nature of this behaviour pattern, and
the need for sources of water especially if high levels of meat are consumed,
predators will not operate well if such landscapes are in arid regions; and (d)
complex social systems are necessary to increase environmental resistance, as
we shall see later. The Eurasian Plain may have been exceptional if the humans
exploiting it were able to melt the readily available snow and ice.
In Chapter 4 I discussed the evidence for climatic adaptation in Neanderthals
and Modern Humans. Here I relate the evidence to geographic distribution
Comparative behaviour and ecology 111

200

700-1000-1300
180
1200-2100-2200

160


140 100-1200-1600


120
Tree Density (Trees/ha)




1400-1800-2000
100


80
0-0-700
MAX
60
MIN
40 MEAN
nigra sylvest pinast halep pinea
Species

Figure 5.2. Present-day density of pine (Pinus) trees in southern Iberia. Boxes
indicate altitude range in metres. nigra, P. nigra; sylvest, P. sylvestris; pinast, P.
pinaster, halep, P. halepensis; pinea, P. pinea. Contrast present distribution of P. pinea
(low altitude) with P. nigra (high altitude). During late Oxygen Isotope Stage 3,
P. nigra was the dominant pine in Gorham™s Cave, Gibraltar, currently at sea level.



patterns. It is widely accepted that Neanderthals were cold-adapted. I have no
doubt that populations of humans (Neanderthals or Moderns) living in cool
climates would have bene¬ted from adaptations that minimised heat loss. The
question is to what extent were such adaptations signi¬cant in de¬ning geo-
graphical range limits? The evidence suggests that they were unimportant. Late
Neanderthal sites are all in Mediterranean or sub-Mediterranean bioclimatic
zones and cool temperate sites are associated with warmer periods (Figure 5.3).
In Chapter 4, I showed that Neanderthals would have evacuated geographical
areas long before the arctic conditions characteristic of the strong glacial pulses
would have reached them. Instead, we have the Moderns occupying the plains of
Eurasia once the Neanderthal range has receded towards the south. The Moderns
too were hit by the glacial cold, but signi¬cantly later and when it reached greater
intensity towards the LGM (once Neanderthals were extinct). The range of the
Moderns only then extended south into the Mediterranean peninsulas of Iberia,
Italy and the Balkans, as suitable habitats closed in the north and opened up in
(a)




(b)
20




10
Number of Sites




Middle East

Mid/South Europe

0 N. European Plain
3 4 5a-d 5e
Oxygen Isotope Stage

Figure 5.3. (a) Occupation of Europe by Neanderthals. (b) The histogram (based on
data in Stringer & Gamble, 1993) illustrates the sporadic occupation of the North
European Plain, only during warm intervals. P, Permanent occupation; S,
semi-permanent occupation; N, never occupied. Bioclimates after Rivas-Mart´nez
±
(1996). Dark grey, Mediterranean; intermediate grey, sub-Mediterranean; pale grey,
temperate oceanic; white, temperate continental (below arrowed line) and boreal
(above arrowed line).
Comparative behaviour and ecology 113

the south. This raises the question of the extent to which the supposed tropical
morphology of the Moderns meant anything to them. They clearly survived the
cold of the north until they were forced south because of habitat and resource
loss. Clearly behavioural and physiological adaptations including the use of ¬re,
shelters, clothing, the inclusion of fat in the diet, food caching and a complex
social system were overriding factors for Moderns just as they are for Eskimos
today. Eskimos today do not differ from central Europeans in their ability to
retain heat. They simply wear suitable clothing and consume large amounts of
animal fat that permits a higher rate of non-shivering thermogenesis (Gisol¬ &
Mora, 2000). This begs the question of the extent to which comparisons with
modern populations are valid. The crucial point is that it was food and habitat,
not temperature, that limited the geographical distribution of Eurasian humans.


Home range, group size and related features

The spatial arrangement of resources, habitat and barriers affects the location,
movement patterns, foraging dynamics and persistence of organisms (Karieva,
1990; Danielson, 1991; Pulliam et al, 1992; Turner et al, 1995). In humans,
the way they move in the landscape exerts strong in¬‚uences on culture and
society (Kelly, 1992). In the case of the polar bear, a species that has evolved
into a High Arctic carnivore from an omnivorous ancestor, movements are

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