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ture. The mule deer is a post-Pleistocene sub-species of the black-tailed deer
O. hemionus thought to have hybridised with white-tailed deer.
It is clear from this brief account that deer have been ubiquitous components
of the ecosystems of much of Eurasia and North America, especially since the
middle Pleistocene. We will see later how particular species, especially red
deer and reindeer, were a major element of the staple diet of humans in these
continents throughout the Pleistocene.

Pronghorn (Antilocapridae)
The pronghorn Antilocapra americana of the late Pleistocene was one of
the two dominant ungulates of the western North American high plains
(Eisenberg, 1981). It is the only surviving species in the family. Four species
in the genus Capromeryx ranged from California to Florida and south to Mex-
ico from the late Pliocene to the late Pleistocene. Two species of Stockoceros
occurred in Mexico, Arizona and New Mexico in the late Pleistocene. Five
Biogeographical patterns 29

species of Tetrameryx occurred in Texas, California and New Mexico in the
Pleistocene. Stockoceros remains have been found in association with humans
dated at 11.5 kyr bp (Anderson, 1984).

Wild cattle and antelopes (Bovidae)
The bovid family radiated extensively in the Pliocene and, particularly, the Pleis-
tocene (Cifelli, 1981). It is the most diverse family and naturally colonised all
continents except South America and Australia. It is also the family that, along
with the deer, has been central as a food resource to the hominid populations
of the Pleistocene. The Bovidae is divided into nine sub-families:

(1) Alcelaphinae: wildebeests, hartebeest; Africa; 7 extant species.
(2) Antelopinae: antelopes, saiga; Africa, Asia; 38 extant species.
(3) Bovinae: kudu, eland, four-horned antelope, bison, wild cattle; Africa,
Eurasia, North America; 24 extant species.
(4) Caprinae: chamois, mountain goat, serow, goral, musk ox, takin, bharal,
goats, sheep; Eurasia, North America, Africa; 32 extant species.
(5) Cephalophinae: duikers; Africa; 19 extant species.
(6) Hippotraginae: sable, roan antelopes, blue buck; Africa, Asia; 7 extant
species.
(7) Peleinae: rhebok; Africa; 1 extant species.
(8) Aepycerotinae: impala; Africa; 1 extant species.
(9) Reduncinae: kob, reedbuck; Africa; 8 extant species.

Representatives of all the sub-families have been found in association with
hominid levels (Anderson, 1984; Bromage & Schrenk, 1999; Tougard, 2001)
suggesting a long and continued interrelationship between hominids and the
Bovidae.



Herbivore distribution patterns

A cluster analysis (Figure 2.2) of the Pleistocene distribution of mammalian
herbivores at the level of sub-family separates geographical regions in the fol-
lowing manner (in order of separation):

(1) Africa.
(2) Desert regions.
(3) India, the eastern mid-latitude belt, South-east Asia and China (South-east
Asia, then China, separate from the cluster).
30 Neanderthals and Modern Humans




Figure 2.2. Separation of geographical regions by similarity of mammalian herbivore
components. (a) Results of cluster analysis; (b) representation of results in map form.
Biogeographical patterns 31


(4) The Holarctic. The Holarctic cluster then separates out, in order, into,
(a) the western and central mid-latitude belt;
(b) North America;
(c) the Eurasian Plains and the Arctic.

It is interesting to note that the separation of the western and central mid-latitude
belt from its cluster occurs earlier than the separation of South-east Asia from
its group, indicating its distinctiveness, and that North America separates from
its cluster before China from its cluster.
There are two underlying patterns to this distribution. On the one hand there
is the historical pattern of the isolation of the various regions. Thus Africa, the
South-east Asian cluster and North America are distinct because of the periods
of isolation caused by deserts, ice and sea barriers. South America and Aus-
tralia, had they been included in the analysis, would have separated out even
earlier for this reason. In this respect North America is the least distinctive and
Africa the most. At another level there is the ecological pattern. The desert re-
gions, independently of geographical location, separate out early on account of
the impoverished faunas. The separation of the Eurasian Plains“Arctic“North
America group from the western and central mid-latitude belt is a combination
of the two patterns. It shows that the Eurasian Plains, in spite of a huge longi-
tudinal spread, are an ecological unit that cluster with the Arctic (a northward
extension of the plains) and North America (an eastern continuation). This unit
is sub-divisible into Eurasian Plains/Arctic“North America for reasons of occa-
sional isolation. The separation of the eastern mid-latitude belt from the rest of
the belt, and its af¬nity particularly with India, re¬‚ects a shared history related
to proximity. The separation of the mid-latitude belt from the Eurasian Plains is
indicative of a marked ecological boundary that, as I shall develop in this book,
has been crucial in human evolution.
Hominids would have encountered a group of similar mammalian herbivore
size classes and ecological equivalents across their geographical range in the
Pleistocene (Table 2.1 & 2.2; Figures 2.3 & 2.4). I will discuss the exploitation
of prey by hominids in Chapter 5. At this stage, I will highlight some macro-
ecological patterns. When we consider the range of open habitats (usually with
few, scattered, trees and shrubs) and shrublands that hominids would have
encountered in different parts of the geographical range we ¬nd recurrent trends.
For huge areas of Africa, India, China, in patches within the mid-latitude belt,
across the Eurasian Plains, in the Arctic and over much of North America similar
mammals would have been encountered. Among the largest forms (i.e. >1000
kg) would have been the elephants (including mammoths) and the rhinoceroses.
In the next size category (500“1000 kg) would have been the large bovids such
as bison Bison spp., wild cattle Bos spp. or African buffalo Syncerus spp. The
32 Neanderthals and Modern Humans


Table 2.2. Frequency of occurrence of mammalian herbivore taxa in African
hominid and non-hominid sites (after Bromage & Schrenk, 1999). This table,
together with Figure 2.4, suggests that the main taxa associated with hominids
were pigs (Suidae), bovids (Bovinae), wildebeests and related forms
(Alcelaphinae), giraffes (Giraf¬dae), horses (Equidae), and hippopotamuses
(Hippopotamidae). The majority are medium-sized taxa (see text).
Hippopotamuses may be associated with humans because many sites are near
wetlands, where hippopotamuses preferentially would have been found, and
need not indicate a hunting preference. The relationship with giraffes is
unclear. A positive but not a statistically signi¬cant relationship is indicated
by parentheses

More than Fewer than
expected in expected in
χ2
Taxon hominid sites hominid sites Signi¬cance
+ <0.005
Suidae 13.53
+ <0.05
Bovinae 8.62
Antelopinae (+) 5.58 NS
Reduncinae (+) 1.31 NS
+ <0.005
Alcelaphinae 10.66
+ <0.025
Giraf¬dae 18.5
+ <0.05
Equidae 8.61
Elephantidae (+) 6.94 NS
Rhinocerotidae (+) 2.54 NS
+ <0.025
Hippopotamidae 9.93
Hippotraginae (+) 7.81 NS
Aepycerotinae (+) 7.53 NS
Camelidae (+) 5.57 NS


predominant prey would have been in the 100“500 kg category. In Africa this
category would have been dominated by bovids such as gnus Connochaetes
sp., hartebeests Damalops and Alcelaphus spp., or kob Kobus sp. In Eurasia
and North America this category was largely ¬lled by the deer Cervus and
Rangifer spp. Horses Equus spp. would have been present throughout. The
smaller category (<100 kg) would have included the smaller African bovids
such as steenbok Raphicerus sp., springbok Antidorcas sp. and the gazelles
Gazella sp. that also occupied large areas of southern Asia. In Eurasia the group
would have included the fallow deer Dama sp. and the saiga antelope Saiga
sp. Pigs such as boar Sus sp. in Eurasia, warthog Phacocoerus sp. in Africa or
peccary Platygonus sp. in North America would have largely fallen into this
category. These animals, occurring as similar or different species across Africa,
Eurasia and North America, would have been those available to all hominid
populations throughout their range.
Biogeographical patterns 33




Figure 2.3. Main mammalian herbivore components by geographical region.

The large tracts of desert would have been impoverished relative to these open
habitats. The main species would have been adapted to arid conditions. Some,
such as gazelles, Gazella spp., would have been encountered in different parts
of the range. Others, such as camels, Camelus spp., in Eurasia or Camelops sp.,
in North America, or oryx Oryx sp., would have been geographically localised.
Dense temperate, tropical and equatorial forests were not regularly-used
hominid habitats (Gamble, 1993). Mammalian herbivores do not reach the high
diversity, densities or levels of aggregation that they do for example in open
grasslands (Bigalke, 1968; Jarman & Sinclair, 1979; Prins & Reitsma, 1989;
Srikosamatara, 1993; Fritz & Duncan, 1994; Fryxell, 1995; Prins & Olff, 1998),
and prey are more dif¬cult to detect and catch. They are often of small size.
It is not surprising, therefore, that forest herbivores, e.g. roe deer Capreolus
capreolus in Eurasia, duikers Cephalophus spp. in Africa or muntjacs Muntiacus
sp. in South-east Asia, do not feature as prominently in hominid sites as do open
habitat herbivores.
Independently of region, and at smaller spatial scales, there are habitats
that have been favoured by hominids on account of high prey densities or
because they hold particular species. Such habitats are readily identi¬able by
their features in the landscape. These are:
34 Neanderthals and Modern Humans


5


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