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America (around 45 Myr) and the late Eocene of Asia (around 35 Myr); Africa
was not reached until well into the early Miocene (around 20 Myr) (Cerde˜ o, n
1998). The family became extinct in North America during the Pliocene and
is therefore not relevant to the present discussion. Rhinoceroses disappeared
from Europe and large areas of Asia in the late Pleistocene, leaving Africa and
South-east Asia with species to the present. The genus Stephanorhinus pre-
dominated in Pleistocene Europe, with S. etruscus in the Pliocene and early
Pleistocene, S. hundsheimensis in the middle Pleistocene and S. kirchbergensis
and S. hemitoechus in the middle and late Pleistocene. There were also at least
three European endemic forms in the Pliocene that did not persist into the Pleis-
tocene (Cerde˜ o, 1998). Stephanorhinus kirchbergensis was a browser of open
n
forest and S. hemitoechus a grazer of open woodland and prairie (Gu©rin &
Patou-Mathis, 1996). The woolly rhinoceros Coelodonta antiquitatis arrived in
Europe from Asia at the end of the middle Pleistocene and also persisted until
the end of the Pleistocene (Cerde˜ o, 1998). This was a typical cold steppe grazer
n
whose geographical range extended across Eurasia, from Korea to Scotland and
Spain (Gu©rin & Pathou-Mathis, 1996). Finally, the large Elasmotherium sibir-
icum/caucasicum was present in eastern Europe during the middle and early late
Pleistocene and occasionally reached western Europe (Gu©rin & Patou-Mathis,
1996).
There was a greater diversity of taxa in Asia than in Europe during the
Plio-Pleistocene: the related genera Rhinoceros and Punjabitherium from
the early Pliocene; Dicerorhinus from the early Pleistocene; Chilotherium
during the Pliocene; as well as the genera described for Europe (Cerde˜ o, n
1998). Rhinoceros unicornis is present in the Pliocene and R. sondaicus from
the early Pleistocene. Both survive today, in India and in South-east Asia
Biogeographical patterns 23

respectively. Two species of Dicerorhinus are present in the early Pleistocene:
D. sumatrensis and D. lantianensis. The former still survives today in South-
east Asia. Stephanorhinus etruscus is present in Pliocene Asia and S. hund-
sheimensis, S. hemitoechus and S. kirchbergensis occur in the middle Pleis-
tocene. Stephanorhinus hemitoechus, unlike in Europe, does not persist into the
late Pleistocene. The other two do but not to the Pleistocene“Holocene bound-
ary. Chilotherium antiquitatis appears in Asia in the middle Pleistocene, earlier
than in Europe, and is preceded by the congener C. nihowanensis in the early
Pleistocene. Elasmotherium persisted from the Pliocene to the late Pleistocene,
E. caucasicum and E. inexpectatum in the late Pliocene/early Pleistocene and
E. sibiricum and E. peii from the middle Pleistocene (Cerde˜ o, 1998).
n
The two extant African species Diceros bicornis and Ceratotherium simum
were present in the Pliocene and have persisted to the present. Three other
species of Ceratotherium were present in Pliocene Africa and another in the
early Pleistocene. In North Africa S. hemitoechus was present in the late Pleis-
tocene (Cerde˜ o, 1998). Humans would have therefore been in contact with
n
rhinoceroses in Africa from the earliest times and in Eurasia throughout. There
would never have been contact in North America.


Artiodactyls (Order Artiodactyla)

Pigs (Suidae)
The Suidae ¬rst appear in the late Eocene in Eurasia and in the Miocene of
Africa. The peccaries (Tayassuidae) underwent a separate American radiation
from the early Oligocene (Cifelli, 1981). In Africa the Suidae underwent a large
radiation during the Pliocene and Pleistocene (Cooke 1968, 1978; White &
Harris, 1977; Bishop, 1999). There were six African Plio-Pleistocene genera.
Koplochoerus of the middle Pliocene to middle Pleistocene is thought to have
been ancestral to the extant African bush pig Potamochoerus porcus and for-
est hog Hylochoerus meinertzhageni (White & Harris, 1977; Bishop, 1999).
The genus Metridiochoerus, with four species from the middle Pliocene to the
middle Pleistocene, is considered ancestral to the extant warthog Phacocoerus
aeithiopicus (White & Harris, 1977; Bishop, 1999), a genus that ¬rst appears
in the Pliocene (Anderson, 1984). The Pliocene Notochoerus that evolved from
the late Moicene Nyanzachoerus, with three species, became extinct in the late
Pliocene/early Pleistocene (Anderson, 1984; Bishop, 1999)
The Eurasian genus Sus ¬rst appears in the early Pliocene (Anderson, 1984)
and is represented today by the widespread middle Pleistocene wild boar
S. scrofa, and South-east Asian bearded pig S. barbatus (and a number of
South-east Asian island forms) and also the Indian S. salvanius. In Europe
24 Neanderthals and Modern Humans

the Villafranchian species S. strozzii persists locally to the start of the middle
Pleistocene (Gu©rin & Patou-Mathis, 1996). In South-east Asia there are three
other middle Pleistocene forms, S. lydekkeri, S. macrognathus and S. of¬cinalis
(Tougard, 2001). Frozen ground and deep snow limited the distribution of the
Suidae, Sus being commonest in Eurasia during interglacials and interstadials
(Anderson, 1984) and they never penetrated the high latitude ˜Bering ¬lter™
(Flynn et al., 1991) to reach North America in the Pleistocene. There is no ev-
idence of human association with the Pleistocene peccaries (genera Mylohyus
and Platygonus) (Anderson, 1984) but there must have been some contact with
the recent forms of Tayassu. Pigs would have therefore been a regular feature of
the mid- and low-latitude open woodland and shrubland landscapes occupied
by humans throughout the Pleistocene in Africa and Eurasia.


Hippopotamuses (Hippopotamidae)
The family originates in the middle Miocene of Africa with the genus
Kenyapotamus (Pickford & Morales, 1994). The genus Hexaprotodon, that
includes the extant pygmy hippopotamus H. liberiensis (also placed under the
separate genus Choeropsis), is of late Miocene origin while Hippopotamus
is a Pliocene genus (Pickford & Morales, 1994). Hexaprotodon was a domi-
nant genus in the Tertiary with a distribution that included Europe, South and
South-east Asia as well as Africa. In Asia the genus persisted into the middle
Pleistocene (Anderson, 1984; Gu©rin & Patou-Matthis, 1996; Tougard, 2001).
Their African decline “ most were extinct by the Plio-Pleistocene boundary “
is related to the rise of the bovids that outcompeted them for food and habi-
tat (Anderson, 1984). On the other hand, the amphibious Hippopotamus did
not compete with the bovids. Four species are identi¬ed from Plio-Pleistocene
Africa, H. gorgops, H. aethiopicus, H. protamphibius and H. amphibius, which
has survived to today (Bromage & Schrenk, 1999). In India, Hippopotamus is
present in the early Pleistocene (Anderson, 1984). In Europe, hippopotamuses
had been absent throughout the Villafranchian but re-appeared in the middle
Pleistocene and persisted until the start of the last glaciation (Gu©rin & Patou-
Mathis, 1996). Hippopotamus major (=antiquus), considered by some authors
to be a large form of H. amphibius (Anderson, 1984), is a middle Pleistocene
form. Hippopotamus incognitos is regarded as a separate middle Pleistocene
species that persisted to the last glaciation (Gu©rin & Patou-Mathis, 1996).
Hippopotamus is an interglacial species outside Africa. Humans would there-
fore have been in potential contact with hippopotamuses throughout the Pleis-
tocene in Africa, for much of it in South-east Asia and elsewhere in Eurasia
during intarglacials. As with the Suidae, hippopotamuses never reached North
America.
Biogeographical patterns 25

Camels (Camelidae)
Contact between humans and camels is unlikely to have been major during the
Pleistocene, and only when humans penetrated open desert and steppic environ-
ments. Camels ¬rst appeared in the Eocene (40“45 Myr bp) of North America
(Stanley et al., 1994) and experienced a major radiation in the Pliocene and
Pleistocene (Cifelli, 1981). Migration into Asia occurred at 3 Myr bp (Stanley
et al., 1994). Camelus, represented by the extant dromedary C. dromedarius
that is no longer found in the wild and by the bactrian camel C. bactrianus that
may survive in Central Asia, was present in Pleistocene North Africa and Asia
(Anderson, 1984).
In North America, the camel Camelops and the llamas Hemiauchenia and
Palaeolama persisted until the end of the Pleistocene but their remains are rare
in Palaeo-Indian sites (Anderson, 1984). The giant camel genus Titanotylopus
became extinct in the Middle Pleistocene.

Chevrotain and mouse deer (Tragulidae)
The Chevrotains ¬rst appear in the fossil record in the middle Miocene (Cifelli,
1981) and they are the ¬rst group to diverge among the ruminants (Nikaido
et al., 1999). There is very little fossil information available. The two extant
South-east Asian species, the greater mouse deer Tragulus napu and the lesser
mouse deer T. javanicus, are present in late Pleistocene sites (Tougard, 2001).
Currently the family has a single representative in Africa and three in South
and South-east Asia. They are forest species and are unlikely to have been
frequently met by humans on account of habitat differences.

Giraffes and okapi (Giraf¬dae)
The Giraf¬dae ¬rst appear in the Miocene of Africa and radiate during the
Pliocene with a signi¬cant decline in diversity in the Pleistocene (Cifelli, 1981).
The genus Giraffa reached southern Europe and Asia in the Pliocene, remaining
in the latter continent into the late Pleistocene and becoming extinct in North
Africa in the Holocene (Anderson, 1984). Five species have been described
from the Plio-Pleistocene of Africa (Bromage & Schrenk, 1999) although their
status as variants of the extant G. camelopardalis is unclear (Anderson, 1984).
The okapi Okapia jonhstoni is a rainforest species from the early Pleistocene
(Anderson, 1984). It shared a common ancestor with the Pliocene O. stillei
(Hamilton, 1978). The gigantic giraf¬ds Sivatherium were present in the
Pliocene of Eurasia and Africa, S. giganteum/maurusium, an open woodland
species, persisted to the middle Pleistocene and has been associated with African
Acheulian sites (Hamilton, 1978; Anderson, 1984). Giraf¬ds, except the rain-
forest okapis, would therefore have been a regular feature of the open landscapes
inhabited by humans in Africa, Asia and, at times, southern Europe.
26 Neanderthals and Modern Humans

Deer (Cervidae)
Deer originated in the mid-Tertiary of the Old World tropics and evolved as
global temperatures and climatic stability declined, culminating with the Pleis-
tocene glaciations. Already in the Tertiary, deer had evolved strategies that
permitted them to thrive in cold climates by taking advantage of seasonal high-
quality plant foods. Modern genera appeared in the Early Miocene grasslands
(Geist, 1998). Three sub-families are recognised, the Cervinae (Old World deer),
the Muntiacinae (muntjacs and tufted deer), and Odocoileinae (New World deer)
(Miyamoto et al., 1990). Some authors add the antler-less musk deer of China
as a fourth subfamily “ the Moschinae (Eisenberg, 1981).
The Old World deer originated in mid-Tertiary Eurasia, splitting from the
muntiacines around 6“8 Myr (Miyamoto et al., 1990), and subsequently spread-
ing to Africa and India (Geist, 1998). The group saw repeated tropics to cold
climate radiations during the Pleistocene. The following account is largely
based on Geist (1998). The forest muntiacines appear in the late Miocene of
China and are today represented by the genera Muntiacus and Elaphodus of
India, China and South-east Asia. It is unlikely that these small forest deer
were in any way signi¬cant as prey to hominids as other deer were, although it
should be noted that they may have been regionally or locally exploited, as evi-
denced by their presence in South-east Asian middle and late Pleistocene sites “
M. muntjak, M. szechuanensis, Elaphodus sp. (Tougard, 2001).
As in the case of human evolution, deer evolution proceeded from forest to
open steppe types via an intermediate, ecotone, stage. This stage is represented
by the sub-tropical, three-pronged, deer represented today by the genera Axis,
Rusa and Rucervus. Fossil three-pronged deer that appear to have originated in
warm climates spread widely across Eurasia in the Villafranchian, represented
by Cervus etuerarium, C. paradinensis, C.philisi, C. perolensis, Nipponicervus,
Rusa and Rucervus. These deer died out at the end of the Villafranchian, or
the early Pleistocene in the case of the Middle East. In Japan, Nipponicervus
survived to the end of the Pleistocene. In general terms, deer were a small
element of the Villafranchian faunas. A number of advanced cervine deer also
arrived in the open landscapes of Europe (some reaching China) during the
Villafranchian (Creuzetoceros ramosus, Arvenoceros ardei and Eucladoceros
spp.) but these did not survive into the middle Pleistocene when the cold climate
megacerines and elaphines, the moose Alces and the roe deer Capreolus spread.
Linked to climate deterioration was a westward intrusion of east Asian species,
C. punjabensis reaching India around 2.5 Myr, that coincided with a major
extinction of the endemic Villafranchian fauna.
Geist (1998) identi¬es four major Old World deer radiations that penetrated
temperate and cold areas:
Biogeographical patterns 27

r The ¬rst is represented by the white-lipped deer Przewalskium albirostris,
a high alpine steppe species of eastern Tibet.
r The second is represented by Pere David™s deer Elaphurus davidianus, of
the Chinese swamps that ¬rst appeared in the Pliocene.
r The third was the radiation of the highly successful megacerines, of which
only the fallow deers (Dama dama and D. mesopotamica “ considered either
species or subspecies by different authors) and the Irish elk Megaceros
giganteus hibernae survived into the Holocene. The history of the megac-
erines can be traced back 1.4 Myr in the eastern Mediterranean with the
¬rst reaching Europe around 700 kyr. They were also present in the Vil-
lafranchian of Japan. Megacerines were widely distributed, in North Africa,
Europe, central Asia and Japan but did not reach eastern Siberia and Beringia
even though M. giganteus inhabited the mammoth steppe. Three lines within
the group entered Europe. These were: (a) the fallow deer Dama, the
¬rst to reach Europe in the mid-Pleistocene being Dama (d.) clactoniana;
(b) the forest sub-genus Megaceroides, represented by M. verticornis and
M. solilhacus in the early Pleistocene, which disappeared early in Europe but
persisted (M. algericus) to the end of the Pleistocene in North Africa; and (c)
the grassland and steppe sub-genus Megaloceros, with the middle Pleisto-
cene M. savini and M. antecedens and the late Pleistocene M. giganteus.
r The fourth, and the most successful, was that of the genus Cervus. The
group was represented by opportunistic, savannah, deer with some species
adapting to open landscapes. Africa was not colonised except north of the
Sahara. The oldest red deer C. elephus remains are 1 Myr and from Alaska.
It is thought that the red deer originated in Asia and the species ¬rst appeared
in Europe in the middle Pleistocene (700 kyr), coinciding with an arrival of
a range of Asian cool temperate species. The success of the red deer, with
its many forms including the North American wapiti, is due to its ecological
plasticity and mixed feeding strategy. Sika deer C. nippon ¬rst appeared in
China in the late Pliocene.

The New World deer originated in the cool temperate and cold zones of North
America in the Pliocene. Miyamoto et al. (1990) put the split from the Old World
deer lineage between 12.4 and 9.3 Myr. There are 10 extant genera: ¬ve South
American and ¬ve North American, with elements in Eurasia. The latter, of con-
cern to us here, are Alces, Rangifer, Capreolus, Hydropotes and Odocoeilus.
The moose (Alces) branch splits off from all other New World deer in the late
Tertiary. In Europe it appeared as A. (Libralces) gallicus in the late Pliocene
around 2.0“1.6 Myr. This was the only deer to survive the Villafranchian ex-
tinctions. The ability of moose as a coloniser with rapid dispersal, related to
28 Neanderthals and Modern Humans

its resource strategies that exploit instabilities in its environment, undoubtedly
made it a successful lineage. Alces gallicus was followed in Europe by the large
A. latifrons around 700 kyr bp, coinciding with the arrival of other Asian species
including the red, roe and fallow deers and the reindeer. This species was already
present in its probable centre of evolution in eastern Siberia/Beringia between
1.2 and 0.5 Myr. It spread widely across Eurasia and North America between
45—¦ and 71—¦ N, suggesting a cold-adapted nature although this seems dif¬cult
to reconcile with its abundance in interglacial deposits from warm areas in the
middle Pleistocene. There is a similar pattern of high incidence of A. alces in
interglacial, and absence in glacial, deposits. Moose with modern characteris-
tics are found in deposits at the beginning of the last cold stage around 100 kyr
but it is not clear if these are derived from A. latifrons. A similar pattern of
expansion is observed in the other two North American genera that penetrated
Eurasia. The reindeer Rangifer tarandus spread to occupy the widest longitu-
dinal (circumpolar) distribution of any ungulate, from 14—¦ W to 5—¦ E. During
cold episodes reindeer reached southern France and even penetrated Iberia, and
were commonly hunted by humans. The roe deer Capreolus is the third North
American cold-adapted deer. Its ¬rst appearance in Europe, as we have seen,
was in the middle Pleistocene and warm climates were colonised successfully
later“for example, France, Iberia and Italy around 200 kyr. Roe deer never pene-
trated into North Africa. Two species are recognised, the European C. capreolus
and the Siberian C. pygargus. In North America Odocoileus brachydontus ¬rst
appeared around 3.9“3.5 Myr and spread southwards reaching South America.
Together with its later form, the current white-tailed deer O. virginianus, this
lineage represents a unique success that is attributed to its opportunistic na-

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