Wil Roebroeks Hominid behaviour and the earliest

Faculty of Archaeology, occupation of Europe: an exploration

Leiden University,
P.O. Box 9515,
2300 RA Leiden,
The Netherlands.
E-mail: W.Roebroeks@
arch.LeidenUniv.nl
Received 12 October 2000
Revision received 23 May
2001 and accepted 23 May
2001
Keywords: Europe,
Pleistocene, colonization,
hominid behaviour, hunting,
palaeolithic.
The last decade has witnessed a heated debate over the age and the
character of the earliest occupation of Europe. This paper addresses
two aspects of the debate, one dealing with the chronology of
occupation, which is put to use in the second issue, an exploration of
the behaviour of the earliest occupants of Europe. The review of the
debate on chronology concludes that a short chronology applies to
Europe north of the large mountain chains of the Alps and the
Pyrenees, where the earliest traces of a human presence date back to
about half a million years ago. In this phased-colonisation model, the
Mediterranean, and especially Spain, saw an earlier occupation,
starting around the end of the Lower Pleistocene. The archaeological
record of these first Europeans suggests that from the first presence in
northern Europe onwards, regular hunting of large game was com-
mon practice among Middle Pleistocene hominids. By situating this
archaeological evidence in the context of findings from a range of
other disciplines I develop a behavioural scenario which suggests that,
at its latest by the Middle Pleistocene, increased forms of social
cooperation, exchange of information within larger groups and in
general forms of behaviour based on a ‘‘release from proximity’’ had
become a standard ingredient of the hominid behavioural repertoire.
 2001 Academic Press

Journal of Human Evolution (2001) 41, 437–461

doi:10.1006/jhev.2001.0499
Available online at http://www.idealibrary.com on

Introduction discussion, the last decade has seen an

The present day distribution of the human
species is the result of a complicated process
of dispersal from the inferred African homi-
nid cradle that started probably around
2 mya. Given the age and the number of
hominid remains from East Africa, few
would question that beyond all reasonable
doubt hominids evolved there (but see
Dennell, 2001 for a critical review). The
timetable of the subsequent dispersal is
hotly debated though, with recent evidence
even pointing towards the possibility of a
Late Pliocene to Early Pleistocene first ‘‘Out
of Africa’’ radiation (Swisher et al., 1994;
Gabunia et al., 2000b; Balter & Gibbons,
2000; Lordkipanidze et al., 2000; Huffman,
2001; but see also Langbroek & Roebroeks,
2000). Within the context of this larger
intensive debate over the age and the
character of the earliest occupation of
Europe (see for instance: Bonifay &
Vandermeersch, 1991; Roebroeks & Van
Kolfschoten, 1994, 1995, 1998; Carbonell
et al., 1995, 1996, 1999a, b; Dennell &
Roebroeks, 1996; Turq et al., 1996; Villa,
1996, 2001; A. Turner, 1999; Oms et al.,
2000; Balter, 2001). The debate is far from
over, and new discoveries both within and
outside Europe have fuelled its intensity and
have led to the formulation of various com-
peting models favouring a wide range of
chronological scenarios. In the middle of all
this uncertainty and controversy, however,
the debate has resulted in a kind of consen-
sus view that the European archaeological
record changes significantly around 500 ka
BP, with an increasing number of sites

0047–2484/01/100437+25$35.00/0  2001 Academic Press

438 . 
probably indicating a more substantial
occupation than in the period before (e.g.,
Roebroeks & Van Kolfschoten, 1994;
Dennell & Roebroeks, 1996; Dennell, 1998;
Bermúdez de Castro et al., 1999; Villa,
2001). Most authors also agree now that
north of the large mountain chains of the
Pyrenees and the Alps the first unambiguous
traces of human occupation date from about
500 ka BP; the Mediterranean witnessed
earlier human presence than the more
northern areas, though the exact age
difference between the southern and more
northern parts of Europe remains to be
established.
This paper consists of two rather different
parts, each addressing an issue related to the
topic of the earlier occupation of Europe.
First, I will give a short review of the past
decade’s debate on the earliest occupation of
Europe, in which I will focus on the short
chronology (Roebroeks & Van Kolfschoten,
1994) and the implications and the signifi-
cance of the Atapuerca TD6 and other
Spanish finds for the debate. The data on
the earliest occupation history of Europe will
be put to use in the second, more explorative
part. This discusses the behaviour of the
hominids responsible for the earliest occu-
pation of Europe, with an emphasis on the
first ‘‘substantial’’ settlement of Europe
from about 500–600 ka onwards. The
past decade has seen a wealth of cross-
disciplinary studies concerning various
aspects of biology and behaviour of Lower
and Middle Palaeolithic hominids. By using
various independent, but convergent, lines
of evidence from a range of disciplines to
contextualise the archaeology of these first
Europeans, I will follow Aiello’s (1998) lead
and try to set limits to archaeological specu-
lations on their behaviour and to sketch a
basic outline of the social organization of
these hominids and the way this might have
related to the colonisation of various envi-
ronments. While the review of the debate on
the first occupation is heavily data oriented,
the part of the paper dealing with the second
issue is considerably more explorative, pull-
ing together data and ideas from a wide
number of sources in an attempt to advance
our knowledge of the issue. The resulting
scenario makes some predictions on the
character of the archaeological record and
hence can be tested in some aspects, but it
also contains elements that are more difficult
to verify, such as statements on language or
demography. The fact that these concepts
are difficult to operationalise in studying
the fossil record should not prevent us
from occasionally attacking these ques-
tions, especially by means of independent
yet convergent lines of cross-disciplinary
evidence.
Earliest Europe—the debate
Reviewing the evidence pertinent to the
first human settlement of Europe, I have
argued earlier that the European archaeo-
logical record changes substantially around
500,000 years ago. Together with T. van
Kolfschoten I have taken this as indicating
that Europe was not colonised until around
half a million years ago (Roebroeks, 1994;
Roebroeks & Van Kolfschoten, 1994, 1995,
1998; Dennell & Roebroeks, 1996; cf.
Dennell, 1983). This so-called short chro-
nology was based on a reassessment of
artefactual and dating evidence from a large
number of European sites, carried out in the
early 1990s. In our reading of the evidence,
there was a difference between the European
record from before 500–600 ka BP and the
later period (500–600 ka BP is the estimated
date of the appearance of a biostratigraphi-
cally important vole, Arvicola terrestris can-
tiana, in northwestern Europe situated in
the second part of the Cromerian complex,
about 500 ka BP). A recent study in Spain
has shown that there, as elsewhere in
Europe, this transition and hence the
Biharian to Toringian boundary occurs
within the Brunhes epoch (cf. Agusti et al.,
        
1999). Before 500–600 ka virtually all finds
come from a coarse matrix, afterwards we
have primary context sites in fine-grained
deposits. The small assemblages dating from
before 500 ka are virtually all the result of
selection by archaeologists of primitive
‘‘worked’’ pieces from natural deposits; by
contrast, younger assemblages are often
excavated from knapping floors. We pro-
posed two basic ways to interpret these
differences. The pre-500 ka assemblages
could reflect the sparse traces of intermittent
occupation of Europe by people with
‘‘primitive’’, ‘‘Oldowan’’-type toolkits differ-
ent from later ones, while substantial colon-
isation of Europe took place from about
500 ka onwards (cf. Turner, 1992). How-
ever, in our reading of the evidence, many
purportedly early archaeological sites did
not contain firm artefactual evidence for an
early human presence. For instance, this is
the case for Le Vallonet in France (De
Lumley et al., 1988; see for critical com-
ments on the lithic assemblage also White,
1995; Santonja, 1996; Villa, 1996) and
the lower levels of Kärlich (A and B) in
Germany (cf. Baales et al., 2000). Other
sites were at least problematic from a
chronological point of view (such as Monte
Poggiolo in Italy, cf. Mussi, 1995; Villa,
2001) and/or shown to be younger than
originally thought, such as Soleihac in
France (Raynal et al., 1995) and Isernia in
Italy (see also Cuenca-Bescós et al., 1999;
Villa, 2001). Hence, in view of the attributes
of the ‘‘artefacts’’ and the contexts of the
pre-500 ka sites (see Roebroeks & Van
Kolfschoten, 1994, 1995), we interpreted
these differences as strongly suggesting
that there was no indisputable proof for
human occupation of Europe prior to about
500 ka BP.
Our scenario had several advantages.
First, it is supported by a body of data
independent of arguments concerning stone
tools: the chronological distribution of
human remains. From about OIS 13
439
onward we had Middle Pleistocene human
remains all over Europe, whereas from the
long period before we did not have a single
uncontested tooth, despite the huge amount
of other mammalian fossils known from this
time range. That is, until the Atapuerca
TD6 finds were discovered (Carbonell et al.,
1995).
As a second advantage, the pattern was
backed by a long history of intensive
research, as can be assessed by a look at the
history of European palaeolithic archaeology
and mammalian palaeontology. The
nineteenth- and early twentieth-century
search for early humans was a pan-European
phenomenon, and every country had its own
searchers and sets of early sites. As in
England, which I have treated in detail
(Roebroeks, 1996), all over Europe count-
less exposures were subjected to close
scrutiny and many presumed ‘‘primitive’’
artefacts were discovered in Early Pleis-
tocene and Tertiary deposits (cf. Boule,
1921; Obermaier, 1912; Sollas, 1924).
Many of these have, over many years,
yielded rich mammal faunas, but never any
convincing traces of human presence. This
applies to such different cases as the Somme
valley in northern France (Tuffreau &
Antoine, 1995), the Lower Pleistocene
Tegelen clay pits in the Netherlands (e.g.,
Dubois, 1904; see also Luttschwager & van
Bemmel, 1962; Peeters et al., 1988), the
Upper Val d’Arno basin in Italy, whose
mammal fauna was already known to the
seventeenth-century naturalist Nicolaus
Steno (cf. De Mortillet, 1883) and the pro-
lific early Cromerian site of Süssenborn near
Weimar (Germany), a site where Goethe
collected fossils (Nolte et al., 1969). The
failure of the many investigations looking for
Lower Pleistocene lithic assemblages and
human fossils strongly suggests the absence
of hominids from, especially, northern
Europe prior to the Middle Pleistocene. The
southern and eastern parts of Europe
have been investigated for as long but not
440 . 

10
23 19/20
6
26 17 22/
4 24/
31
12/15
30

13

21
5/8

2 14 16
28
25 29

1/27

7 11

18
3/9

Figure 1. Locations of the European key sites discussed in the text. 1, Ambrona; 2, Atapuerca; 3, Barranco
León; 4, Biache-Saint-Vaast; 5, La Borde; 6, Boxgrove; 7, Ceprano; 8, Coudoulous; 9, Fuente Nueva; 10,
Hoxne; 11, Isernia; 12, Kärlich; 13, Mauer; 14, Mauran; 15, Miesenheim; 16, Monte Poggiolo; 17,
Neandertal; 18, Notarchirico; 19, Salzgitter-Lebenstedt; 20, Schöningen; 21, Soleihac; 22, Süssenhorn;
23, Swanscombe; 24, Taubach; 25, Tautavel; 26, Tegelen; 27, Torralba; 28, Le Vallonet; 29, Val d’Arno;
30, Wallertheim; 31, Weimar.

nearly as intensively as northern Europe,
and hence there is a greater probability of
surprise there.
The third and most important advantage
of the short chronology was that its basic
elements were very easy to falsify. As we
wrote earlier: ‘‘The find of only one Early
Pleistocene site of primary context would
disprove it, and one would have to conclude
that before about 500,000 occupation
existed (but was largely intermittent). New
studies of some of the sites mentioned
in our survey could lead to such a result’’
(Roebroeks & Van Kolfschoten, 1994: 500).
The finds from Atapuerca TD6 have done
exactly this, while the finds from the Guadix
Baza basin in southern Spain seem to be
even older than Atapuerca, judging from the
fauna associated with the artefacts (Turq
et al., 1996; Oms et al., 2000). The TD6
assemblage was recovered from deposits
that on palaeomagnetic evidence (Parés &
Pérez-González, 1999) and on the basis of
ESR and U-series dating of teeth (Falguères
et al., 1999) have been situated at the very
end of the Lower Pleistocene (cf. Carbonell
et al., 1995). Furthermore, the TD6 fauna
contains the vole Mimomys savini and is
thus earlier than all archaeological sites
mentioned above. Hence the pan-European
value of the short chronology has been
falsified, and the Mediterranean seems to
have seen some earlier hominid presence.
Exactly how much earlier is difficult to say,
though. As stressed elsewhere (Dennell &
Roebroeks, 1996; Van Kolfschoten, 1998),
full acceptance of the Lower Pleistocene age
of the TD6 assemblage is not unproblematic
        
for some palaeontologists: despite the range
of dating techniques applied at Atapuerca,
the TD6 dating evidence is not entirely
consistent with the evidence from the wider
setting of the site, as its fauna contains
elements which elsewhere in Europe
commonly appear above the Brunhes–
Matuyama boundary (Van Kolfschoten,
1998, 1999). Relevant for this discussion is
the stratigraphical age of the species within
the Allophaiomys pliocaenicus–Microtus (Sten-
ocranius) gregalis lineage. M. (st.) gregaloides
and M. gregalis forms date from the earlier
part of the Middle Pleistocene, and the
presence of M. gregaloides indicates that the
reversed part of the Trinchera Dolina fill
may represent one of the intra-Brunhes
reversals, according to Van Kolfschoten
(1999), though this argument has been
countered by Parés & Pérez-González
(1999). There is clearly a biostratigraphical
problem here, which needs to be resolved.
No matter how large the time difference
between the first occupation of southern and
northern parts of Europe will eventually turn
out to be, what is important here is that the
TD6 finds are earlier than any other good
archaeological site known in Europe before
its discovery. The palaeomagnetic and
faunal evidence from Fuente Nueva-3 and
Barranco León in the Guadix-Baza basin in
southern Spain points to an even older pres-
ence of hominids in southern Spain, now
estimated to be pre-Jaramillo, minimally
around 1 million years BP (Oms et al.,
2000).
The Atapuerca TD6 case seems to have
closed the debate around the age of the
Italian archaeological site of Isernia, for a
long time presented as a late Lower
Pleistocene, late Matuyama site on the basis
of both K/Ar and palaeomagnetic dating
(McPherron & Schmidt, 1983; Peretto,
1991; Peretto et al., 1983). Apart from the
somewhat controversial nature of these
dates (cf. Villa, 1996), the presence of
A. terrestris cantiana in the Isernia fauna
441
was difficult to reconcile with a Lower
Pleistocene age of the site (cf. Roebroeks
& Van Kolfschoten, 1994, 1998; Van
Kolfschoten, 1996, 1998; Von Koenigswald
& Van Kolfschoten, 1996; Agusti et al.,
1999; Cuenca-Bescós et al., 1999; Sala,
1999). Others workers stressed the mis-
match between the macrofaunal evidence
and the inferred Lower Pleistocene age
(Petronio & Sardella, 1999; cf. Villa, 2001).
In recent publications the Isernia team has
placed the site in the (early part of the)
Brunhes epoch (Gagnepain et al., 1996;
Peretto et al., 2000), without any discussion
of the debate reviewed here. Coltorti et al.
(2000) recently reported an Ar/Ar date on
sanidine from the archaeological level of
60510 ka, which agrees considerably
better with the fauna from the site than the
previous age estimates. Discrepancies be-
tween various sets of data, as in the case of
Isernia, should be addressed explicitly, as
they show where our knowledge of the issues
at stake is inadequate and hence new insight
can be gained. At the same time the Isernia
case shows how powerful a tool the
biostratigraphy of small mammals is, and
biostratigraphical subdivisions of the
Pleistocene are becoming increasingly more
fine-grained. Recent studies of small mam-
mal assemblages from northern and central
Europe by S. Parfitt (London) and T. van
Kolfschoten (Leiden) have shown that the
middle part of the Middle Pleistocene is
more complex than hitherto envisaged
(personal communication, 2000).
It cannot be overemphasised that
Atapuerca TD6 and the Guadix-Baza basin
constitute, for the time being, a chronologi-
cal ‘‘loner’’ in the whole of Europe. It is to
be expected that more traces of a hominid
presence of about the age of Atapuerca TD6
and the Guadix-Baza basin sites will eventu-
ally be uncovered elsewhere in southern
Europe (e.g., in Turkey, Greece, former
Yugoslavia and Italy), but these simply have
not been found, or rather identified, yet. All
442 . 
other European sites thought to date from
the late Matuyama or the early part of the
Brunhes are problematic, and the new dates
for Atapuerca TD6 cannot refute the earlier
doubts on these sites. Each piece of evidence
has to be considered in its own right, not by
consistency with a fashionable interpret-
ation. The Atapuerca TD6 finds in no way
lend credibility to the disputed stone assem-
blage from Le Vallonet or to the age esti-
mate for Ceprano, Italy, of about 800,000
years BP (Ascenzi et al., 1996, 2000). The
Ceprano skull is a stray find and its age
assessment is based on stratigraphical corre-
lations only (cf. Carbonell et al., 1999a: 668;
Villa, 2001). Villa’s (2001) study of the
Italian evidence agrees in general terms with
the results of Mussi’s (1995) earlier survey.
Of the Italian sites considered as important
occurrences that might date to the upper
Matuyama or the early part of the Brunhes,
none has yielded solid evidence for human
occupation prior to the Isernia–Notarchirico
time range (but see Milliken, 1997–1998).
This means that there is no evidence for an
occupation prior to the Middle Pleistocene
and even prior to the appearance of Arvicola
faunas (cf. Piperno, 1999; Sala, 1999).
If the Iberian evidence really is an excep-
tion, a chronological ‘‘loner’’ (yet), how are
we to weigh this evidence? Are the finds
traces of a late Lower Pleistocene circum-
Mediterranean settlement that gained
demographic momentum (and archaeologi-
cal visibility) around 500,000 years ago,
with the increase in population density lead-
ing to an expansion into new areas? Or are
the TD6 hominid remains the markers of a
Lower Pleistocene hominid incursion into
southern Europe that just failed to survive?
The current archaeological and palaeonto-
logical evidence is too meagre to choose one
of these options. Some scholars, however,
suggest that at least two successive disper-
sals of hominids into Europe can be distilled
from the archaeological and palaeontologi-
cal record (cf. Carbonell et al., 1999b;
Milliken, 2000; Otte, 2000). They see the
more substantial settlement, from about
the Isernia–Notarchirico–Miesenheim time
range, as the result of an influx of new-
comers who carried a toolkit different from
that of the first settlers, referred to as Homo
antecessor (Bermúdez de Castro et al., 1997;
Carbonell et al., 1999b). In this view, the
first settlers’ toolkit was characterised by the
dominance of a core and flake technology
and the absence of handaxes (e.g. Carbonell
et al., 1999b). Indeed, handaxes appear on
the European scene only at about 500–
600 ka, i.e., about one million years after
their first appearance in Africa (Asfaw et al.,
1992) and also considerably later than in
Israel, at ‘Ubeidiya and Gesher Benot
Ya’aqov (cf. Goren-Inbar et al., 2000).
Some see this as evidence for a successive
dispersal of two different cultural traditions
in Europe, with Lower Pleistocene Mode 1
‘‘people’’ preceding Acheuleans and a later
Acheulean wave displacing Mode 1 groups
(Carbonell et al., 1999b). There are some
problems with such a model though, for
instance arguments over the age of many of
these sites (see above, and Villa, 2001), the
disputed artificial character of some of the
key assemblages (see above), the variability
between so-called Mode 1 assemblages
(compare Atapuerca TD6 with Fuente
Nueva-3), and the large and well known
variability between artefact assemblages
from 500 ka onwards (Villa, 2001). Various
workers (e.g., Dennell & Roebroeks, 1996;
Villa, 2001) have also stressed the small size
of some ‘‘Mode 1’’ assemblages, which
makes an attribution to technical tradition
very unreliable: TD6 has yielded 268 arte-
facts (including unmodified pebbles and
chunks, cf. Carbonell et al., 1999a), while
Fuente Nueva-3 and Barranco León yielded
each about 100 artefacts, mostly flakes and
small flaking debris (Oms et al., 2000).
Paola Villa has taken up this very point
recently in a survey of the Italian and West-
ern European evidence, concluding that the
        
evidence for a greater antiquity of core-and-
flake industries is ambiguous, and that also
the interpretation of biface and non-biface
industries as designating different groups of
hominids lacks a sound archaeological basis
(Villa, 2001).
In summary, a short chronology still
applies to Europe north of the large moun-
tain chains of the Pyrenees and the Alps.
When these parts of Europe were first
colonised, occupation took place rapidly,
geologically speaking. The Iberian evidence
suggests that there was at least some inter-
mittent presence around the very end of the
Lower Pleistocene in southern Europe, with
a (possible) hiatus of 200,000 to 300,000
years separating the TD6 activities from
a more substantial human presence
starting around the Isernia–Notarchirico–
Miesenheim time range, i.e., about half a
million years ago [this situation shows a
remarkable similarity to the record from
Atlantic Morocco, as described by Raynal
et al. (1995): while some traces of occu-
pation may date from just before the
Brunhes–Matuyama boundary (Thomas
1-quarry, Level L), the main part of the rich
Acheulean sequence at Casablanca, for
instance, dates from the second part of the
Middle Pleistocene]. Virtually all workers
agree that the European archaeological
record changes significantly after 500 ka BP,
a communis opinio which is certainly one of
the positive results of the past decade’s
debate (e.g., Roebroeks & Van Kolfschoten,
1998; Carbonell et al., 1999b; Turner, 1999;
Villa, 2001). Sites increase in number in the
Mediterranean, while north of the large
mountain chains of the Pyrenees and the
Alps the first unambiguous traces of human
occupation appear. It is clear, finally, that in
order to set limits on speculations about the
when and how of colonisation processes we
need better chronological frameworks than
those at our disposal now. The discrepancies
between various sets of dating evidence
should form the point of departure for refin-
443
ing the timetable of Europe’s earliest settle-
ment, as it is in these problematic cases that
the largest progress can be made. While
many workers suppose that Europe may
have seen various waves of Lower and
Middle Pleistocene immigrants (cf. Goren-
Inbar et al., 2000), it is only through a
careful, multidisciplinary approach towards
the chronology of sites that eventually such
separate phases of colonisation will become
archaeologically visible.
Continuity of occupation?
For the time being, there is little evidence to
suggest (or contradict) that Europe saw a
continuous presence of hominids from the
end of the Lower Pleistocene onwards. A
better case can be made for a continuous
presence from about 600–500 ka. Studies of
the environmental tolerance of the earliest
Europeans north of the large mountain
chains have shown that these hominids were
present in a wide range of environmental
conditions, from full interglacial up to and
including cold climatic conditions and more
open environments, as testified by the evi-
dence from the loess sequence at Kärlich,
Germany (Roebroeks et al., 1992). These
various environmental settings do not
include the coldest phases of glacial–
interglacial cycles and likewise do not imply
that the whole of Europe saw a continuous
presence of hominids in the Middle Pleis-
tocene. Hominid numbers may have fluctu-
ated significantly in a given area from one
century to the next, especially for hominids
near the margins of ecological tolerance.
While the northern areas may have seen
an ebb and flow of human presence (cf.
Gamble, 1986), southern parts of Europe
could have functioned as refuges during the
most severe parts of the glacials. As feasible
as this possibility may sound, is not observ-
able in the archaeological record, where
continuity of human presence is difficult to
argue anyway, especially for the time range
444 . 
at stake here, with all its problems of
chronological resolution.
Recent developments in the field of Nean-
dertal studies can possibly be of help here.
According to workers like Arsuaga et al.
(1997b) and Hublin (1998), the typical
Neandertals of the last glacial were the result
of an accretion phenomenon, beginning in
the middle part of the Middle Pleistocene,
around 500 ka, if not earlier. The major cold
stages of the last half million years increased
the climatic stress on the initial colonisers
and resulted in the isolation of European
hominids and the evolution of cold adap-
tations, already thought to be visible in the
approximately 500,000 years old tibia from
Boxgrove in southern England (Trinkaus
et al., 1999). Added to the founder effect
following an initial colonisation of Europe
by small populations, climatic fluctuations
may have produced genetic drift episodes
resulting in fixation of derived features and
hence a decrease of variability through time
(Maureille, 1994; Hublin, 1998; see also:
Hewitt, 2000). Middle Pleistocene homi-
nids and the Neandertals are increasingly
seen as an ancestral-descendance sequence
of populations without rupture of reproduc-
tive continuity (Arsuaga et al., 2000; but see
also Hawks & Wolpoff, 2001). Hence, mor-
phological studies suggest a continuous and
isolated presence of hominids on the basis of
a kind of allopatric speciation process.
Recent genetic studies of the Neandertal
type specimen from the Feldhofer Grotte in
Germany come to a somewhat comparable
conclusion (Krings et al., 1997, 1999).
These studies indicate that Neandertals did
not contribute mtDNA to modern humans,
at the same time not ruling out the possi-
bility that Neandertals contributed other
genes to modern humans. Using the
mtDNA sequence from the Neandertal
specimen, Krings et al. (1997) estimated the
age for the most recent ancestral sequence
common to the Neandertal and modern
human mtDNA sequences at 550,000–
690,000 years BP. A second study of the
mtDNA sequence of the Neandertal type
specimen yielded an estimate of 465,000
years, with confidence limits of 317,000 and
741,000 years (Krings et al., 1999). Such
estimates are to be treated with caution, as
they rely heavily on the calibration point
of the chimpanzee–human divergence and
have errors of unknown magnitude. How-
ever, they strongly suggest that the age of the
common ancestor of Neandertal and mod-
ern human mtDNAs is four times greater
than that of the common ancestor of mod-
ern human mtDNAs. More important here,
they unambiguously situate the most recent
common ancestor of Neandertal and mod-
ern human mtDNAs in the first half of
the Middle Pleistocene, a date that agrees
very well with the occupation history data
reviewed above.
The archaeological record shows that
Middle Pleistocene hominids were present
in a large range of environments, over large
(but not all—see below) parts of Europe
from about half a million years ago, and we
have many sites from the middle parts of the
Middle Pleistocene onward (Roebroeks
et al., 1992; Gamble, 1995). Combined with
the genetic data and the accretion model
with its focus on the development of
European endemicity, I interpret this as
suggesting that from about half a million
years ago, Europe saw a continuous occu-
pation by—occasionally very small and rather
isolated1—groups of hominids. From about
half a million years ago Europeans, as rep-
resented by the Mauer and Boxgrove fossils,
gradually developed into early Neandertals,
such as those from the Sima de los Huesos
1
No matter how important a chronologically signifi-
cant geographical isolation may have been in this Nean-
dertalisation process, contacts between Europe and
other parts of the Old World must have existed. This is
indicated by the—in Pleistocene terms—almost con-
temporaneous appearance of the Levallois technique in
Europe, Western Asia and Africa at around 300 ka BP.
If not genes, at least techniques were freely exchanged
around that time.
        
at Atapuerca, Spain (Arsuaga et al., 1997a),
and ultimately to the ‘‘classic’’ forms from
the last glacial (it remains to be established
how the Atapuerca TD6 fossils fit into this
scheme of Neandertalisation, both in
chronological and in morphological terms,
cf. Hublin, 1998).
With these results in mind it is legitimate
to address the question of how these settlers
managed to survive over almost half a mil-
lion years. What kind of information does
the archaeological record contain, what
can the inferred continuous presence tell
us about the behaviour of these early
Europeans? For a first answer to this ques-
tion I will turn to the archaeological record
of Europe north of the large mountain
chains.
A look at the northern temperate zone
(and beyond)
While making their way through Europe, the
first settlers were confronted with environ-
ments with shorter and shorter growing sea-
sons as they went north, and they had to deal
increasingly with the problem of the temper-
ate zone: the winter stop in productivity of
the environment. In more general terms, the
major problems of the northern temperate
environments were ‘‘. . . the lower biological
productivity of the land, the greater season-
ality and reduced vegetation season, the
great extremes in temperature, and the
greater expenditures of energy required to
maintain homeostasis and reproduce’’
(Geist, 1978: 271). In such settings hunting
was a strategy that must have increasingly
served an omnivorous primate (cf. Binford,
1981), forced to live largely on animal mat-
ter, including its own fat (Milton, 2000) [a
recent—extreme—example is provided by
Alaskan Eskimos, who get the biggest part of
their total daily energy intake from fat
(50%), 30–35% from protein and 15–20%
from carbohydrates, mostly glycogen from
meat (Ho et al., 1972)].
445
In the last two decades these early north-
ern hominids have been depicted in many
behavioural scenarios as scavengers, surviv-
ing on the leftovers of large predators, that
is, prior to the appearance of modern
humans on the European scene (e.g.,
Gamble, 1987). In a paper published in
1985, Lewis Binford gave a review of the
foregoing two decades of efforts at recon-
structing early hominid behaviour, in which
he paid considerable attention to the north-
ern temperate zone. Starting with the con-
sensus view that early man was a hunter, his
reading of the evidence led him to ‘‘. . . the
inevitable conclusion . . . that regular, mod-
erate to large mammal hunting appears
simultaneously with the foreshadowing
changes occurring just prior to the appear-
ance of fully modern man’’ (Binford, 1985:
321).
When Binford wrote his ‘‘Human ances-
tors: changing views of their behavior’’ paper
(1985), there was indeed little archaeologi-
cal evidence to suggest that Middle and Late
Pleistocene hominids were actively engaging
large mammals. Now, 15 years later, many
new finds and analyses of these, and of older
assemblages, along the lines developed by
Binford’s research unambiguously show that
(late) Middle Pleistocene and Upper Pleis-
tocene hominids were capable hunters of
large mammals (see for example Marean &
Assefa, 1999; Gaudzinski & Roebroeks,
2000). A large number of archaeological
studies has shown that European middle
palaeolithic prey animals included reindeer
from the south of France to Salzgitter
Lebenstedt in the north of Germany
(Gaudzinski & Roebroeks, 2000), bovids at
such sites as Biache-Saint-Vaast (OIS7,
Auguste, 1992, 1995), Coudoulous, La
Borde and Mauran in France (e.g., Jaubert
et al., 1990; Farizy et al., 1994) and
Wallertheim in Germany (Gaudzinski,
1995). Bratlund’s (1999a; see for a sum-
mary 1999b) detailed taphonomic analysis
of the fauna from the OIS5e travertine site
446 . 
of Taubach, near Weimar (Germany),
strongly suggests that the major part of the
woodland rhinoceros assemblage there
(mandibular MNI of 62) was the result of
deliberate hunting of 1–1·5 year old rhino
individuals. Bratlund discusses at length
other workers’ scepticism regarding rhino
hunting in the Palaeolithic; using archaeo-
logical, ethnohistorical and ethnological
information she concludes that this scepti-
cism is not based on empirical evidence but
rooted in the attitude of earlier European
colonial hunters towards these animals
(Bratlund, 1999a: 135–146). The best par-
allel to Taubauch in respect to species rep-
resentation and preservation is the OIS 7
site Biache-Saint-Vaast, mentioned above,
though the focus on age classes differs.
The environmental settings of these sites
are highly variable, from ‘‘cold’’ to warm-
temperate, from open subarctic type of envi-
ronments to full interglacial wooded habitats
(Roebroeks et al., 1992). The sites are situ-
ated in various landscape types, from dis-
sected limestone valley systems in southern
France via the montane regions of the
German Mittelgebirge up to the loess plains
of northern Europe. The large size of many
of the prey animals from these sites, such as
the bovids at the OIS7 site Biache-Saint-
Vaast, is strongly suggestive of full-fledged
cooperative forms of hunting (cf. Geist,
1978: 274–278).
Independently of the archaeozoological
studies mentioned here, studies of the stable
isotopes from Neandertal skeletal remains
strongly suggest that they were top-level
carnivores, animal protein constituting an
important part of their diet (cf. Bocherens
et al., 1999, 2001; Richards et al., 2000).
The evidence for hunting in the Lower
Palaeolithic is not as clear cut as for the
Middle Palaeolithic, though it remains to be
established whether these differences really
reflect behavioural differences or whether
we are simply dealing with sample biases
and processes of preservation as suggested
by the evidence from a few sites, and
especially from Schöningen in Germany
(Thieme, 1997, 1999). The Schöningen site
has become well known through the discov-
ery of a series of wooden throwing spears
which date to about 300,000–400,000 years
ago. These were found amidst the concen-
trated remains of approximately 20 horses,
with almost no bones from other species
present, and together with a few dozen
flint tools as well as a large number of flint
chips, probably the result of resharpening
of tools (Thieme, 1997, 1999; personal
communication, 2000). The faunal remains
are even more exciting than the spears, as
they yield precious evidence on subsistence
strategies of the earliest Europeans unknown
as yet from any other Middle Pleistocene
site. An ongoing taphonomical study of the
assemblage by B. Voormolen (Leiden)
shows that a large number of the horse
bones display cutmarks, and that many
bones have been processed for marrow
extraction. Voormolen’s preliminary results
on body part representation and the virtual
absence of carnivore gnawing marks strongly
suggest that the horse remains are the results
of hominid hunting activities (Voormolen,
personal communication, 2000). The
Schöningen finds were recovered over an
area of approximately 1000 m2, from the
borders of a former lake, which hominids
may have used to ‘‘disadvantage’’ the horses
before killing them at close range with the
spears (L. R. Binford, personal communi-
cation, 2000). The high sedimentary rate
environment, where sedimentation was both
rapid and calm, enabled a perfect preser-
vation of hominid activities. Schöningen is
very clearly an exception, but in this sub-
sistence discussion such an exception carries
more weight than the ‘‘normal’’ type of
Lower Palaeolithic sites, such as the ones
available when Binford wrote his ‘‘Look at
the northern temperate zone’’ (1985: 315):
sites from where we do have indications
for some human interference with faunal
        
remains, though mostly in small numbers
only, and usually accompanied by a ‘‘nor-
mal’’ background fauna with abundant evi-
dence for carnivore involvement. Sites as
Swanscombe (Binford, 1985) and Hoxne
(Stopp, 1993) in England, Kärlich-Seefer
(Gaudzinski, 1998) and Miesenheim (E.
Turner, 1999) in Germany, and Torralba
and Ambrona (Santonja & Villa, 1990;
Villa, 1990) in Spain have complex tapho-
nomic histories, with material introduced by
various agents and accumulated over some-
times longer periods of time than seems to
have been the case for the Schöningen
site under consideration here. Even at high
rate sedimentation sites such as Boxgrove
(Roberts & Parfitt, 1999a), interpreted as
containing evidence for hunting of large
mammals such as horse and rhinoceros
(Roberts & Parfitt, 1999b), the archaeologi-
cal evidence is often not unambiguous
(McNabb, 2000).
Apart from a few open air sites, evidence
from cave sites also suggests that hunting
might have been common practice in the
Lower Palaeolithic, as can be inferred from
Moigne & Barsky’s (1999) study of the
fauna assemblage from Level L of the Caune
de l’Arago at Tautavel, southern France,
which is dominated by reindeer remains,
with an MNI of 40. They interpret the
reindeer as hunted and butchered, with the
bones of adult individuals systematically
processed for marrow extraction.
If, as I believe, one can generalise the
Schöningen evidence for the whole of
Europe around 400,000 years ago, this
implies that by the mid part of the Middle
Pleistocene occupants of Europe were ca-
pable and active hunters of large mammals.
At least by the time we see the first occu-
pants of Europe north of the Pyrenees and
the Alps, these hominids must have solved
the overwintering problem of the temperate
zone by hunting and eating animals that had
solved the overwintering problem long
before the arrival of the first Europeans.
447
The archaeological data for this statement
need to be reinforced, though, by detailed
publication of the faunal assemblages from
relevant sites, Schöningen and especially
Boxgrove, where important taphonomic
issues still need to be clarified. However, the
conclusion fits very well with the ecological
studies of Geist, who paid considerable
attention to the possible adaptations of the
first colonisers of the northern temperate
environments, and in fact predicted the
specific ‘‘top-level carnivore’’ (Richards
et al., 2000: 7663) adaptation, which a wide
range of recent studies have come to
acknowledge:
Man could not sustain himself by gathering
throughout the year. . . . There was no room
for a slow, herbivorous ape. There was also
an abundance of carnivores roaming about,
even in winter, such as cave lions, wolves . . .
cave hyenas and wolverines. Only the super-
carnivore niche is open, since the aforemen-
tioned predators concentrate preferentially
on medium-sized and small-bodied ungu-
lates . . . because ungulates of large size,
such as moose—but also horses, and pre-
sumably bison, mammoth and woolly
rhinos—are dangerous, capable opponents.
. . . The only niche open is that of the
supercarnivore, who can despatch very large,
slow, dangerous herbivores, and also the
large carnivores themselves (Geist, 1978:
281–282).
It is, finally, worthwhile mentioning here
that intrinsic nutritional factors may have
limited—at least seasonally—the extent to
which hominids could rely on these animal
products for calories (e.g., Speth, 1987,
1991). Total fat levels in temperate and
northern latitude large ungulates may drop
by late spring to 2–3% (Speth & Spielmann,
1983), and during cold seasons recent
hunter-gatherers may focus more specifi-
cally on the acquisition of fat or carbohy-
drates, while high protein food resources,
especially lean meat, can take on a very
marginal role or even be avoided entirely
(Speth, 1987).
448 . 
Contextualising the archaeological
evidence
How can we go further from the inference
that from about half a million years ago
hominids were hunting large mammals in
Europe? What kind of information can we
possibly distil from these observations on the
lives of these earliest colonisers and the
broader social setting within which these
hunting activities were situated? I suggest
following Aiello’s (1998) important lead and
taking an exploratory look at other disci-
plines dealing with the history of our species,
in an attempt to contextualise the archaeo-
logical data with the independent results of
other approaches and to develop new
insights on—or at least set limits to specu-
lations about—the social life of the earliest
Europeans.
For instance, as we know from animal
studies, the first settlers’ greater dependence
on hunting must have had consequences for
many aspects of their lives, including the size
of their home ranges: among primates
higher quality diets tend to be associated
with larger home ranges, and carnivores
have much larger ones than herbivores and
omnivores. Also carnivore ranges are larger
in high altitudes than in lower ones (Geist,
1978, passim; cf. Gamble & Steele, 1999).
This is related to the general trophic pyra-
mid of any environment, and together with
hunter-gatherers ethnographic data show a
general increase in the area of land exploited
as dependence on hunting increases: moving
north, resources tend to become more spa-
tially segregated along a gradient of decreas-
ing temperature, and the average distance
moved per residential move of hunter-
gatherers tends to increase with decreasing
temperature (Kelly, 1995). As hunting
becomes more important towards the poles
and most mammals need larger territories to
support themselves in colder latitudes, total
areas exploited in general increase consider-
ably. This suggests that the first Europeans
north of the large mountain chains were
hunters who operated over large areas.
There is, however, no archaeological evi-
dence on the specific size of these hypo-
thetical ‘‘large’’ areas. The small amount
of data we have for raw material transfers in
the European Lower Palaeolithic (Féblot-
Augustins, 1997) indicate that raw material
movements were generally short, and can
only provide minimum estimates of the
areas over which early hominids operated.
The database is somewhat better for the
early phases of the Middle Palaeolithic,
where we see transported lithics connect
such different regions as the northern Euro-
pean plains with the German Mittelgebirge
from at least OIS6 onwards, with stone
artefacts being moved over distances of
more than 120 km as the crow flies
(Roebroeks et al., 1988; Féblot-Augustins,
1999; Gamble & Steele, 1999). However,
these Middle Palaeolithic data also provide
only minimum estimates.
A very important contextualisation of the
hunting data is afforded by the so-called
Expensive Tissue Hypothesis. Developed by
Aiello & Wheeler (1995; see also Aiello,
1998), this hypothesis focuses on the large
brains of modern humans and states that
study of the cost of these large brains can be
very informative about the evolution of the
human niche. Human brains constitute on
average only 2% of total body weight, but
they use about 20% of all energy. With
young children the situation is even more
extreme: in the final phases of pregnancy the
growth of the foetal brain consumes about
70% of all energy afforded by the mother,
and in the first year after birth the brain still
needs about 60% of the child’s total energy
expenditure. Aiello & Wheeler posit that the
way in which humans can maintain these
costly large brains without also having high
basal metabolic rates is simply by reducing
the size of one of the other expensive tissues,
the gastrointestinal tract, the gut. This is
only possible by adopting a higher quality
        
diet, which would release the constraints on
encephalisation (Aiello & Wheeler, 1995).
As detailed by Aiello (1998), the relation-
ship between the energy requirements for
the growth of the brain and a high quality
diet ‘‘. . . is likely to have had additional
profound implications for hominin life-
history and social behaviour and, therefore,
for the human niche’’, encompassing
changes in gender relationships.
Women, especially, paid a high price
for the increase in brain size, becoming
increasingly dependent upon the help of
others—grandmothers, males—in the pro-
curement of high quality food for themselves
and their offspring (cf. O’Connell et al.,
1999). The increase in brain size occurring
after 500 ka BP would have put both
females and juveniles under increased
energetic stress, and it is around this time
that increased cooperation between caring
females and high quality food—in this
context: fat-rich game (cf. Speth &
Spielmann, 1983; Speth, 1990; Kelly,
1995)—provisioning males is predicted by
the hypothesis.2 It is also around this time
that the archaeological record, as I have
discussed above, contains the first evidence
for hunting of large mammals, as illustrated
by the Schöningen evidence. Though the
morphology of the Schöningen spears
resembles to a high degree that of the
current Olympian sportswoman’s javelin
(Rieder, 2000; Steguweit, 1999), Aiello’s
(1998) line of reasoning indicates that males
might have been the more common users of
these weapons.
The Expensive Tissue Hypothesis’ predic-
tions about female–male cooperation are not
2
Such a ‘‘bringing home the bacon’’ scenario needs
to be put in the context of nutritional studies, though.
For instance, as detailed by Speth (e.g., 1991), preg-
nant women may have to obtain up to 70% of their
daily calories from nonprotein sources; high protein
intakes would especially have posed a problem during
winter and early spring in the northern latitudes, with
pregnant women being a vulnerable segment of the
population.
449
easy to test, as there are relatively little data
on this topic. However, a recent statistical
analysis of the rich Middle Pleistocene
hominid material from the Sima de los
Huesos at Atapuerca (Arsuaga et al., 1997a)
corroborates to some degree the hypothesis’
predictions, by showing that the degree of
sexual dimorphism of this group of about 30
individuals—mostly young adults—is com-
parable to that of modern human popu-
lations (Arsuaga, 2000). This is important
evidence for a ‘‘modern’’ type of male–
female relationships in the middle part of the
Middle Pleistocene. While the Sima de los
Huesos material dates from about 300 ka
BP, more specific information relating to the
sexual division of labour comes from studies
of Late Pleistocene Neandertals only, and
even there the evidence is very limited.
Analysis of Neandertal humeri suggests that
males and females used their arms differ-
ently, with males routinely engaged in activi-
ties that needed considerable strength from
their right arms (Ben-Itzhak et al., 1988).
Differences in traumas of upper body and
head between male and female Neandertals
form another possible signal of a sexual
division of labour (Berger & Trinkaus,
1995).
While the (not uncontested: Hladik et al.,
1999) Expensive Tissue hypothesis offers a
valuable tool for contextualising the hunting
evidence and interpreting it in terms of the
social behaviour of the early Europeans,
Tooby & DeVore (1987) have likewise
developed an interesting behavioural
ecology-perspective on the social setting of
hunting. Hunting of large game is a boom-
or-bust activity. The variability in hunting
success and the fact that, unlike vegetable
resources, meat comes mostly in chunks in
excess of what an individual capturer can
readily consume, indicates that there must
have been a strong evolutionary selection for
‘‘food sharing, food exchange and risk shar-
ing through deferred reciprocation among
the larger social group’’ (Tooby & DeVore,
450 . 
1987: 224). In Winterhalder & Smith’s view
it was only with the evolution of reciprocity
or exchange-based food transfers that it
became economical for individual hunters to
target large game (Winterhalder & Smith,
2000). And for sharing the positive and
negative outcome of hunting activities
between independently foraging subunits to
take place, people had to have meeting
places: ‘‘Such a meeting area to exchange
food makes sense if the supply is irregular:
either more (in the case of success) or less
(in the case of failure) than the hunters
would need for themselves’’ (Tooby &
DeVore, 1987: 224). Hunting made food
sharing necessary, while food sharing made
hunting feasible.
Active cooperation between individuals
seems also indicated by the hunting activi-
ties themselves, as already pointed at above.
Compared to most other hunting animals,
humans are equipped with limited olfactory
and auditory capacities, humans cannot out-
run most game, and have to outsmart prey.
As discussed at great length, for instance by
Geist (1978: 308–310 and passim), modern
hunters are successful by virtue of knowing
the animals and the various traces they leave
behind in the landscape, and by using this
knowledge of animal behaviour as a ‘‘tool’’
(cf. Liebenberg, 1990). It is obvious that
knowledge of animal behaviour must have
likewise played a key role in successfully
hunting large mammals in the Palaeolithic,
especially with the limited technologies at
stake here. Even if experiments suggest that
the Schöningen throwing spears were superb
hunting weapons usable up to a distance of
about 25 m (Steguweit, 1999; Rieder,
2000), the most important ‘‘tool’’ must have
been an extensive knowledge of a wide range
of animal behaviour. Combined with a keen
insight in a variety of other factors such as
diurnal changes in wind directions—human
scent triggers the immediate departure of
most game animals—that knowledge ena-
bled them to get close enough to a large
mammal, considerably less than 25 m, for
some certainty of a hit. As stressed by
Guthrie (1997) such stalking activities are
more difficult by an order of magnitude out
in the open, away from tree-stands in the
forest or forest edge. Indeed, many workers
have emphasised the difficulties and com-
plexities of hunting, for instance Geist (1978
and passim) and Frison:
Animal procurement requires a body of
knowledge that comes only with long-term
involvement in predator–prey relationships
and familiarity with animal behavior under
all conditions and at all seasons of the year.
The factors that control the success or failure
of any animal procurement event are com-
plex and ever-changing. Every animal
species and individuals within the species
demonstrate unique behavioral characteris-
tics depending on internal conditions of sex,
age and animal condition and on external
conditions of time of year, weather, topo-
graphic features, and vegetative cover. Most
of these conditions can and often do change
rapidly: for example, a decrease in rainfall
and/or deep winter snow affects the avail-
ability of grasses, which subsequently affects
the location of the animals and ultimately
determines the optimum procurement
strategy (Frison, 1998: 14,578).
Hunting the variety of species we now
know from Lower and Middle Palaeolithic
contexts, in a variety of situations ‘‘. . .
required considerable experience, quality
education, and years of intensive practice.
From what we know about hunting today,
we can see that such hunting was very
difficult’’ (Guthrie, 1997: 107; see also
Geist, 1978). Hence, again, in all likelihood
there was a strong evolutionary stimulus on
cooperation, and on sharing information
between the older members of a group and
the young—and judging from the evidence
on their life history pattern these hominids
may have had a prolonged pattern of matu-
ration similar to that of modern humans
(Bermúdez de Castro et al., 1999), while a
        
recent study of the Boxgrove tibia suggests
that archaic humans may have reached
quite high ages (Streeter et al., 2001). The
‘‘quality education’’ needed to become an
expert Pleistocene hunter could not do
without a complex form of information
transmittal interaction in which the tran-
scendence of the here and now, a release from
proximity (Rodseth et al., 1991) played a
key role.
This brings me to the last exercise of this
exploration in contextualising the hunting
data, i.e., to a scenario for the social origin
of language, based on comparative mor-
phology and the ethology of primates, which
fits well with the implications of the archaeo-
logical evidence for hunting in the Middle
Pleistocene as discussed above. Dunbar’s
(e.g., 1992, 1993, 1996) research has shown
that there is a clear relationship between
the relative size of the neocortex of the brain
of primates, their ‘‘cognitive group’’ size
(Dunbar, 1998) and the time they invest in
physical contact, in grooming. The relation-
ship between neocortex size and group size
holds up in at least four other mammalian
taxonomic groups, an important support for
what has become known as the Social Brain
Hypothesis (Dunbar, 1998), according to
which primates’ relatively large brains arise
from the information-processing demands
of their complex social systems. Starting
from the relationship between the relative
size of the neocortex, group size and time
invested in individual contact, Aiello &
Dunbar (1993; cf. Dunbar, 1996) suggest
that by the time of the appearance of the
genus Homo, groups had become so large
that a nonphysical form of social contact—
language—became necessary for mainten-
ance of the group’s social cohesion. While a
rather rudimentary language may have
been at stake with early Homo, the Middle
Pleistocene encephalisation mentioned
above would indicate a group size not much
smaller than is common with modern
humans.
451
Using the human neocortex size ratio and
extrapolating a value for human group size
from the primate equation produces a value
in the order of 150 for modern humans.
This number agrees well with actual obser-
vations of the number of individuals humans
tend to associate with. As stressed by
Dunbar (1998), with modern humans these
larger groups are not (or only very rarely)
conspicuous as physical entities, but they do
have important social connotations for the
individuals concerned: 150 is the number of
individuals most people know as an individ-
ual on a personal basis, in fact the definition
used with other primates (Dunbar, 1998;
see also Steele, 1996). The observed
number of 150 also fits very well with
the results of Wobst’s (1974) computer
simulations to determine the minimum size
of a viable human breeding population,
which suggest that biological survival is
dependent on a group size of minimally
175.3 The actual size of such ‘‘cognitive’’
groups and prehistoric breeding populations
must have varied widely, in various complex
ways, determined by environmental factors
amongst others (cf. Kelly, 1995). The key
point here, though, is that there are good
arguments to infer a minimum ‘‘cognitive’’
3
While these larger groups are only conspicuous as
physical entities to the ethnographer during periodical
aggregations of the well-documented minimum bands
of about 25 individuals (Kelly, 1995), the archaeo-
logical visibility of any group size is an entirely different
problem. Recent studies of some classical Upper
Palaeolithic sites thought to reflect the remains of
aggregations of smaller groups into larger ones (for
ritual or other purposes), e.g. Dolni Vestonice and
Pavlov in Czechia, Mal’ta in Siberia and Oelknitz in
Germany, show that such earlier interpretations are not
unproblematic, and that the rich and large sites in fact
have to be reduced in the first instance to accumula-
tions of isolated depositional events, without any clear
relationship between the individual events. In other
words, small groups could have reoccupied much the
same locations in successive years, leading to the accu-
mulation of large amounts of material (respectively,
Verpoorte, 2000; Vasil’ev, 2000, S. Gaudzinski, per-
sonal communication, 2000). In this sense large group
sizes remain invisible ‘‘on the ground’’ until the estab-
lishment of the first sedentary communities with an
architectural record.
452 . 
group size of around 130 individuals by the
middle part of the Middle Pleistocene.
Following Aiello & Dunbar (1993), with
such a large group size it is impossible to
influence other members’ behaviour
through physical contact. Group integration
cannot be performed on a personal contact
basis along, and a modern way of grooming,
vocal grooming, language, must by then,
i.e., about half a million years ago, have
emerged for developing and maintaining
stable social relations. According to the
Social Brain Hypothesis language came
into existence as a kind of social glue that
enabled displacement in both place and time
(see Kay et al., 1998 for a comparable
‘‘chronology’’, based on a very different
dataset).
In summary, we can situate the archaeo-
logical indications for hunting of large mam-
mals within the context of the findings from
other disciplines, i.e., the physiological need
of a regular supply of high quality food and a
strong selection pressure on the sharing of
information and the products of hunting and
gathering activities. The three independent
sets of evidence reviewed here, (1) occu-
pation history, (2) archaeological evidence
for hunting and (3) comparative anatomy,
converge in the suggestion that the small
and mobile groups of individuals, which we
know from the archaeological record from
this period, were integrated into larger units
such as Dunbar’s ‘‘cognitive groups’’. The
individual members of such loose larger
groups collected information on the where-
abouts of other individuals and smaller
groups and the availability of food resources
as they roamed the landscape in a kind of
embedded learning, and shared this infor-
mation with other members of the larger
‘‘cognitive’’ group. Language made such
exchange of information about resources
and coordination of foraging activities poss-
ible. At its latest in the middle part of the
Middle Pleistocene, intensive, language
based cooperation between individuals and
between local small groups had become a
standard ingredient of the social behaviour
of early hominids. While Dunbar’s larger
groups are not visible ‘‘on the ground’’, in
the actual patterning of finds on the
archaeological site level, I suggest that their
existence is at its latest indicated by the
presence of hominids in temperate Europe
north of the large mountain chains,
half a million years ago. And so we have
gradually moved from a simple question
about the diet of these early settlers to the
question of what they did in between meals,
simply because both topics are inextricably
intertwined.
Hominid behaviour and the
chronology of colonisation
Where does this explorative exercise take us
in terms of the larger picture of the colonis-
ation of the Old World, or the long-term
trajectory of hominid adaptations? From
sites earlier than the first occupation of the
northern temperate zones we have no unam-
biguous evidence for hunting à la Schöningen
yet, and the fascinating evidence for canni-
balism at Atapuerca TD6 (Fernández-Jalvo
et al., 1999) is difficult to interpret in this
context. However, from the very first
appearance of stone artefacts in East Africa
onwards we see hominid interference with
bones in the form of cutmarks and conchoi-
dal fractures (De Heinzelin et al., 1999),
and somewhere in the more than 2 Ma
separating these earliest archaeological sites
(Semaw, 2000) from Schöningen, hominid
hunting à la Schöningen must have devel-
oped. If not for the freak conservation con-
ditions at Schöningen, the European earlier
Middle Pleistocene record would resemble
the earliest Palaeolithic record in terms of
human involvement with animals to a
high degree. While one can make a good
point that the Schöningen taphonomical
exception signals the behavioural rule
around 400,000 years ago, it remains to be
        
established when and where such behaviour
started and, again, possibilities should
not be confused with observables in this
context.
We run the risk of becoming trapped in
a chicken-or-egg problem when asking
whether hunting capacities enabled the
move to the north or whether northern envi-
ronments stimulated the development of
new adaptations. The evidence from Isernia
does show some hominid involvement with
the rich faunal remains there (Anconetani,
1999), but its exact character is too ambigu-
ous to be of help in choosing between these
two options, and the same applies to other
sites in Italy (Mussi, 1995). On the other
hand it is clear that in due time the archaeo-
logical record might yield information to
distinguish between cause and effect—for
instance, further excavations at Atapuerca
TD6, or the current studies of the well-
preserved faunal remains from Gesher
Benot Yaqov in Israel (Goran-Inbar et al.,
2000). However, for the time being it is
impossible to choose, and there is the
additional problem that we have little to no
idea how to evaluate the role of factors such
as demographic developments, adaptive
limitations, environmental changes or basic
physical obstacles in human range expan-
sion, certainly as long as the chronology of
that expansion is still unclear in so many
important aspects.
To illustrate the importance of a solid
chronology I can sketch a scenario starting
from the communis opinio that the Mediter-
ranean was colonised earlier than Europe
north of the large mountain chains, and that
the chronological differences between the
first appearance of hominids in both areas
may have been in the order of about
200,000–300,000 years. A comparable time
difference seems to exist between the first
traces of occupation in northwest and cen-
tral Europe and its easternmost parts, where
the first sites date from the later part of the
Middle Pleistocene (Gamble, 1995; Praslov,
453
1995—research bias may play a role here
though). If this is a correct chronology,
Europe was not settled more or less at the
same time, as implied by an earlier version
of the short chronology. Instead, the chro-
nology of occupation of the various regions
conveys an ecological message, as it enables
us to relate the history of colonisation to
the variations in key resources observed
within Europe (cf. Gamble, 1986, 1995). In
general, resources tend to become more
spatially segregated along a gradient of
decreasing temperature (Kelly, 1995: Fig-
ures 4–7). On a European scale, going from
the Mediterranean to the north of Europe
distances between resource patches become
greater, and are more easily stretched by
changes in the environment. Distances be-
tween feeding patches are critical for mobile
hominids, increases in these distances in-
creasing the size of their regular moves,
and the size of the areas over which groups
foraged. In the sketched chronology we see a
late Lower Pleistocene hominid presence
starting in the diverse environments of
southern Europe, where for Italy Mussi
(1995) has stressed the high variety of envi-
ronments over short distances, a factor also
called upon to explain the rich Lower and
Middle Pleistocene record of the Sierra of
Atapuerca. A few hundred thousand years
later hominids also show up in northern
Europe, even in the northern plains, in a
wide range of environments, from temperate
woodland conditions to colder steppic
settings as documented at Kärlich H
(Germany) (Roebroeks et al., 1992). I have
related their presence and aspects of their
adaptive strategies to large group sizes and
hence to the large spatial scales of their net-
works. For later phases of the Palaeolithic,
raw material studies have made abundantly
clear that in the Middle, as well as in the
Upper, Palaeolithic raw material transfers
tend to be consistently larger in the eastern,
continental parts of Europe than in the west
(Roebroeks et al., 1988; Féblot-Augustins,
454 . 
1993, 1997, 1999). The more continental,
harsher conditions increased the size of the
areas over which groups had to hunt and to
forage and hence the spatial scale of the
networks these hominids regularly main-
tained. The fact that a few hundred thou-
sand years seem to have passed between a
first presence in the western–central parts of
Europe and the first hominid presence in
the Russian plains, around OIS 9 (Praslov,
1995), might indicate that the spatial scale of
the networks initially did not match the en-
vironmental conditions of the continental
parts of Europe, with their extreme seasonal
temperature ranges. The differences in dis-
tribution and movement of animal resources
in these continental environments would
have required matching mobility strategies
and information gathering over larger areas
than necessary in the more western parts,
where the oceanic influence created more
mosaic environments (Gamble, 1995). We
can follow this chronological pattern into
Siberia, where even more extreme seasonal-
ity must have confronted the first settlers
with new challenges. As recently shown by
Goebel (2000), there is little evidence
that hominids entered Siberia during the
Middle Pleistocene. Early in the Upper
Pleistocene, middle palaeolithic populations
colonised Siberia’s southernmost latitudes,
but these were generally limited to the
heterogeneous environments of the south,
below 55 degrees north, as were the early
Upper Palaeolithic inhabitants of Siberia
(Goebel, 2000).
The palaeoenvironmental data from the
Lower Pleistocene site Dmanisi (Georgia),
at the gates of Europe, fits very well into the
chronology of the scenario outlined above.
The site was in the middle of a highly
productive ecotone rich in animal and plant
resources. The upland environment com-
bined diverse landscapes within short dis-
tances from each other, a vertical zonation
reflected in both faunal and floral remains
which indicate the prevalence of a warm and
moderately dry ‘‘Mediterranean’’ type cli-
mate (Gabunia et al., 2000a). Whereas in
earlier work the hominid remains were situ-
ated at the base of the sediments overlying
the basalt and were associated with more
forested conditions, hominid remains and
artefacts are now seen as intrusive into these
deposits (Gabunia et al., 2000b), which in
my reading implies that they date to periods
when the forested areas were considerably
reduced in size (see Table 2 in Gabunia
et al., 2000a). One site does not constitute a
pattern, but the Dmanisi evidence might
indeed be read as indicating that ‘‘the range
expansion of hominids from east Africa was
associated with adaptations to middle lati-
tudes where the upland environments
featured very diverse resources’’ (Gabunia
et al., 2000a:799).
This is certainly a too simple scenario,
which furthermore assumes that there was
continuity of occupation in Europe from the
Lower Pleistocene onwards. I have already
discussed above that the early Iberian traces
of occupation may have been the result of
one or more incursions that failed to
survive within Europe, and that the more
substantial settlement after about 500,000
could have resulted from another migratory
pulse into Europe. As stressed by Dennell &
Roebroeks (1996:540–541), hominids seem
to have occasionally moved into southern
Europe well before half a million years ago,
as and when conditions permitted, but with-
out living there ‘‘continuously’’. In this
scenario, the earliest hominids remained
confined to areas south of roughly 45 north,
whether in Iberia, Georgia or China. The
absence of Lower and early Middle Pleis-
tocene sites north of the Pyrenees and Alps
suggests that even if hominids were around
the Mediterranean perimeter from the late
Lower Pleistocene onwards, it required sig-
nificant changes in their behaviour to take
them north, across the many inland barriers
of Europe. Alternatively, the evidence from
areas outside of Europe seems to suggest
        
that there may have been other factors
involved in the colonisation of Europe.
Work in Atlantic Morocco indicates
that there too, a marginal hominid
(‘‘Acheulean’’) presence just prior to the
Brunhes–Matuyama boundary was separ-
ated from a substantial occupation in the
second half of the Middle Pleistocene
(Raynal et al., 1995). The similarity to the
European patterns suggests that there may
have been more at stake than adaptations to
northern latitudes per se. Potts’ Variability
Selection hypothesis offers an interesting
perspective here, as it relates both African
and European developments to the increas-
ing amplitude in environmental fluctuations
apparent in deep sea records and in terres-
trial regional records, especially from the
Middle Pleistocene onwards (Potts, 1996,
1998). Potts has pointed to the temporal
correlation between these environmental
changes and the spurt in hominid brain
expansion described above. In his view, this
correlation is one expression of the evolution
of complex anatomical, cognitive and social
factors capable of processing and respond-
ing to intricate and variable environmental
conditions: in short, the evolution of the
kind of adaptive flexibility indicated by
the various types of environments docu-
mented in the European Lower Palaeolithic
record.
Conclusion
This review of the past decade’s debate on
the earliest occupation of Europe concludes
that a short chronology still applies to
Europe north of the large mountain chains.
The southern parts of Europe must have
seen some earlier incursions, around the end
of the Lower Pleistocene, as shown by the
evidence from Spain. The absence of other
sites from that time range strongly suggests
that this was a marginal, intermittent occu-
pation. A substantial settlement of Europe
took place from about half a million years
455
ago. From that date onward, a good case
can be built for a continuous presence of
occasionally small and isolated groups of
hominids, gradually developing into the
classic Neanderthals of the last glacial. The
archaeological record suggests that these
Neanderthals were capable hunters of large
game, most probably from the very first
occupation of northern Europe onwards.
This higher dependence on hunting had
important consequences for the life of these
hominids, as I have shown in a contextuali-
sation of the archaeological data. At about
500 ka BP, increased forms of social
cooperation over large areas, exchange of
information between individuals within
larger groups, and in general forms of behav-
iour based on a ‘‘release from proximity’’
had become a standard ingredient of
the behavioural repertoire of these first
Europeans. At the same time the chronology
and ecology of the colonisation of the
various European regions suggest that the
size and the scale of these networks were
subject to long-term changes, which in the
long run may have enabled the occupation
of the eastern, more continental parts of
Europe—together with changes in other,
e.g., technological, domains, not discussed
here.
The settlement history of Europe may
have seen several pulses of immigration,
including the arrival of modern humans
about 40,000 years ago. The geographical
impact, i.e., the size of the areas settled,
must have been dependent on various
factors such as the adaptive capacities of the
immigrants and their demographic make up.
This last factor especially is extremely diffi-
cult to grasp in an archaeological context. It
seems clear beyond any doubt, however,
that around half a million years ago both
demographical momentum and behavioural
capacities enabled a substantial colonisation
of large parts of Europe, and a virtually
continuous hominid presence of more than
400,000 years.
456 . 
Acknowledgements
Thijs van Kolfschoten (Leiden) kindly
provided me with results of his biostrati-
graphical research, as did Boudewijn
Voormolen (Leiden) with his work on the
Schöningen fauna. An anonymous Journal of
Human Evolution Associate Editor and three
likewise anonymous reviewers made exten-
sive helpful comments on an earlier draft of
this paper, as did Sabine Gaudzinski
(Monrepos). I want to thank Juan Luis
Arsuaga (Madrid), Lewis Binford
(Dallas), Raymond Corbey (Leiden), Paul
Hennekens (Maastricht) and Alain Tuffreau
(Lille) for inspiring discussions on some of
the topics addressed in this paper. Robert
Franciscus (Iowa) provided me with some
very useful literature references. I am
grateful to Marco Langbroek, Boudewijn
Voormolen (Leiden) and a dozen Leiden
students for their active participation in a
2000 seminar series ‘‘Food for Thought’’,
where some of the ideas presented here were
tried out.
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