_UDK 903i12/i15(4)''633/634'':575-17_
Documenta PraehistoricaXXXIII (2006)
Gene-flows and social processes:
the potential of genetics and archaeology
Julian Thomas
School of Arts, Histories and Cultures, University of Manchester, UK
Julian.Thomas@manchester.ac.uk
ABSTRACT - During the past four decades, genetic information has played an increasingly important
part in the study of the Mesolithic-Neolithic transition in Europe. However, there sometimes seems
to be a degree of disjunction between the patterns revealed by genetic analysis and the increasingly
complex social and economic processes that archaeology is starting to identify. In this contribution,
I point to the multiplicity of identities, subsistence regimes and patterns of social interaction involved
in the introduction of the Neolithic into northern and western Europe, and consider the implications
for genetic research.
IZVLEČEK - Genetske informacije so v zadnjih štirih desetletjih igrale pomembno vlogo pri študiju
mezolitsko-neolitske tranzicije v Evropi. Včasih se zdi, da prihaja do odstopanja med vzorci, ki jih
razkrivajo genetske analize in med kompleksnimi socialnimi in ekonomskimi procesi, ki jih prou-
čuje arheologija. V tem prispevku poudarjam mnogoterost identitet, prehrambenih režimov in vzor-
cev socialne interakcije, ki so bili vpleteni v uvajanje neolitika v severni in zahodni Evropo in oce-
nim, kakšne so implikacije za genetske raziskave.
KEY WORDS - archaeogenetics; demic diffusion; agency; frontiers; cultural hybridity
Scales and units of analysis
Over the past forty years, a fascinating dialogue has
been developing between archaeology and gene-
tics, specifically in relation to the question of the
dispersal of domesticated plants and animals into
Europe and its relationship with the movements of
human populations. This debate has often been mar-
ked by a degree of mutual confusion, owing largely
to the different temporal and spatial scales at which
the two disciplines operate, and the different ques-
tions that they address (Brown and Pluciennik 2001.
101). While genetics generally concerns itself with
the global or continental scale, archaeology is often
more focused on the regional and the local, with the
result that phenomena that are described at differ-
ent levels of magnitude may appear to contradict
each other. Some common ground is now beginning
to emerge, but from an archaeological point of view
it is especially interesting to ask whether the fine-
grained patterns that we think we can discern in the
evidence can be accommodated by the broader
sweep of the genetic information, or whether there
is a degree of dissonance between the two, whose
investigation might prove fruitful.
We are indebted to Albert Ammerman and Luca Ca-
valli-Sforza (1971; 1973) for initially stimulating
debate with their discussion of the expansion of
agriculture into Europe through demic diffusion, de-
veloped in the first instance in relation to radiocar-
bon dates from Neolithic sites, and later used as a
means of explaining the distribution of genetic mar-
kers across the continent. The model of farming com-
munities gradually expanding as their population
rose, fuelled by the productivity and reliability of
their subsistence base was explicitly differentiated
from migrationary arguments, in which communi-
ties are imagined to leave one area in order to set-
tle in another (Cavalli-Sforza 2002.82; Bellwood
2002.17). None the less, subsequent debate has ge-
nerally been framed in terms of the contrast between
demic and cultural diffusion; the physical spread of
farming societies versus the transmission and adop-
tion of Neolithic innovations amongst indigenous
hunting societies. While Ammerman and Cavalli-
Sforza have been careful to acknowledge the diver-
sity of the processes that might have been involved
in the Neolithic transition in Europe, and to recog-
nise a role for indigenous peoples in that process,
some of the geneticists who have followed in their
wake have tended toward a more categorical view
(Cavalli-Sforza 2003.303). Recent publications by
Barbujani and Dupanloup (2002), and by Chikhi
(2002), seem to imply that demic and cultural diffu-
sion are polar opposites, and that if the presence of
substantial Near Eastern genetic material attribut-
able to a post-Palaeolithic horizon can be identified
in Europe, the Neolithic must have spread into the
continent primarily, or exclusively, by population
movement.
At a philosophical level, many of the difficulties that
we encounter in trying to reconcile archaeological
and genetic evidence arise from the ways in which
we conceptualise past human communities and po-
pulations. Here, archaeology must take much of the
blame for developing and perpetuating the image
of human groups as self-contained and bounded
entities. It is instructive to remember that Gordon
Childe formalised the notion of the archaeological
culture precisely in the context of his study of the
early agricultural societies of south-east and central
Europe, in work that led up to the publication of The
Dawn of European Civilisation (Childe 1925) and
The Danube in Prehistory (Childe 1929). While
Childe was adamant that culture and race did not
coincide (1950.1), and even though he was an en-
thusiastic proponent of cultural diffusion, he none
the less instituted the expectation that a variety of
different aspects of human identity should be con-
gruent, so that definable distributions or assemblages
of artefacts could be identified as the material signa-
tures of 'peoples' who existed in the past (Jones
1997.17). I would argue that this is an understan-
ding that arises from the modern experience of liv-
ing within the nation-state, where political, ethnic,
linguistic and expressive entities are generally boun-
ded at the same level. This, then, grounds our expec-
tation that in prehistory we should be dealing with
neatly bounded social units (Thomas2004a.Ch. 5).
Of course, such an emphasis on bounded and inter-
nally homogeneous social wholes was by no means
exclusive to culture-historic archaeology: it was also
a hallmark of much processual archaeology (Brum-
fiel 1992).
Indigenism and evolutionism
One of the most innovative and attractive aspects of
Ammerman and Cavalli-Sforza's work was that it
challenged both the culture-historic image of boun-
ded ethnic groups in prehistory, and the processual
vision of autonomous communities in parallel evo-
lution, influenced only by environmental selective
pressures. As they point out, their initial papers
were written at the high point of anti-diffusionist
sentiment in archaeology, the time of 'the autonomy
of the south-east European copper age' and of 'Wes-
sex without Mycenae' (Cavalli-Sforza 2003.299).
More recently, Albert Ammerman has explicitly jux-
taposed the internationalist aspect of a perspective
based on demic diffusion, spreading agriculture
across the continent in a wave of advance, with what
he calls 'indigenism' (Ammerman 2003.4). Indige-
nism holds not only that each society is independent
and autonomous, but also that its development can
be attributed to internal processes, or to its specific
relationship to its environment. As such it chimes
with the culture-historic image of the bounded cul-
tural unit, and shares its affinity with nationalistic
approaches, which often hark back to a mythical gol-
den age of ethnic homogeneity (Gellner 1983.57).
Each nation has its own tribal ancestors, who were
responsible for their own independent domestica-
tions, and their own rise to statehood. So at a time
of heightened friction between the western and Isla-
mic worlds, Ammerman is absolutely correct to em-
phasise the importance of a Near Eastern contribu-
tion to the European genetic inheritance.
But on the other hand, there is the equal and oppo-
site danger of a crude social evolutionism which pre-
sents hunters and farmers as representatives of dif-
ferent stages of cultural development (Borić 2005;
Warren 2005; etc.). While demic diffusion is a popu-
lation model, the peril is of casting hunters and ga-
therers as passive and powerless in the face of the
oncoming Neolithic steamroller, while the farmers
are active and dynamic, even if only in demographic
terms (a position apparently adopted by Rowley-
Conwy 2004.97). Mesolithic populations are thus
understood as being 'absorbed', 'incorporated' or
'recruited'. Here again, we are faced with a problem
of scale, for while there is a virtue in describing and
explaining a pattern at a pan-European level, it
leaves little space for any consideration of agency
and contingency: outcomes that could have been
otherwise, and decisions made in the context of
inherited historical conditions. So the fate of hunt-
ing and gathering groups appears determined and
unavoidable. Similarly, by discussing the change
from Mesolithic to Neolithic at a continental scale,
such models run the risk of relying on stereotypical
and over-generalised models of hunters and farmers.
Yet as we are well aware, prehistoric Europe con-
tained colossal variation, between mobile and seden-
tary hunter-fisher-gatherers, and between tell-based,
longhouse and broad-spectrum forms of the Neoli-
thic (Zvelebil 2004.45). It may be that in the near
future the questions that genetic information will
be able to help us unravel are ones concerned with
the kinds of interactions that took place between
these diverse communities.
Of course, most authorities now maintain that the
Neolithic transition in Europe involved some combi-
nation of population movement and acculturation,
as in the framework that Marek Zvelebil describes as
'integrationism' (2002.397). Zvelebil suggests that
we should imagine a mosaic of different processes,
ranging from demic diffusion to leap-frog colonisa-
tion, frontier mobility, contact and exchange. I would
very much concur with him, but would wish to add
a further element to the argument. Zvelebil described
a series of different mechanisms by which agro-pas-
toral farming expanded. I would like to question
whether it was necessarily the same thing that was
expanding throughout, or whether the Neolithic did
not undergo a series of fundamental transformations
in the course of its translocation (see, for example,
Arias 1999.445). The key question is whether there
Fig. 1. Synthetic map of the first principal component of variation
in 95 classical genetic markers (from Cavalli-Sforza et. al. 1994,
copyright Princeton University Press).
was a single, constant causal motor for change
throughout, such as population growth. I am inclined
to doubt this. While in some areas a strong argument
can be made for density-driven expansion (cf. Van
Andel and Runnels 1995.498), in others it may be
that processes were at work to which agriculture was
no more than incidental. That is to say, in some parts
of Europe the Neolithic may have represented as
much an identity process as an economic package.
The indigenous component
One of the original reasons why the demic diffusion
model was considered pertinent to the spread of the
Neolithic was that the process was evidently so slow,
apparently progressing at around 1.1 kilometers per
year over a period of more than two and a half mil-
lennia (Ammerman and Cavalli-Sforza 1971.684).
By contrast, the cultural transmission of innovations
might be expected to have proceeded much faster.
But again, this neglects the agency of hunter-gathe-
rers, who appear to have resisted the adoption of
agriculture under some circumstances. In order to
spread from one community to another, a cultural
innovation needs to confer some perceived advan-
tage on the recipient. Agriculture is somewhat ambi-
valent in this respect: it brings the advantages of
higher yield and lower risk, but it might easily be
recognised as corrosive of a hunter-gatherer way of
life in restricting mobility and personal autonomy,
imposing greater labour investment, and transfor-
ming property relations. There is certainly good
ethnographic evidence of communities resisting eco-
nomic and technological change for social and cultu-
ral reasons (e.g. MacCormack 1978).
Now, of course, the first principal
component of protein genetic mark-
ers shows a gradient across Europe,
from south-east to north-west (Fig.
1), and Cavalli-Sforza explains this
in terms of the combination of demic
diffusion and recruitment or accultu-
ration, so that as the wave of ad-
vance swept westwards the expand-
ing Neolithic population would
have been characterised by a greater
and greater proportion of indigenous
genes (Cavalli-Sforza 2002.82). At
the peripheries, Neolithic people
would have been largely 'Mesolithic'
in genetic terms, and there appears
to be a level of agreement between
classical markers, mitochondrial DNA
and the Y-chromosome in suggesting roughly a 20%
Near Eastern Neolithic contribution to the overall
European gene pool (Underbill 2002; Richards
2003). However, this is another instance in which
the broad picture of human genetics and the fine
grain of archaeology rub up against one another,
because for a specialist in the study of the British
Neolithic the critical issue is that of what transpired
when these population processes washed up against
the Atlantic facade. At this point the Near Eastern ge-
netic inheritance becomes so etiolated as to be vir-
tually invisible. Neither classical markers nor Y-chro-
mosomes detect any 'Neolithic' presence in Britain,
yet it is of considerable importance to consider what
this non-presence represents, and what the proces-
ses might have been that gave rise to it.
For instance, in proposing a 'staged population inter-
action' wave of advance, Colin Renfrew (2002.100)
argues that a process of demic diffusion and the in-
corporation of hunter-gatherers might have contin-
ued uninterrupted beyond the point where any of
the genes of Near Eastern framers were present.
Martin Richards, however, indicates that this is not
compatible with the mitochondrial DNA evidence
(2003.153). On the other hand, there is the sugges-
tion that the classical marker plots represent palimp-
sests of a series of north-westerly population move-
ments throughout pre- and proto-history, which po-
tentially reduces the impact of Neolithic population
movement (Zvelebil 2002.385). Perhaps most inte-
resting is Richards' suggestion that mitochondrial
haplogroups J1a and J1b may relate to the very swift
dispersals of the Cardial complex and the Linear-
bandkeramik, at speeds swifter than those predicted
from demic diffusion (2004.152; Sykes 2003.323).
This is significant because it harmonises both with
the idea of the Neolithic transition as a patchwork
of diverse population processes, and with that of
alternating phases of rapid expansion and prolon-
ged standstill. Some of these episodes of expansion
may represent so-called leap-frog colonisation, which
I take to be comparable with the modified form of
demic diffusion described by Van Andel and Runnels
(1995.495) in the earliest Neolithic of Greece: the
selective and targeted colonisation of optimal areas.
However, we should be wary of assuming that such
processes were homogeneous, for while there may
be a case for such colonisation in Thessaly, sites like
Franchthi Cave may indicate a degree of continuity
from Mesolithic to Neolithic, although the extent of
this is open to debate (Thissen 2000). Similarly, the
very particular locational preferences demonstrated
by LBK settlements in central and western Europe,
and the extensive unoccupied areas between set-
tlement cells might be indicative of swift expansion
into favourable landscapes rather than slow, popu-
lation-driven movement (Lüning 1982.14; Bakels
1982).
Recently, Chris Scarre (2002.400) has suggested that
the Villeneuve-St-Germain sites of Normandy and
the Loire might represent small and dispersed pio-
neer agricultural groups, who moved into areas of
the landscape which complemented those occupied
by hunters and gatherers (Fig. 2) (although other
authorities argue that the VSG groups were them-
selves indigenous, and that their somewhat dispa-
rate long-houses represent copies of Danubian proto-
types: Jeunesse, pers. comm.). Scarre implies that
these communities were effectively absorbed by the
Mesolithic population, but it is arguable that both
groups contributed the subsequent formation of the
Cerny group (Cassen 1993; Scarre 1992). Similarly,
in east-central Europe, Nowak (2001.582) describes
a situation in which distinct enclaves of Neolithic
settlement existed in the period between 5600 and
4800 BC. These were migrant Linearbandkeramik
groups and their successors, who settled on areas of
highly fertile soil, creating 'small islands of farmers
in the immense sea of foragers' (Nowak 2001.590)
(Fig. 3). In a process analogous to that in western
France, it was only when the indigenous communi-
ties began to make extensive use of domesticates
and Neolithic artefacts that a more culturally homo-
geneous landscape began to develop, with the for-
mation of the TRB.
In Britain, similar arguments have been made con-
cerning the arrival of pioneer Neolithic groups (She-
ridan 2000; 2003; 2004, for example), but it is ar-
guable that they are far less convincing. In contrast
to the west French or east European examples, there
is no clear evidence for the coexistence of Mesolithic
communities and Neolithic enclaves, and the start of
the Neolithic was abrupt and uniform. Indeed, it is
the complete and sudden disappearance of the Me-
solithic assemblage in Britain that is the most remar-
kable aspect of the period, and it is my belief that it
can only be explained by a transformation that the
Mesolithic population were themselves instrumen-
tally engaged in. While this may not have involved
the movement of entire population groups as distinct
entities from the continent to Britain, it is extremely
likely that the exchange of personnel between groups
took place both during and prior to the transition.
For, while the manufacture of new stone tool types
and conceivably aspects of animal husbandry might
have been learned by indigenous people, the tech-
niques of potting and cereal cultivation are more
likely to have been transmitted by inter-marriage or
prolonged visiting and apprenticeship. While it has
conventionally been maintained that Britain and
Ireland had no contact with the European continent
during the later Mesolithic (e.g. Jacobi 1976), this
argument has been made on the basis of the morpho-
logical distinctiveness of microlithic assemblages. Yet
the degree of similarity of artefacts cannot be taken
as an index of the degree of interaction between so-
cial groups, while there is now evidence of the pre-
sence of domesticated cattle in Mesolithic Ireland
(Woodman and McCarthy 2003), demonstrating
that indigenous hunter-gatherers did have some deal-
ings with continental Neolithic groups (see Thomas
2004b for more detailed discussion). This means that
the transfer of domesticates and Neolithic material
culture into Britain is likely to have taken place in
the context of long-established relationships with
continental communities. In this connection, it is
important to remember that most of the Neolithic
groups that existed along the Atlantic coasts facing
the British Isles by 4000 BC (in Brittany, Normandy,
the Low Countries, Northern Germany, Denmark and
southern Sweden) were most likely indigenous peo-
ples who has adopted a new way of life through
acculturation or appropriation (Arias 1999.432).
Their connections with British hunter-gatherer socie-
ties are therefore likely to have been long-established.
Frontiers, interaction and hybridity
These arguments suggest that our investigation of
the genetics of prehistoric communities in Europe
needs to take more account of what happens when
personnel are exchanged between spatially juxta-
posed communities which are not bounded but per-
meable. After all, recent strontium isotope analysis
by Bentley et. al. (2003) on skeletons from Vaihin-
gen and other LBK sites in Germany suggests that
between 30 and 50% of the burials there were of
non-local origin, putatively Mesolithic people who
had married in to the community. This was happe-
ning not in the context of expansion, but of a period
of prolonged stasis in which some form of interac-
tion took place between Mesolithic and Neolithic
communities. It is these extended periods during
which the Neolithic did not expand that I want to
emphasise. Recently, Dušan Boric (2005) has offered
a cogent critique of the post-colonial assumptions
implicit in the notion of a 'frontier' between Mesoli-
thic and Neolithic groups in Europe. He argues that
such a model reinforces a dichotomous relationship
between two essentialised and ahistoric ideal types,
hunters and farmers, and that it implies a social evo-
lutionary scheme in which the replacement of forag-
ing by farming is an inevitable outcome. As an alter-
native, Boric presents an account of Lepenski Vir in
which multiple, complex identities existed side by
side (2005.99). Undoubtedly, Boric is correct to re-
ject the view that the Mesolithic and the Neolithic re-
Fig. 2. Villeneuve-Sain-Germain longhouses in northern France (from Scarre 2002/
presented two fixed and opposed
entities. However, it may still be
worth retaining the terminology
where we acknowledge that sixth-
fourth millennium BC Europe was
a patchwork of different kinds of
Mesolithic and Neolithic societies,
each of whose social and econo-
mic arrangements were continu-
ally open to transformation.
Wherever formally 'Mesolithic'
(that is, indigenous hunter-fisher-
gatherer) and 'Neolithic' (having
access to domesticates, ceramics
and polished stone tools) com-
munities came into spatial juxta-
position, the potential was creat-
ed for social interaction and cul-
tural innovation. The 'frontier'
between such groups is best seen
as a zone in which unpredictable
social and cultural exchanges
might take place: possibly vio-
lent (Gronenborn 1998), but also
potentially creative and transfor-
mational. It is unhelpful to think
of such interactions as taking place under the sway
of a 'law of cultural dominance' (Sahlins and Service
1960.69), in which the 'inferior' Mesolithic was
always influenced or dominated by the 'superior'
Neolithic. Rather, the encounter might affect either
group equally, through a process of hybridization or
creolization of cultural repertoires that were never
'pure' to begin with. Thus, while hunter-gatherer
groups in central and northern Europe adopted ce-
ramics and other Neolithic innovations, it is arguable
that in the post-Bandkeramik era, the various forms
of Neolithic that emerged incorporated elements of
a Mesolithic cultural inheritance (Arias 1999.445).
Simply because societies that used ceramics, polished
stone tools and domesticates eventually replaced ace-
ramic hunting and gathering groups across much of
Europe, we should not accept the teleological argu-
ment that this was the only possible outcome.
I suggest that through these episodes of interaction
at relatively long-lived 'frontiers', indigenous groups
acquired and assimilated new cultural and material
resources, but in addition the Neolithic was itself re-
peatedly transformed. There are a series of reasons
why such situations of contact and interaction might
persist over lengthy periods: where pioneer farmers
operated under conditions in which land and re-
Fig. 3. Poland at the time of the formation of the TRB, showing
'enclaves' of SBK/Lengyel Neolithic settlement in relation to Late
Mesolithic sites (from Zvelebil 2004).
sources were plentiful, and had no need to expand
further; where populations of hunter-gatherers were
dense, and operated elaborate subsistence regimes;
where physical circumstances restricted the expan-
sion of farming economies; and where economically
diversified Mesolithic groups would have perceived
no benefit in adopting new resources and techno-
logy or being incorporated into different cultural or
symbolic regimes.
There are at least three distinct areas in which we
can identify such phases of standstill and interaction.
The first would be following the following the estab-
lishment of the Neolithic in the northern Balkans,
where Esther Banffy (2004.57) points to prolonged
contact between Starčevo farmers, Körös communi-
ties who may have been of indigenous origin but
who combined the use of wild and domesticated
resources, and Mesolithic hunters in Transdanubia
during the earlier sixth millennium BC (see also
Whittle 2005). Out of this interaction emerged the
earliest Bandkeramik, which combined elements of
the Starčevo ceramic tradition with timber long-
houses. The longhouse implies the formation of an
entirely new mode of sociality: not the imposition
onto central Europe of a Balkan model, but some-
thing that developed in the protracted negotiation
between hunters and farmers, and which facilitated
the subsequent pioneer expansion into the loess
country of Europe north of the Alps and Carpathians
(Gronenborn 1999). Similarly, in the period follo-
wing the Bandkeramik expansion we can identify
parallel processes in western France and the North
European plain which began with the exchange of
ceramics, stone tools, livestock, furs, and presum-
ably also personnel between Neolithic and Mesoli-
thic communities, and culminated in the formation
of hybrid forms of sociality which drew on both tra-
ditions, with Cerny and the earliest TRB (Doman-
ska 2003; Nowak 2001; Zvelebil 2004.51). Just as
the formation of the LBK introduced an entirely new
social focus and forum in the shape of the longhouse,
so with Cerny and TRB it was monumental funerary
architecture that was central to a new kind of social
life (Midgley 2005.36). Again, I would stress that this
was a transformation of the Neolithic, arising out of
interaction, and introducing elements which had
simply not been there before.
It was the development of a new form of Neolithic,
in which kin relations were expressed through mor-
tuary monuments as much as in the domestic con-
text, that facilitated the final phase of the Neolithic
expansion in Europe: into Scandinavia and the Bri-
tish Isles. Here the process was more thoroughly
one of acculturation than elsewhere (Price 2000.
299). It is important to note that the significance of
domesticated species altered subtly at this point.
Where agriculture spreads by population movement,
we might expect domesticated plants and animals to
form an integrated food-production system, and to
represent staples. However, where we have popula-
tions of hunter-gatherers who have a stable econo-
mic base of their own, the initial occurrence of dome-
sticates beyond the 'agricultural frontier' is likely to
be as exotica and novelties. Interestingly, there is a
growing pattern of the identification of domesticated
cattle in pre-Neolithic contexts in southern Scandina-
via, southern Brittany, the Rhine Basin, the Alpine
foreland, northern Poland and Ireland (Zvelebil
2004.49; Woodman and McCarthy 2003). In the
Mesolithic context, the acquisition of a single dome-
sticated cow and its slaughter for communal con-
sumption might have had an appreciable impact on
local social relationships, in terms of status, prestige
and personal obligation. It is arguable that the pene-
tration of north-west European Mesolithic societies
by Neolithic systems of consumption and prestige
was one of the mechanisms that led to their trans-
formation. But as we have stressed above, it should
not be presumed that this was a one-way process,
for the acquisition of goods and raw materials, and
the incorporation of personnel from Mesolithic com-
munities would have held a transformative potential
for Neolithic societies.
Clearly, the societies that inhabited northern and
western Europe between the sixth and fourth mil-
lennia BC were diverse in terms of their various com-
binations of hunting, gathering, fishing, herding and
horticulture, their material culture, and their social
organisation. The contacts and relationships between
these groups will have been more complex still. The
articulation of social reproduction and social inter-
action will, under these circumstances, have resulted
in elaborate sequences of non-reversible historical
change. I have dwelt on all of this complexity because
each of these processes will have had their own de-
mographic consequences and correlates. The pat-
terns that we observe in the DNA evidence are the
outcome of these processes, overlaid with four mil-
lennia of further developments. There remains a
massive contribution that human genetics can make
to the study of this period, but it may be that it is
now time for it to address a finer-grained picture of
the Neolithic transition.
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