_UDK 903'1(4)''632'':575.113_
Documenta Praehistorica XXXIII (2006)
The Late Glacial ancestry of Europeans:
Combining genetic and archaeological evidence
Clive Gamble1, William Davies2, Paul Pettitt3, Lee Hazelwood2, Martin Richards4
1 Centre for Quaternary Research, Department of Geography, Royal Holloway University of London, UK
Clive.Gamble@rhul.ac.uk
2 Centre for the Archaeology of Human Origins University of Southampton, UK
3 Department of Archaeology, University of Sheffield, UK
4 Institute of Integrative and Comparative Biology, Faculty of Biological Sciences, University of Leeds, UK
ABSTRACT - Chronometrie attention in the Late Glacial of Western Europe is turning from the dating
of archaeological cultures to studying how the continent was re-populated at the end of the last ice
age. We present results from a survey of all available radiocarbon determinations (the S2AGES data-
base) which show that when calibrated, and compared to the GRIP stratotype of climatic events, the
data can be interpreted as five population events in the 15ka prior to the onset of the Holocene. The
fine-grained climate record provides an opportunity to study the impact of environmental factors on
a human dispersal process that not only shaped subsequent European prehistory, but also the gene-
tic makeup of modern Europeans. The population events have implications for archaeologists and
molecular geneticists concerning the timing, direction, speed and scale of processes in Western Euro-
pean demographic history. The results also bear on the role of climatic forcing on the expansion and
contraction of human populations and in particular the correlation of ice core and terrestrial re-
cords for the onset of warming in the North Atlantic.
IZVLEČEK - Kronometrična pozornost se v poznem glacialu zahodne Evrope odmika od datiranja ar-
heoloških kultur k proučevanju ponovne poselitve kontinenta ob koncu zadnje ledene dobe. Predstav-
ljamo rezultate pregleda vseh dosegljivih radiokarbonskih sekvenc (podatkovna baza S2AGES), ki ka-
žejo, da lahko datume, potem ko jih kalibriramo in primerjamo z GRIP stratotipom klimatskih dogod-
kov, interpretiramo kot zaporedje petih populacijskih dogodkov v času 15ka pred pojavom holocena.
Podrobno strukturiran klimatski zapis nam omogoča proučevanje vpliva okoljskih faktorjev na pro-
ces človekove razselitve, ki je sooblikoval evropsko prazgodovino in genetski zapis modernih Evro-
pejcev. Populacijski dogodki so pomembni za arheologe in molekularne genetike, ki se ukvarjajo s
časovnim usklajevanjem, smerjo, hitrostjo in obsegom procesov v zahodnoevropski demografski zgo-
dovini. Rezultati kažejo na pomen klimatskih pritiskov pri širjenju in krčenju človeških populacij in
še posebej na korelacijo med lednimi in kopenskimi zapisi ob začetku segrevanja v severnem Atlantiku.
KEY WORDS - Radiocarbon; Calibration; Late Glacial; Settlement patterns; mitochondrial DNA;
GRIP; Archaeological Taxonomic Unit
Introduction
A major population expansion occurred in Western
Europe during the Late Glacial (15-11.5ka CAL BP)
as the OIS2 ice sheets retreated and unglaciated
areas in the north became available for re-settle-
ment. Phylogeographic analysis using molecular evi-
dence assigns 60% of the European mitochondrial
DNA lineages (Richards et al. 2000), and an even
higher proportion of West European Y-chromosome
lineages (Semino et al. 2000), to a population bot-
tleneck prior to an expansion from southwest to
northern Europe (Torroni et al. 1998; Torroni et
al. 2001; Achilli et al. 2004; Rootsi et al. 2004; Pe-
reira et al. 2005). A potential sig-
nal of the bottleneck has also been
detected in patterns of linkage dis-
equilibrium in the autosomes (Reich
et al. 2001).
Population estimates (Bocquet-Apel
and Demars 2000) based on the
changing distribution of archaeolo-
gical sites indicate an increase in
the Western European metapopula-
tion, excluding Iberia, from 9000 to
40 000 persons with the correspon-
ding occupied areas augmented from
0.55Mkm2 to 1.12Mkm2. The demic
expansion is associated with the Ma-
gdalenian, a time-space archaeologi-
cal taxonomic unit (ATU), which is
found in Western Europe from c.22
to 13ka CAL BP and is known for its
abundant cave and mobiliary art.
However, the temporal sub-divisions
of this ATU, based on artefact type
fossils and stratigraphic sequences,
remain problematic (Laville et al.
1980; Thevenin 1995) such that the
timing, direction and pattern of the Late Glacial de-
mic expansion are imprecise. Moreover, estimates of
the age of the timing of this expansion derived from
a molecular clock model for mitochondrial DNA to
be c.16ka BP (Torroni et al. 2001) rely upon a num-
ber of assumptions and need to be substantiated by
independent radiometric means.
To address this imprecision, and to examine the
correspondence between archaeological and genet-
ic evidence for a population expansion, we present
here results from S2AGES, an OIS2 database of radio-
carbon determinations from Europe, the Near East
and North Africa. The sub-sample described here
comprises 2255 determinations from 1200 archaeo-
Fig. 1. Late Glacial Western Europe showing sampling regions,
the two geographical refugia and probable expansion routes.
logical assemblages in Western Europe (Fig. 1) (Pet-
titt et al. 2003). Figure 2 presents, using CALPAL
(Weninger and Jöris 2004), the calibrated radio-
carbon determinations against the GRIP 818 curve
for Iberia, France and North Central Europe.
We interpret the changing frequencies of calibrated
determinations as a proxy for the timing and direc-
tion of demic expansion as well as relative levels
of human activity between regions. This method,
using dates-as-data, is an established technique to in-
vestigate a process such as population dispersal into
un-occupied habitats (Holdaway and Porch 1995;
Ross 2001). We recognise five major population
events for Late Glacial Western Europe (Tab. 1).
Population event	Settlement pattern	Dominant settlement type	Phylogeography	GRIP Stratotype	GRIP Ice core years BP
i. Refugium	Dispersed	Sheltered	Low population size	LGM - GS-2c	25ka - i9.5ka
2. Initial demic expansion	Pioneer	Sheltered and Open		GS-2b - GS-2a	i9.5ka - i6ka
3.1	Main demic expansion 3.2	Main demic expansion	Residential Residential	Sheltered Open	Founder effect and expansion	GS-2a GI-ie	i6ka - i4.7ka i4.7ka - i4ka
4. Population stasis	Nucleation	Open		Gl-id - GSia	i4ka - i2.9ka
5. Population contraction		Open		GSi	i2.9ka - ii.5ka
Tab. 1. Late Glacial population history of Western Europe as reconstructed from archaeological, radiocar-
bon and genetic evidence.
Population event 1: Refugium, 25ka-19.5ka BP
Existing refugium models for Western Europe pre-
dict that the lowest metapopulation and the small-
est inhabited area co-incided with the last glacial
maximum (= LGM 25-21.8ka) (Soffer and Gamble
1990; Strauss 2000a). By contrast, refugium is used
here to describe the geographical distribution of po-
pulation (G refugium), and the contraction, irrespe-
ctive of geographic extent, of the size of the meta-
population (M refugium) within Western Europe.
Iberia, with radiocarbon deter-
minations predominantly from
Cantabria and Portugal (Strauss
2000c), was the major G refu-
gium into which population
contracted throughout the LGM
and continued there until GS-
2b (Fig. 2). By contrast, human
presence was markedly less in
France during this long period
even though the Southwest,
Aquitaine, has often been pro-
posed as a major G refugium
(Jochim 1987; Soffer and Gam-
ble 1990; Demars 1996). North
Central Europe was never en-
tirely abandoned as shown for
example by the site of Wiesba-
den-Igstadt dated to GI-2 in
the Rhineland (Terberger and
Street 2002) but the calibrated
curve indicates a low, probably
intermittent, human presence
in this region.
When the M refugium is exami-
ned using the S2AGES data, we
find it lasted much longer than
the LGM, sensu stricto and into
GS-2b. Moreover, a radiocar-
bon database compiled for the
Stage 3 project (van Andel et
al. 2003) shows very low hu-
man presence in North Central
Europe for at least the 10ka
preceding the LGM in spite of
a further six Dansgaard-Oesch-
ger oscillations in this time in-
terval (Walker et al. 1999).
The Solutrean has been propo-
sed as the ATU of the LGM re-
fugium (Strauss 2000b) (Fig. 3). The typological dif-
ferences between the Solutrean and Badegoulian
ATUs are regarded by archaeologists as too great to
derive the latter from the former (Djindjian et al.
1999). As recently demonstrated by Terberger and
Street (2002) the Badegoulian is most closely related
to ATUs in Central and Eastern rather than West-
ern Europe. This suggests the spatially extensive use
of the French reindeer steppes by hunting parties
with predominantly dispersed settlement patterns
and whose population focus lay to the east, rather
than the south. The eastern origins of the Badegou-
Fig. 2. Overview of Late Glacial radiocarbon dating probabilities for
archaeological sites and assemblages, calibrated using CALPAL. The
GRIP data form the stratotype for the Late Glacial-Last Termination
event based stratigraphy. This recognises, post the Last Glacial maxi-
mum (LGM), two warm events (Greenland Interstadials 1 and 2 and
two cold events (Greenland stadials 1 and 2). Currently GS-2 is subdi-
vided into three sub-stages and GI-1 into five. In conventional terms
GS-1 broadly corresponds to the Younger Dryas, GI-1e to the Bölling
and GI-1c-1a to the Aller0d interstadials. Other authors follow GISP2
for the period 25-16ka CAL BP. N = number of calibrated radiocarbon
determinations in the frequency curve.
lian may be reflected in the distribution of mtDNA
haplogroups H and pre-V. These are inferred to have
an origin around 20-30ka BP either in eastern Eu-
rope or the Near East, and to have re-expanded from
southwest Europe circa 16ka BP (Richards et al.
2000; Torroni et al. 2001; Pereira et al. 2005). The
Y-chromosome cluster R1 shows a similar pattern
(Semino et al. 2000; Kivisild et al. 2003).
Settlement type in both Cantabria and Southwest
France during Population event 1 is dominated by
rockshelters as opposed to open sites. Within Popu-
lation event 1 we include the relatively short lived
population expansions from Cantabria (Solutrean)
and Central Europe (Badegoulian) into southern
France (Fig. 3). We expect that similar demographic
pulses were a regular feature of
long-term human refugia.
Population event 2: Initial
demic expansion 19.5ka-
I6ka BP
The radiocarbon determinations
in Figure 4 form the proxy data
to infer significant demographic
change during GS-2b and the
first half of GS-2a. In all areas
this is associated with Magdale-
nian ATUs (Fig. 4). The change
takes the form of an initial step
and plateau (Fig. 2) in all re-
gions outside the Cantabrian G
refugium before the major rise
in the frequency of calibrated
determinations during the sec-
ond half of GS-2a. This occurs
during Heinrich Level 1 (17.6-
l4.9ka BP) when very low sum-
mer sea surface temperatures
(SST) have been recorded in
core MD95-2040 125km off the
Portuguese coast (de Abreu et
al. 2003).
In both France and Iberia Popu-
lation event 2 corresponds (Fig.
4), to a pronounced peak in di-
rectly dated cave art. The num-
ber of sites is small (N = 14) but
it is noteworthy that an earlier
peak in France corresponds to
the Solutrean population expan-
sion we have identified (Fig. 3)
and which also corresponds to low summer SST in
the Iberian Atlantic during HL2 (24.3-23.1ka BP)
(de Abreu et al. 2003). Therefore, cave art may be
associated in some regions with small-scale demic
diffusion out of the G refugium. This contrasts with
earlier interpretations that linked the appearance
of cave art to increased population density as people
moved into refugia during the LGM and used art as
a means to establish territorial rights to key resour-
ces such as salmon runs (Jochim 1983).
We favour an alternative, first proposed by Housley
et. al. (1997), that this small demographic step infer-
red from the increased number of radiocarbon de-
terminations represents a stage of pioneer settle-
ment in the process of demic expansion as regions
Fig. 3. Population events 1 and 2 (LGM to GS-2b) in Iberia and
France. Calibrated curves are shown by ATUs. In Iberia the Solutrean
is well represented from the LGM through GS-2c. In France the relative
presence of the Solutrean, as indicated by the number of radiocarbon
determinations, is substantially less suggesting a small demic diffu-
sion north followed by population contraction back to the Cantabrian
refugium. The wider context for the development of the Badegoulian
(= Early Magdalenian) is therefore the contraction of population
back into Iberia after GI-2.
Fig. 4. Population events 3-5 (GS-2b to GS-1) radiocarbon dating pro-
babilities by ATU. Later Magdalenian = ATU1; Epipalaeolithic = ATU2;
Mesolithic = ATU3. It is expected that an audit of the dates by precision,
accuracy and their archaeological integrity will reduce the outliers in
earlier population events (Pettitt et al. 2003).
librated determinations in all
regions between 16ka and
14.7ka CAL BP. During this
1300 year period the dominant
ATU is the Late Magdalenian.
Furthermore, the sequence in
the radiocarbon curves (Fig. 5)
confirm earlier archaeological
studies and the interpretation
of genetic evidence that with-
in Western Europe demic ex-
pansion was from the south to
the north and associated with
a rapid increase in the metapo-
pulation (Demars 1996; Hous-
ley et al. 1997; Bocquet-Apel
and Demars 2000; Strauss
2000a).
Phase 3.2 of the main demic ex-
pansion is represented in two
of the three regional radiocar-
bon curves (Fig. 4) as a plateau
that lasts throughout GI-1e (=
Bölling interstadial). These 700
years partly coincide with a
small plateau in the calibration
curve (Weninger and Jöris
2004). This is not considered
significant because at this time
in southern Iberia the frequen-
cy of determinations continues
to rise (Fig. 5).
beyond the G refugium were explored, assessed and
utilised in a more systematic fashion and on a more
regular basis. This settlement pattern occurred over
much of Western Europe where the main population
focus now included southwest France as well as re-
gions in Iberia outside Cantabria/Portugal. Settle-
ment type now comprises both open and naturally
sheltered sites.
Population event 3: Main demic expansion
I6ka-l4ka BP
The principal Late Glacial demic expansion occurred
in two phases in Western Europe (Fig. 5) over a 2000
year period in GS-2a and GI-1e. In phase 3.1 (Tab.
1) there is a significant increase in the number of ca-
Following Housley et al. (1997),
we describe the archaeological
settlement pattern in Popula-
tion Event 3 as residential.
Large-size, open-air campsites are known (Strauss
et al. 1996) from Lake Neuchatel and the Neuwied
and Paris basins and these are matched by substan-
tial rock shelter occupations in the Rhine-Danube
watershed, the uplands of Southern Germany, Thu-
ringia and Belgium. Smaller-scale rock shelter occu-
pations are known from the periphery in the Bri-
tish Isles (Barton 1999) and are comparable in site
scale to the pioneer settlement of earlier Population
event 2. No doubt small open settlements extended
into other northern areas, such as Doggerland (Coles
1998), now inundated by the North Sea.
Archaeological evidence suggests two dispersal cor-
ridors for Population event 3 to the west and east of
the Massif Central (Thevenin 1995). The eastern cor-
ridor following the Rhöne-Saöne-
Rhine rivers is supported by evi-
dence for the long-distance trans-
fer of raw materials including Medi-
terranean shells and Baltic amber
(Floss 2000) and is studded with re-
sidential settlement sites (Street et
al. 2001).
Population event 4: Population
stasis, I4ka-12.9ka BP
The interpretation of this popula-
tion event as stasis rather than con-
traction illustrates the importance
of combining radiocarbon determi-
nations, when used as proxy data
for past demography, with archae-
ological information on settlement
patterns. The face-value interpreta-
tion from the calibrated curves (Fig.
5) is that after 14ka CAL BP popu-
lation declined in most regions of
Western Europe.
However, Figure 6 shows that when
the radiocarbon determinations
from Northern Europe (Fig.1) are
compared for two classes of sites,
open and rock shelter/cave, then
the reason for the decline in the
proxy population curve becomes
apparent. Naturally sheltered loca-
tions were dominant in both pop-
ulation events 2 and 3. However,
this settlement type did not form a
significant part of the continuing
occupation of the region. On the
contrary, open-air campsites conti-
nued in importance throughout
GI-1 and remained largely unaf-
fected by climate change until the
last sub stage GI-1a. We interpret
these frequency data as changes in
regional settlement patterns, for
example from dispersed to nuclea-
ted, rather than a decline in the size of the metapo
pulation.
Fig. 5. Population events 3-5 (GS-2b to GS-1) radiocarbon fre-
quencies by region. ATUs represented are Later Magdalenian, Epi-
palaeolithic and Mesolithic. As in Figure 4 the dates are unaudited.
Population event 5: Population contraction,
12.9ka-11.5 BP
The impact of GS-1 (= Younger Dryas) on population
size in northern Europe was considerable (Fig. 6)
and is confirmed by archaeological surveys (Strauss
et al. 1996). However, this northern area was nei-
ther abandoned nor used as infrequently as in Po-
pulation events 1 and 2 (Fig. 2). This suggests a
higher metapopulation for Western Europe than
the refugium minimum (Bocquet-Apel and Demars
2000) and from this deme came the Holocene/Meso-
lithic recovery and population growth.
Comparison with genetic data and the environ-
mental framework
Phylogeographic analysis of modern Europeans indi-
cates a major founder effect in the Late Glacial (Ri-
chards et al. 2000), with an age estimated from
using the mitochondrial DNA molecular clock to
circa 16 000 years ago (Torroni et al. 2001; Pereira
et al. 2005). The radiocarbon chronology presented
here provides the first independent assessment of
this estimate with calibrated determinations (Tab. 1,
Fig. 2). However, it is not yet possible to be precise
about the duration of the constriction itself, except
to suggest from the proxy radiocarbon data for re-
gional demography that the neck of the bottle was
particularly elongated. Estimates (Bocquet-Apel and
Demars 2000) that the metapopulation increased
from 9000 to 40 000 in the Late Glacial of Western
Europe, excluding Iberia, now have to be considered
as demographic growth, independent of in-migra-
tion, which occurred in Population event 3.1 over
an estimated 1.3ka.
The S2AGES proxy data for demic expansion can be
considered in the light of two current interpretations
of the Greenland ice-core record and its significance
for the North Atlantic (Walker et al. 1999; Lowe et
al. 2001). At issue is the role of climate forcing in the
expansion of human populations. Ameliorating cli-
mate at the end of the European ice ages is believed
to drive the process of northward expansion of many
species from southern refuges
(Hewitt 1996; Hewitt 1999;
Willis and Whitaker 2000).
For this model to stand for
human expansion it is neces-
sary to interpret the radiocar-
bon and GRIP data as time-
transgressive. Multi-proxy en-
vironmental evidence from
terrestrial locations in north-
ern Europe (Walker et al.
2003) suggest this may be the
case. These data indicate war-
ming was underway by 15.5ka
CAL BP and that a time-trans-
gressive interpretation be-
tween palaeoclimate archives
at different latitudes is appro-
priate. If correct, the GRIP S18
signal lags behind climatic
warming and human, plant
and animal migration in the
oceanically controlled clima-
tes of Western Europe. Using Population event 3.1
as indicative of the effect of this warming on the di-
stribution of human populations we can estimate
the lag between climate change in Western Europe
and its register in GRIP. Cross correlation Corr (t) of
the climate and radiocarbon curves identifies a posi-
tive peak only in the Northern Europe and French
data. This indicates a significant response in the fre-
quency of radiocarbon determinations some 700
years before climatic warming took place as indica-
ted by GRIP. The width of the peak makes it diffi-
cult to ascertain precisely its statistical significance
but its simultaneous occurrence in the two regions
strongly suggests that it represents the same popula-
tion event, probably related to biomass changes un-
der climatic amelioration. Furthermore, the Iberian
data show a negative correlation with climate change
which is consistent with the continued refugium sta-
tus for this region.
However, an alternative model that expansion is not
directly linked to ameliorating climate and that the
two records are time-locked is supported by marine
evidence. High resolution records from an Iberian
ocean core (de Abreu et al. 2003) shows in-phase
oscillations with GRIP and GISP2 during the last
glacial cycle. Significantly Heinrich Level 1 dated
17.6-14.9ka BP represents the coldest sea surface
temperatures off Iberia in the last 70ka and coinci-
des with the major human expansion during popula-
tion events 2 and 3 (Tab. 1). Therefore, environmen-
Fig. 6. Open and sheltered sites from northern France and North Central
Europe (regions 6-9, Fig. 1) in population events 3-5.
tal forcing on population expansion was linked in-
stead to colder climatic conditions in Western Europe
and the earlier Solutrean pulse (Fig. 3) during HL2
confirms this.
One solution to the conflicting evidence over the
role of climate in human expansion might rest in the
model of southern refuges. Climate modelling using
sea surface temperatures (van Andel et al. 2003) and
evidence from the palaeobotanical (Willis et al.
2000) and faunal (Stewart 2003) archives points to
the existence of cryptic refuges during cold phases
north of Europe's continental divide. These refuges
may have been small in extent and limited in dura-
tion but would have acted as an attractor to human
populations with an extensive settlement network.
By comparison to human expansion, the contraction
we document in Population event 5 appears in phase
with the GRIP stratotype but is not related to any
Heinrich Level. This association supports Foley's
(1994) larger analysis of environmental change and
evolutionary events among hominins, where the pri-
mary influence of climate is on extinction rather
than speciation. We conclude that the expansion
and contraction of mobile Late Glacial populations
in Western Europe may be responding to fine-scale
climate change in similar ways and where, at these
smaller temporal and spatial scales, population dis-
persal and decline are analogous to speciation and
extinction (Gamble et al. 2004).
-ACKNOWLEDGEMENTS-
The compilation of S2AGES was supported by a grant
from the Leverhulme Trust (F/00180/I). Additional
funding from the British Academy for a Research
Readership and Research Grant was awarded to CG.
Nick Barton, Robert Foley, Mike Walker and Cheryl
Ross provided valuable comments.
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