Plant subsistence and environment at the Mesolithic site Ta˚gerup,

Plant subsistence and environment at the Mesolithic site Ta˚gerup,
Veget Hist Archaeobot (2012) 21:1–16
DOI 10.1007/s00334-011-0299-x
Plant subsistence and environment at the Mesolithic site Tågerup,
southern Sweden: new insights on the ‘‘Nut Age’’
Mats Regnell
Received: 31 August 2010 / Accepted: 25 April 2011 / Published online: 7 May 2011
Springer-Verlag 2011
Abstract Pollen was analysed from a sediment sequence
collected in the close vicinity of the Mesolithic settlement
Tågerup, southern Sweden. Macroremains were also
retrieved from numerous samples taken at the site of the
archaeological excavations of Kongemose and Ertebølle
settlement phases, 6700–6000 B.C. and 5500–4900 B.C.
respectively. Plants and other organic remains were well
preserved in the refuse layers from the settlements
embedded in the gyttja. The pollen record includes no clear
indications of human impact on the vegetation during the
Mesolithic. The occurrence of charcoal particles and pollen
of grass and herbs associated with nutrient-rich soils are
contemporaneous with the Kongemose settlement. The
Ertebølle settlement phase, although characterised by
considerable dwelling activities less than a hundred metres
from the pollen sampling site, is scarcely seen in the pollen
data. Numerous finds of crushed dogwood stones from the
Kongemose phase, often partly carbonised, suggest that
these stones were used for the extraction of oil. Other
plants found in the Kongemose refuse layers that may have
been used are apples, cherries, raspberries, acorns and
rowan-berries. Based on the abundance of hazelnut shells
found at the studied site and in other studies of Mesolithic
sites in southern Scandinavia it is proposed that these
remains may testify to an important food supply rather than
just use as a supplement to animal protein. It is also
hypothesised that a regional decrease in hazel populations
Communicated by M. Latałowa.
M. Regnell (&)
Department of Physical Geography and Quaternary Geology,
Stockholm University, 10691 Stockholm, Sweden
e-mail: [email protected]
and thus hazelnut availability at the end of the Mesolithic
may have motivated the adoption of Neolithic subsistence.
Keywords Plant macroremains Pollen Mesolithic Environmental history Plant subsistence Hazelnuts
When literature on Mesolithic plant use in Northern Europe
is considered, three important aspects can be perceived.
First, pollen analysis alone seems to be inadequate to gain
specific information on plant use. Second, studies on plant
macroremains have, when wood working is not involved,
identified few intentionally used plant species. Third, hazel
nuts seem to have been the most important plant food
resource. Pollen analysis is a useful method to reconstruct
past vegetation and landscape development, but it has
obvious limitations if the objective is to investigate the past
environment at community or species level. This is apparent
for instance when plant usage is the focus. Plant macroremain analysis is a valuable method for supplying detailed
information on local vegetation and crucial for distinguishing plants that have been collected and utilized. There
are many reports on human disturbance of vegetation in
northern Europe during the Mesolithic from pollen analytical studies (e.g. Vuorela 1986; Kolstrup 1990; Regnell et al.
1995; Poska 2001) and on possible pre-Neolithic agriculture
(Edwards and Hirons 1984; Göransson 1988; Klassen 2004;
Poska and Saarse 2006). However, few of these studies
discuss the use of specific plant species and several arguments against Mesolithic agriculture in central and Northern
Europe have been advanced (Behre 2006). Mesolithic use of
specific species concluded from pollen analysis involves
hazelnuts (e.g. Simmons and Innes 1987; Huntley 1993) and
Veget Hist Archaeobot (2012) 21:1–16
ivy (Simmons and Dimbleby 1974). In previous investigations of Stone Age sites in southern Sweden a combination
of pollen and plant macroremain analyses has proved
rewarding (Göransson 1983; Regnell et al. 1995). The
results show, for example, that Mesolithic settlement,
although relatively extensive and of long duration, had a
limited impact on forest vegetation. Adjacent to the Mesolithic site Bökeberg III pollen analysis of a peat sequence
revealed small changes in tree and herb pollen frequencies,
although microscopic charcoal was abundant during periods
of settlement. It was assumed that the habitation was
established without any forest clearance or other disturbance
of the vegetation (Regnell et al. 1995). Retrieved plant
macro remains (excluding wood) from Scandinavian
Mesolithic sites include ca. 70 species (Regnell 1998). Most
of them represent contemporaneous vegetation at the
archaeological sites. It can be argued that a few species were
being collected and used, with hazelnut as the most obvious
example. However yellow water lily, dogwood (Andersen
et al. 1982), acorns and hawthorn have also been pointed out
as having been consumed (Grøn 1997).
This paper presents results from one of the sub-projects
within the archaeological West Coast Line project, which
was launched by the Swedish National Heritage Board in
connection with the construction of a new railway track. The
study area is situated in Scania, the southernmost province of
Sweden, at the outlet of the Baltic Sea into the Atlantic
Ocean (Fig. 1). The Mesolithic site Tågerup presented here
has been the focus of massive investigations. Adjacent to the
Mesolithic settlement, numerous samples from marine
deposits were studied using palaeoecological methods. The
results from the extensive archaeological excavations and
also some of the most important palaeoecological conclusions have been published in a series of books (e.g. Karsten
and Knarrström 2001, 2003; Svensson 2003). In this paper,
two sets of data are presented and discussed:
Pollen stratigraphical investigations of a sediment
sequence accumulated in close connection to the
Mesolithic site at Tågerup
Plant macroremains collected from a range of
contexts during the archaeological excavations.
In this study, pollen analysis was used to reconstruct
general vegetation changes around the settlements at Tågerup and plant macroremain analysis was used to identify
specific plants that might have been used by humans.
The aims within the study have been:
To reconstruct the vegetation around the settlements
and specifically to study human impact on vegetation
during the phase of significant human settlement
Fig. 1 Map of the study area and the investigated site Tågerup.
a Pollen analysed sites mentioned in the text and in Fig. 5: 1—
Hassing Huse Mose Bog, 2—Kurarp, 3—Lake Krageholmssjön, 4—
Lake Bjärsjöholmssjön; sites with Mesolithic plant remains mentioned in the text: 5—Bökeberg, 6—Huseby Klev, 7—Balltorp, 8—
Holmegard Mose; green area with settlements from the Linear Band
Pottery Group dated between 5500 and 4700 B.C. (from Lüning 2000
and Larina 2009). b Map with the sea level shown as 1 m above the
present average level, i.e. the assumed sea level at the end of the
Kongemose settlement phase (Regnell and Risberg unpub.). c Black
dots show the coring sites along the profile presented in Fig. 2. T1
The analysed sequence (pollen). Area of excavation shown with thick
line. Light blue area—excavated area with in situ remains from the
Ertebølle settlement phase. Dark blue—extension of the refuse layers
belonging to the Kongemose settlement phase
To provide information on the use of plants during the
To evaluate the importance of hazelnuts within consumption strategies.
Veget Hist Archaeobot (2012) 21:1–16
Geological and archaeological settings
Scania is situated at the boundary between the Fennoscandian archaic Baltic Shield and the central European
sedimentary basin. Geologically, the north-eastern part of
Scania belongs to Scandinavia and the south-western part,
where Tågerup is situated, to continental Europe. The
topography surrounding the site Tågerup is characterised
by flat land surfaces barely reaching 20 m a.s.l. Sand is the
most common Quaternary deposit along the coast; further
inland clayey till dominates. The stratigraphical distribution of Quaternary deposits is complicated by complex
vertical variations, e.g. thin accumulations of postglacial
sand often overlying clayey till. Moreover, bodies of silt
occur locally in otherwise sandy areas (Ringberg 1976;
Adrielsson 1984). This three-dimensional large variation in
lithostratigraphy explains the complexity of the hydrology
of the area and also the environmental diversity important
for plants and animals.
The minerogenic portion of the Quaternary deposits
mainly originates from the southern part of the Baltic,
which means chalk and limestone predominate. Around
Tågerup the soils are very fertile and the area is characterised by a rich agriculture (Erlandsson 1999), an aspect
that may be assumed to have been also important during
Tågerup is located where the small rivers of Saxån and
Braån converge in a larger stream. Both rivers can be
followed inland about 15 km and represent together a
drainage area of approximately 150 km2. Prior to the
confluence, the rivers run south and west of a hill (Fig. 1a),
where abundant stray finds from all prehistoric periods
have been found. The archaeological excavations at Tågerup were performed during three seasons in 1997–1999
along the southern rim of this hill, parallel to the course of
the present River Saxån (Fig. 1c). The investigations have
revealed Mesolithic occupations dated to two phases (all
dates below are given in calibrated years B.C.):
The Kongemose phase, dated to 6700–6000 B.C., is
represented by finds in aquatic and telmatic sediments, i.e.
in refuse layers deposited close to the contemporary shore
and below water at distances up to 20 m from the former
shore-line. Finds from marine gyttja include flint and bone
tools (some of them decorated), wooden leisters, prongs
and fish traps, and bone remains from humans and 48
animal taxa. At about 40–60 m from the former shore-line,
five graves and eight probable burials were found. One hut
from the Kongemose phase was found (Karsten and
Knarrström 2001, 2003).
The Ertebølle phase, dated to 5500–4900 B.C., revealed
the same type of remains as the older settlement phase,
although with each category generally in smaller numbers.
In addition to finds in gyttja, artefact-rich cultural layers on
dry land were found, including houses and huts with
strikingly preserved construction details. Surface finds
surrounding the excavated area at Tågerup indicate that the
promontory was also inhabited after the Mesolithic, and a
number of dates indicate activities at the site throughout
prehistory (Karsten and Knarrström 2003).
Materials and methods
Close to the excavated area, cores with a 10 cm diameter
Russian sampler were taken along an east–west profile
(Fig. 1c). In this area Quaternary deposits reach a thickness
of almost 20 m (Fig. 2). The deposits consist of fine and
coarse minerogenic late-glacial sediments, laminated minerogenic sediments representing an early Holocene riverbed, and Mid-Holocene transgression sediments. In the
central part of the profile a representative sequence (T1),
was extracted and sub-sampled at 10 cm intervals and
analysed for multi-proxy data. The pollen stratigraphy from
T1 is presented in this paper. Analyses of plant macroremains, together with siliceous microremains, sediment
chemistry and mineral magnetic parameters (Regnell et al.
2001), will be discussed in a forthcoming paper in which
sea-level changes will be the focus.
Pollen analysis
Samples for pollen analysis were treated according to
standard procedures (Berglund and Ralska-Jasiewiczowa
1986). The analyses were performed at 9320 and 9800
magnifications, and phase contrast was used when necessary. A minimum of 1,000 pollen grains per sample was
counted. In general, pollen identifications were based on
the pollen keys by Erdtman et al. (1961), Faegri et al.
(1989), Moore et al. (1991) and Punt et al. (1976). In
addition, the reference collection at the Department of
Quaternary Geology, Lund University was consulted.
Pollen taxonomy and nomenclature follows the pollen keys
mentioned above. The results are presented as percentages
of the sum of terrestrial taxa; aquatics, spores of Pteridophytes and Sphagnum, the alga Pediastrum and herbs of
wet soil (Typha latifolia, Sparganium erectum-type and
Cyperaceae) are excluded from the calculation sum. The
programs TILIA and TILIAGRAPH (Grimm 1990) were
used for calculation, drawing the diagram and zonation. To
facilitate the description and interpretation of the pollen
diagram in terms of vegetation changes, five local pollen
assemblage zones were distinguished (LPAZ T1:1–T1:5),
based on the CONISS cluster analysis (Grimm 1987) as
implemented by the TILIA program (Grimm 1990). Each
zone boundary denotes significant changes in the pollen
assemblages and hence represents major changes in
Veget Hist Archaeobot (2012) 21:1–16
Fig. 2 The litho-stratigraphy at
Tågerup T1. The section is
drawn along the profile shown
in Fig. 1c. Limits between
stratigraphical units
representing Late-glacial, Early
Holocene and Mid-Holocene
deposits are indicated by thicker
vegetation. A number of taxa with low pollen counts and of
little significance to the discussion have been excluded
from the diagram (Fig. 4).
Macroremain analysis
The macroremain analyses were performed on separate soil
samples from the cultural layers, on various constructions
from the terrestrial environment and from refuse layers in
marine, waterlogged, organic sediments. Sampling was
mainly performed in collaboration with the excavating
archaeological staff. No sampling grid has been used, i.e.
each sampling was decided on the basis of contextual
aspects. Sample volume was calculated in a graduated
beaker where a known volume of water was added. All
samples were diluted with 1–2% NaOH. Waterlogged
samples characterised by high organic content were diluted
for a longer time, and/or were heated to approximately
60C and washed through a 0.25 mm mesh sieve. Minerogenic soil samples were handled with a modified flotation
method. Diluted samples were placed in a 10 l bucket.
While constantly pouring water into the bucket that was
inclined with slight variations in angle, smaller minerogenic particles and organic material were decanted into the
0.25 mm sieve. Originally waterlogged samples were kept
in water after preparation, but minerogenic samples were
dried in room temperature. Macro remains were sorted and
identified under a binocular microscope at 96 to 980
magnification using the keys and atlases by Katz et al.
(1965), Berggren (1969, 1981), Beijerinck (1947), Jacomet
et al. (1989) and Anderberg (1994). The majority of samples that originated from minerogenic soils often contained
fresh seeds, rootlets, insects etc. These fresh specimens
were considered as recently incorporated in the soil and
were not regarded as sub-fossil finds, but their presence
was noted. Nomenclature and present distribution of plants
are according to Den Nya Nordiska Floran [The new
Nordic flora] (Mossberg and Stenberg 2003).
The chronology of the sediment sequence T1 was constructed from 11 AMS radiocarbon dates on macro remains
of terrestrial origin (Table 1). The plant remains used for
dating were retrieved from sequence T1, parallel to the
levels that were analysed for pollen. These samples were
prepared and analysed with the same methods as described
above. Dating was performed at the Ångström laboratory,
Uppsala University, and 14C dates were calibrated using
OxCal 4.1. A time-depth model was established applying a
linear interpolation through the individually plotted calibration curves (Fig. 3).
Veget Hist Archaeobot (2012) 21:1–16
Table 1 Radiocarbon dates from Tågerup 1 (T1)
Lab. no.
Depth cm
Cal. age yrs B.C. 1 r-range
Cal. age yrs B.C. 2 r-range
5,495 ± 80
4,448–4,413 (13.5%)
4,518–4,225 (89.6%)
4,406–4,316 (39.9%)
4,205–4,162 (3.2%)
C-age yrs B.P.
4,298–4,262 (14.7%)
4,130–4,112 (0.9%)
4,101–4,072 (1.6%)
4,780 ± 95
4,960 ± 90
3,650–3,501 (54.4%)
3,762–3,724 (2.1%)
3,428–3,380 (13.8%)
3,715–3,362 (93.3%)
3,912–3,878 (9.9%)
3,804–3,649 (58.3%)
3,964–3,632 (94.0%)
3,558–3,538 (1.4%)
6,140 ± 110
5,220–4,938 (68.2%)
5,320–4,796 (95.4%)
6,360 ± 75
5,468–5,402 (23.4%)
5,484–5,207 (95.2%)
5,388–5,297 (42.2%)
5,088–5,084 (0.2%)
5,241–5,232 (2.6%)
6,840 ± 75
5,792–5,657 (66.3%)
5,892–5,621 (95.4%)
5,651–5,646 (1.9%)
7,360 ± 75
6,354–6,294 (17.4%)
6,393–6,069 (95.4%)
6,266–6,202 (23.1%)
6,194–6,099 (27.8%)
8,565 ± 90
7,705–7,698 (1.8%)
7,936–7,927 (0.3%)
7,681–7,522 (66.4%)
7,916–7,900 (0.6%)
7,866–7,860 (0.2%)
7,840–7,454 (94.0%)
8,755 ± 80
7,952–7,658 (68.2%)
7,392–7,381 (0.4%)
8,200–8,110 (8.5%)
8,093–8,072 (1.6%)
8,066–8,039 (2.0%)
8,006–7,597 (83.3%)
7,875 ± 65
6,981–6,975 (1.1%)
7,030–6,874 (23.1%)
6,908–6,886 (4.5%)
6,866–6,596 (72.3%)
6,828–6,638 (62.6%)
8,040 ± 75
Results and interpretation
The age-depth model presented in Fig. 3 was used to
construct the time-scale of sequence T1 (pollen diagram,
Fig. 4).
Pollen-inferred vegetation development
The size of the area for which the local pollen assemblages
are representative is difficult to estimate since the studied
sediments were deposited in a bay of the sea. In addition,
the Braån and Saxån Rivers may have transported pollen
into the bay. Therefore, it is assumed that the pollen source
7,076–5,825 (68.2%)
7,180–6,691 (95.4%)
area is relatively large and that the pollen data therefore
represent mainly the regional vegetation.
In the following description, characteristic features for
each local pollen assemblage zone are indicated. Mesolithic and Neolithic periods are according to Brinch Petersen (1993) and Nielsen (1993) respectively.
Estimated accuracies for the stated ages below are in the
order of ±100 years (Fig. 3).
T1:1: 1,050–885 cm, ca. 7000–6500 B.C. This zone
corresponds to the Late Maglemose and Early Kongemose
cultures. The pollen spectra suggest a landscape dominated
by nemoral trees (Corylus, Ulmus, Quercus and Tilia).
Pinus was present during the early part of the period and
became more common in the later part. Ulmus was slightly
less common in the younger part of the period. Alnus was
Fig. 3 Time-depth model for the sediment sequence T1. The
calibration curve for each date (Table 1) is projected. A linear
interpolation of the age intervals with highest density of likelihood is
drawn (red).The three dates (Ua-26047, Ua-25941 and Ua-26051) are
considered to represent reworked, older material and are excluded
Veget Hist Archaeobot (2012) 21:1–16
from the time-depth model. Boundaries between pollen zones are
shown with hatched lines. The classical elm-decline as dated by Skog
and Regnéll (1995) is indicated by a thick line and a white circle. The
settlement phases at Tågerup are shown below the time axis
Fig. 4 Percentage pollen diagram of the sequence T1. The pollen zones are based on CONISS cluster analysis (Grimm 1987) implemented by
the program TILIA (Grimm 1990)
Veget Hist Archaeobot (2012) 21:1–16
abundant on peaty or waterlogged ground, together with
some Betula and Salix. Wetland communities, represented
by Cyperaceae, Sparganium erectum-type and Filipendula
were probably confined to shorelines. During the oldest
part of the period, highly variable values are found for taxa
such as Alnus, Corylus and Pinus. These fluctuating values
may be caused by redeposition of sediment and pollen
grains, perhaps in connection with a transgression of the
sea at the study site (cf. Digerfeldt 1975). Microscopic
charcoal particles are present in the middle part of the zone,
before the first settlement phase. These charcoal particles
should be interpreted together with occasional radiocarbon
datings of archaeological findings preceding the defined
Kongemose phase and thus suggesting activities at the site
before the main occupation.
T1:2: 885–775 cm, ca. 6500–6100 B.C. The zone corresponds to the Kongemose culture. The lower part of the
zone shows an increase in Quercus, Tilia and Fraxinus.
The landscape was dominated by a nemoral forest where
Corylus, Quercus, Ulmus, Tilia and Pinus were common
trees and shrubs. On waterlogged ground, Alnus and Betula
were abundant, but some Salix shrubs also occurred. The
zone corresponds to an increase of Artemisia, Chenopodiaceae, Poaceae undiff. \40 lm, Sparganium erectum-type
and Cyperaceae. These taxa may, if Poaceae undiff.
\40 lm is at least partly inferred as being Phragmites,
correspond to an expansion of communities close to the
shore. Pediastrum colonies are found in enhanced numbers
through the zone, possibly indicating a stronger influence
of freshwater from the small rivers Braån and Saxån. High
frequencies of microscopic charcoal particles are found in
the zone, correlating to the Kongemose settlement phase.
The occasional presence of Rumex acetosa/acetosella may
indicate either patches of open ground within the forest or
seashore vegetation. Decreases in Viscum and Hedera helix
and the presence of Populus also signify a more open and
disturbed forest.
T1:3: 775–565 cm, ca. 6100–5100 B.C. The zone covers
the Late Kongemose and Early Ertebølle cultures. Trees
and shrubs such as Corylus, Quercus, Ulmus and Tilia were
common. Pinus shows lower frequencies than in previous
zones. Alnus was the dominant species on wet soils, but
Betula and Salix shrubs may have occurred. During this
period, a reduction of open vegetation communities is
indicated. A continuous expansion of Quercus and Fraxinus can be seen as well as a decrease in Corylus frequencies. Microscopic charcoal particles occur only
sporadically, most likely indicating a reduced occurrence
of camp-fires. As a consequence of the change in disturbance regime, Pinus and grasses were disadvantaged and
Quercus and Ulmus could expand. In the upper part of the
zone there is an increase of Artemisia, Aster and Chenopodiaceae that may be an effect of an expansion of shoreline plants.
Fig. 4 continued
T1:4: 565–185 cm, ca. 5100–3600 B.C. The zone is
divided into two subzones where T1:4a (565–275 cm, ca.
5100–3900 B.C.) represents the later part the Late Ertebølle
culture and T1:4b (275–185 cm, ca. 3900–3600 B.C.) the
earliest part the Early Neolithic culture (EN A and B
according to Nielsen 1993). Just above the lower boundary
there is a decrease of Corylus from nearly 40–25%, which
is accompanied by an equivalent increase in Alnus. The
beginning of the zone, ca. 5100–4900 B.C., corresponds to
the most intense phase of Ertebølle settlement, as indicated
by radiocarbon datings of archaeological finds and chronologically significant artefact types (Karsten and Knarrström 2003). In the uppermost part of zone T1:4a, Corylus
increases again to similar values as previously. Paralleling
the changes in Corylus percentages described above,
Quercus and Fraxinus show increasing values in the lower
part of subzone T1:4a while other tree pollen maintains
comparable percentages through the subzone.
The lower part of subzone T1:4b is marked by the classical
Ulmus decline recognised in many areas of north-west Europe
that was dated to 3790–3745 B.C. in southern Sweden (Skog
and Regnéll 1995). It is hard to infer a precise timing for the
elm decline in this study, but the time-depth curve (Fig. 3)
suggests one of ca. 3900 B.C. The regular occurrence of Apiaceae, Compositae/Cichorioideae (Lactucae), Caryophyllaceae and Filipendula may represent an expansion of open
vegetation. However, the forest disturbance indicated by the
decline of broad leaved trees may not have favoured an
expansion of open vegetation since Betula appears to have
responded quickly as a pioneer tree expanding in the areas of
disturbed forest, maintaining a relatively dense tree cover,
however with an altered species composition. Open vegetation communities were probably still confined to shorelines.
Later, Quercus and Tilia may have replaced Betula. Microscopic charcoal particles are scarce in the sediment suggesting
that probably no camp fires occurred in the closest vicinity of
the study site. Therefore, it is considered that signs of human
impact in T1:4b are weak, which implies that the Ulmus
decline rather was a consequence of natural factors such as
climate change and elm disease (Peglar 1993; Peglar and
Birks 1993; Digerfeldt 1997). Ulmus and Fraxinus probably
regained their previous abundance in the later part of the
subzone, as indicated by pollen values as high as those prior to
the Ulmus decline. The structure and composition of the forest
were probably similar at the end of subzone T1:4a and at the
last part of subzone T1:4b. Open vegetation communities
probably had a restricted occurrence in the area.
T1:5: 185–80 cm, ca. 3600–3200 B.C. This zone represents the late Early Neolithic (ENC) and the earliest part of
Middle Neolithic (MNAI-II?). The pollen record reflects a
landscape dominated by a nemoral forest, where Quercus
and Corylus were abundant elements and Ulmus, Tilia and
Fraxinus occurred regularly. Acer was present, but it
Veget Hist Archaeobot (2012) 21:1–16
probably had a low abundance. On wetter sites, Alnus was
dominant, possibly intermixed with Betula. The pollen data
also indicates an increase in Corylus, Tilia and Alnus, and a
reduction of Quercus and Fraxinus in the uppermost part of
the zone. An increase in Artemisia, Filipendula, Plantago
lanceolata and Rumex acetosa/acetosella clearly indicates
the expansion of open vegetation. The occurrence of
Plantago lanceolata suggests that grazing might have been
introduced into the area, possibly as forest grazing.
Microscopic charcoal particles are not common in the
record which suggests that natural fires and forest clearance
using fire were not occurring in the study area.
Plant macroremains
Nearly 50 samples representing 36 litres of gyttja from the
refuse layers were analysed. The plant remains from the
layers ascribed to the Kongemose occupation phase are
listed in Table 2. The composition of the plant macroremains is dominated by aquatic plants. Fruits of Ruppia
maritima and Zannichellia palustris and oospores of Chara
sp. are most common. These plants, as well as different
species of Potamogeton, grow along shores of the Öresund
Strait today, while Najas marina is rarely found along the
Scanian coasts but is more common along the Swedish
eastern coast. Apium inundatum also has a limited occurrence along the Scanian coast today, but the plant was
found close to Tågerup in the 1980s (Weimarck and
Weimarck 1985). Today, it is mainly characteristic of
inland lakes, ditches and ponds.
The most numerous remains from plants characteristic of
shore-lines (telmatophytes) are from Phragmites australis
and Typha sp.; Glaux maritima is also common together with
Moehringia trinervia. A few remains of Schoenoplectus lacustris/Bolboschoenus maritimus and Tripolium vulgare
were found. All these plants grow today along the shores of
the Öresund Strait, although M. trinervia and S. lacustris are
more commonly found along lake shores. Thus, some plant
species from the Kongemose phase indicate aquatic and
telmatic vegetation typical of waters with lower salinity than
the modern conditions at the site. However, the occurrence of
these species might also be explained by transport from
upstream sources via the tributaries.
The wetland plants found at Tågerup probably derive
from wet or moist soils along the seashore or from similar
environments upstream. Alnus was the dominant wetland
tree and probably grew close to the sampling site. Internodes of Equisetum palustre and achenes of Eupatorium
cannabinum are relatively common in the Tågerup sediments and these species are also represented in the modern
surrounding vegetation. Other frequent remains are of
fruits of Urtica dioica and seeds of Juncus sp. All these
wetland species are still widespread in western Scania.
Veget Hist Archaeobot (2012) 21:1–16
Table 2 Plant macroremains from 49 samples from the refuse-layers
dated to the Kongemose phase at Tågerup. The finds are diaspores if
nothing else is specified
Finds (n)
Table 3 Summary of carbonised plant remains (diaspores if not
otherwise specified) found in terrestrial soil samples associated with
the Ertebølle phase at Tågerup
Grave Grave
A6504 A40186
Vol. (l)
Aquatic species
Apium inundatum
Cerealia indet.
Najas marina
Hordeum vulgare coll.
Potamogeton sp.
Ruppia maritima
Hordeum vulgare var.
Zannichellia palustris
Chara sp.
Wetland and shore line species
Alnus sp.
Alnus sp. (flower spindles, leaf scars etc.)
Bidens tripartita
Carex sp.
Crambe maritima
Equisetum palustre (internodes)
Eupatorium cannabinum
Galium uliginosum
Glaux maritima
Juncus sp.
Lychnis flos-cuculi
Lycopus europaeus
Moehringia trinervia
Persicaria hydropiper
Phragmites australis
Urtica dioica
Terrestrial species
Betula sp.
Chenopodium sp.
Cornus sanguinea
Corylus avellana (no. nutshell fragments)
Fallopia convolvulus
Malus sylvestris
Persicaria maculosa
Polygonum aviculare
Prunus avium
Rubus idaeus
Rumex acetosa
Quercus sp. (no. acorn fragments)
Triticum spelta
Triticum dicoccum/T.
Corylus avellana (no. of
nutshell fragments)
bulbosus (tuber)
Poaceae undiff.
Chenopodium sp.
Rumex acetosa
Stellaria palustris
Typha sp.
Schoenoplectus lacustris/Bolboschoenus maritimus
Tripolium vulgare
Silene nutans
Silene vulgaris
Sorbus aucuparia
Stellaria media
Poaceae indet.
Potentilla sp.
Of the terrestrial plants the highest numbers of finds are
from birch Betula sp., Quercus sp., Malus silvestris, Sorbus
aucuparia, Corylus avellana and Cornus sanguinea. Remains
from Chenopodium sp., Persicaria maculosa, Polygonum
aviculare and Stellaria media indicate nutrient rich soils
probably in the moist environment along the shore. Alternatively, the occurrence of such species at the site might indicate
more nutrient-rich conditions at and around the Mesolithic
settlement. In addition, pollen zone T1:2, that is contemporaneous with the Kongemose settlement phase, is characterised by an increase in Apiaceae, Artemisia, Chenopodiaceae
and Rumex acetosa/acetosella. These taxa also indicate
nutrient-rich soils and a situation with more open vegetation.
The finds of Fallopia convolvulus in Mesolithic depositions at Tågerup (Table 2) should probably be interpreted
as contamination from younger layers. F. convolvulus is
probably not native to Scandinavia or the British Isles
(Webb 1985) but was introduced during the Neolithic. The
species has previously been found at about a dozen Stone
Age sites in Scandinavia, and only one of these is Mesolithic, the rest belonging to the Neolithic (Regnell 1998).
The Mesolithic finds are from a coastal site in southern
Norway and although the stone tools and radiocarbon dates
are indisputably of Mesolithic origin (Østmo 1976) the find
description in the original report cannot rule out contamination. It should be pointed out that apart from the finds of
F. convolvulus there could be other examples of redeposition of younger remains into the Kongemose layers.
At Tågerup the gyttja refuse-layers from the Kongemose
settlement phase provided by far the largest amount of
plant remains. Additional plant remains were retrieved
from 519 samples originating from the terrestrial part of the
younger Ertebølle settlement phase. Table 3 summarises
the finds of carbonised plant remains from three houses and
three graves, all of Mesolithic origin. All terrestrial
deposits are related to the Ertebølle phase dating to c.
5500–4900 B.C. In relation to the large volume of soil
analysed, find concentrations were low. In House 2, for
example, 179 litres of soil were analysed but only 13 plant
remains were found. The presence of cereal grains can be
explained by post-depositional processes, as radiocarbon
dating of the grains show that they are contemporaneous
with an Iron Age settlement at a higher elevation. However, the presence of hazelnuts, although in low concentrations (maximum of 34 nuts in House 3), might be
associated with the Mesolithic remains.
This discussion is focused on the consumption of plants,
but other uses will also be mentioned. In such a context it is
very important to define what is meant by plant use and
how deliberate collection and usage of plants can be
inferred from palaeo-records. There are three major situations where archaeobotanical records can be interpreted in
terms of utilization of wild plants; (1) occurrences in very
large quantities, (2) presence in an environment in which
the plant does not belong naturally (i.e. appears as ‘exotic’), and/or (3) obvious signs of processing by humans.
Moreover, ethnological information usually provides
further understanding. Ethnological analogies are of course
important within many aspects of archaeology, and ethnobotany is perhaps an especially rich source of information on human habits and behaviour within the field of
The Mesolithic period is highlighted as the era of hunter-gatherers and therefore wild plants are integrated in
virtually all models and concepts dealing with the Mesolithic (e.g. Clarke 1976; Rowley-Conwy 1983, 1986; Price
1989; Zvelebil 1994). Food strategies during the Mesolithic
are described in a large number of publications, however
the references above put forward the idea that gatherers not
only made a passive use of what nature provided, but also
deliberately maintained plants of particular interest for
them and consciously transformed the landscapes/vegetation for various purposes. Pollen and charcoal analyses
from several sites in Europe suggest that people during
Early and Middle Mesolithic induced forest fires (Tolonen
1985; Patterson et al. 1987; Edwards 1990; Simmons
1996). Some authors consider fire during these periods as a
deliberate method of creating clearances that attracted
game animals and thus made meat a more achievable food
Veget Hist Archaeobot (2012) 21:1–16
component (e.g. Mellars 1976). Nevertheless, other studies
show that natural climate-induced fire was probably a
major factor in the forest dynamics of the Early and Mid
Holocene in southern Sweden (Greisman and Gaillard
2009; Olsson et al. 2010). During the last decades discussions have included the possibilities that plants could have
been important for consumption during the Mesolithic in
northern Europe (e.g. Larsson 2003). Seven species that
fulfil one or several of the indicator criteria for utilization
were found in the refuse layers at Tågerup.
Cornus sanguinea (dogwood)
Among the land plants found in samples from the former shore
zone at Tågerup, fruit stones of dogwood (Cornus sanguinea)
were found in spectacular numbers. In Table 3 only finds from
the processed soil samples are presented, but very large
quantities of dogwood stones were retrieved from wet sieving
during the excavations. In total, more than 300 stones were
found and, interestingly, most of these were cracked. There is
also one find of a probable digging stick or soil-working tool of
dogwood at the site. Dogwood has previously been documented at Mesolithic sites in southern Scandinavia (Regnell
1998). However, it is only at Bökeberg in Scania that large
amounts of fragmented stones of dogwood have been found
(Regnell et al. 1995). What makes the dogwood stones from
Tågerup remarkable is their fragmentation, an obvious result
from processing by people. One possible explanation for the
fragmentation might be that the stones are a by-product of oil
extraction. The dry weight of dogwood stones comprises as
much as 50% oil. A simple way of extracting the oil from the
fruit stones is to crush them, boil them in water and skim off
the oil from the water surface. The oil is excellent as lamp-oil
or as an impregnator for wood or leather, and was commonly
used in later times (Nyman 1868). Dogwood is probably one
of the most convenient terrestrial oil-resources of nondomesticated origin in Scandinavia. There are of course
alternative oil-sources such as seal blubber in coastal areas. As
with plant oil, it is difficult to prove the use of animal fat in
prehistory, although biochemical analyses may provide
important information (e.g. Isaksson 2000). Interestingly
there are far fewer seal bones found at Tågerup, when compared to other coastal sites dated to the Mesolithic period in
Scandinavia (Eriksson and Magnell 2001).
Malus sylvestris (crab apple)
A pip from apple (Malus sylvestris) was found in a sample
from the Kongemose phase at Tågerup. It cannot be ruled
out that the single, uncarbonised pip from this site could be
a result of natural deposition since crab apple trees can
grow adjacent to shores. The finds of crab apple from
Huseby Klev and Balltorp along the Swedish west coast
Veget Hist Archaeobot (2012) 21:1–16
represent carbonised remains and should be interpreted as
conscious use of the fruit (Larsson 2000). When the single
remain from Tågerup is added to the collection of Mesolithic findings of crab apples in Sweden, the few records
indicate that apples were not commonly used.
Prunus avium/cerasus (wild cherry/sour cherry
or dwarf cherry)
Two stones of cherry (Prunus avium/cerasus) were found in
the cultural layers. Both specimens are damaged and their
shapes are hard to use for more specific determination. The
distinction between the two species Prunus avium and P.
cerasus is not easy. P. avium has previously been defined as
being a natural consituent of the European decidious forest
whereas P. cerasus should be native to West Asia (Van
Zeist et al. 1994). Recent studies on distribution and
genetics suggest that the two species are very closely related
and that hybridisation between the species P. avium, P.
cerasus and P. fruticosa is responsible for present phenotypes (Badenes and Parfitt 1995; Brettin et al. 2000).
Although both P. avium and P. cerasus have evolved in
Europe, P. avium is native to southern Europe and P. cerasus to central and northwest Europe (Dirlewanger et al.
2009; Bortiri et al. 2001). The oldest find of cherry known in
Sweden (Prunus sp.) is from early Holocene peat from a
bog in Bohuslän (Hjelmqvist 1963). Because neither of the
two cherry species grows naturally on wet soils or on seashores both finds (from Bohuslän and Tågerup) may
therefore be considered as ‘exotics’ in the sediments. The
finds of cherry from Tågerup are the first documented from
a Scandinavian Stone Age archaeological site. On the
European continent the earliest record of cherry (P. avium)
seems to be described from a Mesolithic site in Southern
France (Vaquer and Ruas 2009). More frequent records start
with the Neolithic (Zohary and Hopf 2001).
Quercus sp. (oak acorns)
In the samples from the cultural layers belonging to the
Kongemose culture, 22 fragments of acorns (Quercus sp.)
were found. All fragments were carbonised or partly carbonised. During wet sieving at the excavation site, ca. 100
whole or cracked pieces of acorns were found, half of which
were carbonised. Acorns, as well as the bark of the oak, are
rich in tannins, which makes them unpleasant in taste and
unusable. The bitter tannins can be leached by soaking in
water or denaturised by heat (Källman 1993). The fact that
the acorns found at Tågerup were carbonised strongly indicates heat treatment as a means to make them edible. Acorns
and hazelnuts are easily identified during archaeological
excavations. They are often handled as any other artefact and
included in registration, documentation and publication.
This may be a reason why acorns are so rarely discussed in
the literature specifically reporting archaeobotanical finds. In
a compilation of plant remains from archaeological sites of
the Nordic countries (Regnell 1998) astonishingly few
acorns are included, i.e. from three Mesolithic and four
Neolithic sites. In other parts of Europe acorns are more
prominent in the archaeobotanical literature, in particular for
the Mesolithic period (Mason 1995; Vencl 1996).
Rubus idaeus (raspberry)
There are numerous finds of raspberry (Rubus idaeus) from
the cultural layers, and samples from the Kongemose refuse
layers contained 78 fruit-stones. It is not conclusive that finds
of raspberry indicate gathering since it normally grows on
fresh nitrogen-rich soil adjacent to seashores and wetlands
and may be deposited through natural processes. However,
samples with high numbers of raspberry fruit-stones (e.g. 55,
16) indicate an unusual high deposit of remains at the same
spot. The latter together with numerous acorns found in the
same sample suggest that raspberries might well have been
gathered. Fruit-stones of raspberries have been found earlier
in Stone Age sites in Scandinavia. Mesolithic finds were
made at Holmegaards Mose (Denmark) and Bökeberg
(Scania) but also at Rognelien in southern Norway (Regnell
1998). A large number of raspberry fruit-stones were found
in the stomach area of a Neolithic skeleton from a young
female often referred to as ‘the raspberry girl’, buried in peat.
It was assumed that the numerous remains were residues
from her last meal (Geijvall et al. 1952).
Sorbus aucuparia (rowan)
Two pips of rowan (Sorbus aucuparia) were found in the
Kongemose cultural layers. Rowan was found earlier in the
Mesolithic sites of Balltorp in Bohuslän (Larsson 2000)
and Bökeberg in Scania (Regnell et al. 1995). Rowan
grows on fairly moist soils and is favoured by unshaded
conditions at the edge of woodlands, and it may have
grown close to the seashore at Tågerup. In historical times
it has been used as a remedy for scurvy and kidney stones.
Fresh berries have also been used to make cider and vinegar or dried for jam (Nyman 1868; Høeg 1974) and are
recognised as an important source of vitamin C (e.g.
Oberdorfer 1990; Källman 1993).
Corylus avellana (hazelnut)
In the West Coast Line project shells of Corylus avellana
(hazelnut) are by far most abundant in samples from the
Mesolithic period (Table 4). In four out of five samples of
Mesolithic age where plant remains were found, hazelnut
shells were present.
Veget Hist Archaeobot (2012) 21:1–16
Table 4 Frequencies of Corylus avellana (hazelnut) shells within the
West Coast Line project, percentages of total preserved plant remains
in samples dated to different archaeological periods
Hazelnut shells (%)
Bronze age
Early iron age
Late iron age
Nutshells of hazel are the most common plant remains
found at the Mesolithic sites of Scandinavia. Approximately
20 Mesolithic and 25 Neolithic sites with finds of hazelnut
shells have been reported earlier (Regnell 1998). The
Mesolithic sites are characterised by higher frequencies of
hazelnut shells than the Neolithic sites, and roughly half of
the sites include refuse layers with both non-carbonised and
carbonised nutshells. In contrast, at the Neolithic sites the
nutshells are mainly carbonised. Therefore it may be proposed that hazelnuts not only played an important role for
foraging during the Mesolithic, but were also an important
subsistence base. This argument has recently been put forward in relation to finds from the Early Mesolithic of
northern Germany (Holst 2010). Modern hazelnuts contain
about 60% fat and 20% carbohydrates and 100 g nuts equal
660–720 kcal (Amaral et al. 2006). Thus, a few handfuls of
hazelnuts represent a substantial part of the daily requirement of energy uptake. Following the definitions of
Zvelebil (1994), the exploitation of hazelnuts during the
Mesolithic in temperate Europe was not opportunistic and
incidental but systematic and intensive. At Tågerup, nutshells were present in over 80% of the samples that contained preserved plant remains and they were, with few
exceptions, fragmented in a characteristic way, different
from the fragmentation caused by birds or rodents. Carbonised nutshells represented only a few percent of the total
amount. The occurrence of carbonised hazelnut shells in
sites of the Stone Age has been interpreted as due to their
being roasted for longer storage (Larsson 1983; Holst
2010), or as a result of their use as fuel (Kubiak-Martens
1999). Experiments showed that moderate roasting of
hazelnuts could prolong storage without significant loss in
nutritional value (Kirbaşlar and Erkmen 2003). Since
osteological data suggest year-around occupation at Tågerup (Eriksson and Magnell 2001), roasting of hazelnuts
might have been useful. However, the relatively low number of carbonised nutshells at Tågerup does not argue in
favour of roasting as a common procedure at the site.
When hazel is cut, new shoots appear that provide a richer
development of nuts. This phenomenon must have been
noted by Mesolithic people. Probably humans during that
time also knew that scattered hazelnuts sprouted and
produced new plants. It has been suggested that the rapid
expansion of hazel in Europe during early Post-glacial times
was at least partly an effect of intentional spreading by
humans (Iversen 1973; Bogucki 1988; Kuneš et al. 2008).
The rapid expansion has also been explained by natural
causes (Tallantire 2002). The benefits from induced growth
in hazel stands did not only come as increased harvests of
nuts but also with the production of long, slender branches
sprouting from coppiced stands. Hazel branches were indeed
used at Tågerup, for example in several of the fish traps.
On the basis of the present study and previous investigations it may be proposed that the hazelnut was a food
product the availability of which played an important role
for the introduction of farming in Scandinavia. The presence of numerous settlements of the Neolithic Linear Band
Pottery (LBP) culture around the mouth of the River Oder at
the southern Baltic coast, centuries before agriculture was
introduced in southern Scandinavia, has long been known
(Grygiel and Bogucki 1993; Lüning 2000). From these
settlements and other contemporaneous ones further east in
the area of Cuiavia in Poland (Czerniak 1998), there is
scattered but significant evidence of farming and animal
husbandry (Heussner 1989; Bogucki 2000). The Neolithic
settlements in the area south of the Baltic are dated to a
period between 5500 and 4700 B.C. (Persson 1999), clearly
predating the adoption of farming in southern Scandinavia,
which did not occur before ca. 3900 B.C. Artefacts from the
same period imported from—or locally produced but
influenced by—LBP culture were found in Sweden and
Denmark, such as the axe-types ‘Schuhleistenkeilen’ and
‘Breitkeilen’. Moreover, based on a thorough investigation
of Swedish aurochs remains, Ekström (1993) concluded
that this species became extinct in Sweden slightly after
7000 B.C. The aurochs disappeared from eastern Denmark at
the same time, although it survived in Jutland, western
Denmark, until the Early Subboreal (Aaris-Sørensen 1980).
Therefore, the presence of aurochs tooth beads in the graves
of the Scanian Mesolithic cemeteries Skateholm I and II
dated to ca. 5300–4700 B.C. (Jonsson 1988) suggests import
from continental Europe where aurochs populations still
existed. The practice of agriculture along the southern coast
of the Baltic, less than 150 km from the Scanian coast, and
occurrences of artefacts in Scania with provenance from the
LBP culture, indicate that the hunter-gatherers of southern
Scandinavia already had knowledge of farming practices at
that time, but neglected their necessity. The LBP culture
disappeared from the area close to the Baltic ca. 4700 B.C.
and retreated further south (Lüning 2000). When a later
expansion of Neolithic people arrived at the Baltic coast of
present day Germany and Poland at ca. 3900 B.C., farming
and animal husbandry became rapidly introduced to Denmark and vast areas of Sweden (Ammerman and Biagi
2003; Fischer and Kristiansen 2002).
Veget Hist Archaeobot (2012) 21:1–16
Fig. 5 Correlation of Corylus pollen curves from Denmark and southern Sweden. Positions of sites are shown in Fig 1. Curves represent pollen
percentages except for the Kurarp diagram which is expressed as concentrations i.e. number of pollen/cm3. See text for further explanations
There are not sufficient archaeobotanical data from
Mesolithic sites in southern Scandinavia to make a precise
quantification of the use of hazelnuts throughout this entire
period. The marked decrease in Corylus in the lowermost
part of zone T1:4a in the pollen diagram from Tågerup
could be interpreted as a diminishing hazel population due
to harvesting of nuts and especially to cutting sticks for toolmaking. A substantial proportion of the wooden artefacts
from the Ertebølle phase are made of hazel (Regnell and
Sjögren 2006). At the end of the settlement phase Corylus is
at first regaining its previous values but shows during the
following 600–700 years a slow decrease during which the
frequencies are reduced by half. However, a decrease in
Corylus pollen during this specific period has counterparts
in other investigations from southern Scandinavia. Here it is
suggested that a reduced availability of hazelnuts, as
interpreted from the decrease in Corylus pollen percentages
during the later part of the Mesolithic, may have motivated
adaption to farming (Fig. 5). The decrease in Corylus pollen
may be caused by climate change. The beginning of this
decrease is dated to about 4700 B.C. This coincides with
indications of increased precipitation and lowered summer
temperature in southern Sweden from ca. 4700 B.C., as
interpreted from charcoal data inferring decrease in forest
fire intensity (Greisman and Gaillard 2009; Olsson et al.
2010), as well as from stable isotope records (Hammarlund
et al. 2003). Wetter and colder climate may have affected
the hazel populations during this phase, since a positive
correlation between temperature during the flowering season and pollen accumulation rate for Corylus has been
found (Nielsen et al. 2010).
When Neolithic groups were present in the southern
Baltic area, 5500–4700 B.C., Corylus show frequencies
between 20 and 40% and high concentrations (Fig. 5).
Pollen values decrease from 5500 B.C. reaching a minimum
that occurs well after the disappearance of Neolithic groups
from the region. The prominent decrease of Corylus pollen
between c. 5500 and 4500 B.C. might be related to a
reduction in nut production. When the next expansion of
farming approached the coast of southern Baltic, in ca. 3900
B.C., Neolithic adaptation to farming took place rapidly and
simultaneously in southern Scandinavia. At the first contact
with Neolithic cultures, people in southern Sweden had an
abundance of hazelnuts and perhaps relied on them as a
staple food resource and, therefore, farming appeared less
attractive. It is important to state that the decrease in hazel
nut availability may have been one of several factors behind
the adaptation to/introduction of farming in Sweden.
Despite considerable settlement activities at Tågerup, the
pollen record does not indicate any substantial human
impact on the vegetation during the Mesolithic. The presence of charcoal particles and pollen from grass and herbs
associated with nutrient-rich soils are contemporaneous
with the Kongemose settlement. The Ertebølle settlement
phase, although characterised by considerable dwelling
activities less than a hundred metres from the pollen
sampling site, is scarcely seen in the pollen data. Apart
from the changes in tree species composition during the
Early Neolithic there is no indication of agriculture or other
human impact on the vegetation during this period. In
contrast the pollen composition during the latest part of the
Early Neolithic and the earliest part of the Middle
Neolithic (3600–3200 B.C.) suggests a more open landscape
than earlier and an increase in grazed areas (animal husbandry), however, still with no indication of agriculture.
Numerous finds of crushed dogwood stones suggest that
they were used for the extraction of oil. Several other forest
plants were found and some may have been specifically
used for consumption, i.e. apples, cherries, raspberries,
acorns and rowan-berries.
The importance of hazelnuts as a food source during the
Mesolithic may have been underestimated in the literature.
At the time of establishment of farming communities from
continental Europe along the coasts of the southern Baltic
between 5500 and 4700 B.C., Corylus exhibits pollen values
between 20 and 40% in pollen diagrams from southern
Scania and Denmark. The occurrence of imports from
continental Europe (e.g. specific axe-types and aurochs
beads) at Mesolithic sites in Scania implies that people in
Scania might have been conscious of the farming practices
in present northern Germany but did not yet adapt to a
Neolithic way of life. During a later phase of the Mesolithic
period, a regional decrease in Corylus as suggested by lower
pollen percentages may indicate a decrease in the availability of hazelnuts. The latter may, in turn have led to a
shortage in protein and energy resources in people’s consumption, which may have forced them to alter their subsistence habits. When the next expansion of farming
economy reached the Baltic coast of present day Germany,
at about 3900 B.C., it led to an almost simultaneous adaption
to farming and animal husbandry in Southern Scandinavia.
Despite the pollen records suggesting that the hazel populations had regenerated considerably in the Early Neolithic,
people had reasons to be less reliant on hazelnuts at around
3900 B.C. However, this hypothesis should be tested further
through studies of the occurrence and types (carbonised,
non-carbonised, fragmentation etc.) of hazelnut shells at
settlements dated to the Middle and Late Mesolithic.
Acknowledgements I am grateful to Leif Björkman and Joachim
Regnéll who performed the pollen analysis and made an initial
description of the pollen diagram, to Sven Karlsson who assisted with
the diagram construction, to Ingrid Bergenstråhle who provided
remarks on archaeological aspects and to Małgorzata Latałowa, Jan
Risberg, Marie-José Gaillard and two anonymous reviewers who
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