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ORIGINAL ARTICLE
New insights into vegetation dynamics and settlement historyin Hummling, north-western Germany, with particular referenceto the Neolithic
Annette Kramer • Felix Bittmann • Daniel Nosler
Received: 2 October 2012 / Accepted: 6 June 2013
� Springer-Verlag Berlin Heidelberg 2013
Abstract Palynological investigations on two well-dated
peat profiles provide insights into Neolithic vegetation and
settlement history from Hummling in north-western Ger-
many. The site selections allow comparisons between local
and regional vegetation changes and are used to estimate
the extent of Neolithic influence on the vegetation. The
interpretation of the fossil spectra relied on radiocarbon
dating, evaluation of pollen indicator taxa, non-pollen
palynomorphs and multivariate techniques. During the late
Mesolithic the vegetation was dominated by mixed oak
forests while openings in forest cover were detected, with a
decline in elm reflected in the regional pollen record
around 4250 cal. B.C. The presence of humans is shown by
settlement indicators that are first recorded at ca. 3800 cal.
B.C. Vegetation changes were small between 4300 and
3600 cal. B.C. This suggests that regional vegetation was
relatively resilient to small-scale disturbances. Possible
indications of grazing were recorded in the spectra of the
local pollen profile but there is no clear-cut evidence for
Neolithic activity. Between 3520 and 2260 cal.
B.C. decreases in forest cover were inferred from both
profiles and increases in settlement indicators reflect
farming activity. These changes coincide with the emer-
gence in the area of the Funnel Beaker Culture and the
subsequent Single Grave Culture. Both profiles suggest that
settlement probably ceased between ca. 3230 and 3050 cal.
B.C. This lull or cessation in activity was probably regional
in character. After 2260 cal. B.C. human impact on the
vegetation decreases and woodlands regenerate. The
longevity of the regeneration phase—ca. 690 years—was
probably connected with the low resilient capability of the
vegetation on the poor soils.
Keywords Neolithic � Pollen analysis � Human impact �Northern Germany � Funnel Beaker Culture �Single Grave Culture
Introduction
Today, the Hummling, an old morainic ridge in the Emsland,
is characterised by cultivated land and parcels of semi-nat-
ural mixed oak–birch forests. This is largely the result of
prolonged farming activity. Farming was introduced by
Neolithic settlers in northern Germany ca. 4100 cal.
B.C. (Kalis and Meurers-Balke 1998; Hartz et al. 2000;
Muller et al. 2010) while intensification of farming took
place between ca. 3600 and 3400 cal. B.C. during the phase of
megalithic tomb construction (Hartz et al. 2007; Kirleis et al.
2012). The megalithic tombs of the Hummling are assignable
to the west group of the Funnel Beaker Culture (Trich-
terbecherkultur, TRB), the earliest Neolithic culture known
from north-western Germany. These peoples occupied the
area during the middle Neolithic (EN II–MN V according to
the Northern European Plain chronology) between 3400
and 2800 cal. B.C. and were subsequently displaced by
representatives of the Single Grave Culture (SGC;
Communicated by M. O’Connell.
A. Kramer (&) � F. Bittmann
Lower Saxony Institute for Historical Coastal Research,
Viktoriastr. 26/28, 26382 Wilhelmshaven, Germany
e-mail: kramer@nihk.de
F. Bittmann
e-mail: bittmann@nihk.de
D. Nosler
Landkreis Stade Archaologische Denkmalpflege, Schloss
Agathenburg, Hauptstraße 45, 21684 Agathenburg, Germany
e-mail: daniel.noesler@landkreis-stade.de
123
Veget Hist Archaeobot
DOI 10.1007/s00334-013-0409-z
ca. 2800–2200 cal. B.C.) (Brindley 1986; Muller et al. 2010).
Little is known about the Neolithic peoples who settled in
Hummling and the way they used and influenced their
environment. Limited information about settlement pattern,
houses and related features is provided by investigations on a
few domestic sites (Nosler et al. 2011). Macrobotanical data
from domestic sites that might be expected to contribute to
our knowledge of subsistence strategies are hence scarce.
Most of the available information from the wider region
relates to the Elbe-Weser area where charred cereal remains,
mainly Hordeum vulgare, Triticum dicoccum and T. mono-
coccum, have been recorded in the TRB settlement Flogeln
(ca. 3050 cal. B.C., details in Behre and Kucan 1994).
The present perception of the Neolithic in north-western
Germany differs from that in neighbouring regions such as
the Netherlands and Schleswig–Holstein in that these
regions have evidence for Neolithic cultures that predate the
middle Neolithic (Raemaekers 1999; Out 2009, 2012; Out
and Verhoeven 2013; Hartz et al. 2007; Muller et al. 2010).
This is not easily explained given that present-day political
boundaries can have had no bearing on these prehistoric
cultural developments (Raemaekers 2013). Preservation
may be important insofar as the sandy soils of north-western
Germany do not favour preservation or indeed detection of
Neolithic sites (Nosler et al. 2011). The study of vegetation
change as detected by pollen analysis, on the other hand, can
be a powerful tool in that pollen data give clear insights into
human impact and especially farming activity (Behre and
Kucan 1994; Kalis and Meurers-Balke 1998; Bakker 2003).
Palynological investigations to date in the Emsland, how-
ever, have rather poor sample resolution and chronological
control is often inadequate or indeed unsatisfactory (Koch
1934; Jonas 1935, 1941, 1943; Kramm 1981) and so the
available data are insufficient to enable a critical and reliable
reconstruction of early prehistoric farming impact. Investi-
gations at the Dummer Lake, ca. 60 km to the south-east,
show early human impact in connection with archaeological
finds at the settlement site Hude I and a wooden trackway
dated to ca. 4800 cal. B.C. (Schutrumpf 1988; Bauerochse
2003; see Kampffmeyer 1991 for further consideration of
the function and chronology of the site). Detailed palyno-
logical investigations in the Elbe-Weser area (Dorfler 1989;
Behre and Kucan 1994; Heider 1995) have provided evi-
dence for Neolithic settlement periods that parallel trends
observed in other parts of northern Europe (Troels-Smith
1954; Iversen 1941; Rasmussen 2005; Wieckowska et al.
2012) though here also some aspects of the chronology
might be further refined. These investigations show the
crucial importance of short distances between coring loca-
tion and Neolithic settlement sites for the reflection of
human activity in pollen spectra (Behre and Kucan 1986).
In this study, two well-dated peat pollen profiles from
Hummling are presented with a focus on reconstructing
Neolithic impact. Holschkenfehn, that is expected to reflect
mainly local vegetation development and farming activity,
was selected because of its close proximity to several
megalithic tombs and two possible settlement sites
(Fig. 1c). The pollen profile from Bockholter Dose, on the
other hand, is expected to provide information on vegeta-
tion development and Neolithic impact at a regional level.
Study sites
Hummling, the study area, is in Emsland, Lower Saxony,
between the river Ems and its tributaries Hase and Ohe
(Fig. 1). The landscape is largely defined by an east–west
orientated morainic (Geest) ridge formed during the Saale,
i.e. the penultimate glaciation (Hauschild and Luttig 1993).
The landscape is also influenced by later geological
Fig. 1 Maps relating to the study area and the wider region:
a overview of north-western Germany showing location of the study
area and Flogeln (indicated by rectangles with non-dashed and
dashed boundaries, respectively). NL, Netherlands; LS, Lower
Saxony; SH, Schleswig–Holstein: b map of Hummling showing
main features including relevant archaeological sites. Hatching
indicates present-day bog distribution. 1 Bockholter Dose, 2
Holschkenfehn; c Detailed map of sampling site Holschkenfehn.
Grey shading indicates afforestation. Mapping of megalithic tombs
and settlement sites based on current knowledge based on ongoing
research
Veget Hist Archaeobot
123
developments. Melting of ice sheets after the last glacial
maximum led to a sea-level rise that resulted in a higher
groundwater-table. This, in turn, favoured the development
of fens and reed swamps in low-lying areas (Behre 2007).
At the same time, the progressively advancing North Sea
shoreline resulted in a more oceanic climate that favoured
the formation of ombrotrophic bogs (raised bogs) on the
sandy soils after ca. 6000 cal. B.C. (Petzelberger et al.
1999). Expansion of bog resulted in part displacement of
the dominant oak-rich forests that included Betula, Tilia,
Ulmus and Acer. The climate is humid (rainfall 815 mm/
year), summers are relatively cool and winters are mild
(mean annual temperature 9.0 �C; meteorological data
relate to the meteorological station at Loningen, Deutscher
Wetterdienst 1996).
The present-day woodlands consist mainly of birch and
oak (Betulo-Quercetum roboris). In areas with podsols and
acid gley soils, Pinus sylvestris is more common on drier
stands but also grows on peatlands, while Alnus glutinosa
is common on water-saturated mineral soils. On fertile (but
podsolised) luvisols, mixed-oak forests are dominated by
Fagus sylvatica (Fago-Quercetum petraeae; Pott 2002).
F. sylvatica occupies the niche filled by Tilia cordata in the
former Atlantic forests and hinders the spread of weak
competitors such as Carpinus betulus which is generally
pushed to the limit of its ecological range on water-satu-
rated soils that are unsuitable for beech (Pott 2002).
Today, areas no longer suitable for farming due to soil
degradation and leaching carry heathlands with Juniperus
communis or heather-rich communities with Calluna
vulgaris and Erica tetralix (Menke 1963). This process of
degradation probably started during the Neolithic when
there was substantial activity. More than 80 megalithic
tombs are known from the area which, in itself, points to a
substantial Neolithic TRB population (Fig. 1b). Heede is
the only Neolithic settlement site investigated in Emsland
that can with certainty be ascribed to the Neolithic (Froh-
lich 1991). It is situated west to Hummling and the river
Ems and has been dated by archaeological finds to TRB
and SG cultures. Based on surface finds, five areas with
potential settlement sites have recently been identified in
Hummling (Fig. 1b). The small number of possible
domestic sites is possibly due to destruction by deep
ploughing (Kramer et al. 2012).
Holschkenfehn is a nature reserve characterised by
birch-pine woodlands that include oak and heaths that are
dominated by Calluna vulgaris (Fig. 1). Within a wood-
land area, an ephemeral pond (52�46055.500N,
7�30049.1400E; Fig. 1c; dimensions: *300 m 9 80 m) was
sampled. The pond today lacks open water but local people
are aware of ice skating taking place there as late as the
1990s. Within 100–250 m of the sampling site, seven
megalithic tombs and two chambered graves, attributable
to TRB, West Group (Schlicht 1972), have been recorded.
In the vicinity of the sampling site, two potential TRB
settlement sites had been identified based on find scatters
(Fig. 1c). Features to the south of the site were originally
regarded as evidence for a TRB house (Kaltofen 1991) but
this is now regarded as unlikely (Nosler et al. 2011).
Peat from the nature reserve Bockholter Dose was used
as source for a regional pollen diagram (52� 510 5900 N, 7�440 5500 E). Today, this bog occupies about 130 ha but was
much more extensive prior to drainage in the context of
intensive agriculture (LBEG 2012). The area today sup-
ports heath with C. vulgaris and M. caerulea and is the
subject of restoration by raising the prevailing water-table.
Materials and methods
Sampling and dating
At the Holschkenfehn a 1.5 9 2 m2 pit was trenched after
several test drillings to locate an appropriate sampling
point. A 1.40 m-long peat profile was recovered in over-
lapping segments at the deepest point within the former
pond from a clean profile wall, using three zinc boxes
(50 9 10 9 5 cm3). Beneath the top soil, highly decom-
posed peat was recovered with some sand in the deeper
parts. The base of the profile (below 108 cm) consisted of
humic sand.
At the Bockholter Dose, a 3.50 m-long peat core was
collected using a Russian corer (chamber dimensions:
50 9 4 cm). The core was recovered close to the northern
border of the nature reserve. In the present study the
interval 66–220 cm, consisting of low to medium-decom-
posed peat, was analysed.
Seven AMS 14C dates were obtained for the profile from
Bockholter Dose and eleven from Holschkenfehn. The
samples from Holschkenfehn were soaked in KOH (5 %).
Macro-remains suitable for dating were picked from the
material, retained by a 200 lm-mesh sieve. Samples from
Bockholter Dose yielded insufficient macro-remains for
dating. Bulk peat samples were therefore used, rootlets
having been removed with the help of a binocular
microscope.
Age-depth models were constructed using OxCal 4.1
(Ramsey 1995) based on IntCal09 (Reimer et al. 2009).
The programme uses Bayesian statistics that incorporate
the prior model (depth and deposition order) and the 14C
dating information for the construction of the age-depth
model. The applied P_Sequence approaches the unknown
deposition rate by introducing the parameter k which rep-
resents the number of accumulation processes per depth
unit. The parameter was set to 1 deposition unit per cm to
avoid circular reasoning and in cognizance of the
Veget Hist Archaeobot
123
randomness in deposition processes (Ramsey 2008). In the
Holschkenfehn profile, the increase of Secale pollen to
more than 1 % was used as an additional date for the
construction of the age-depth model. This implies that the
upper two samples relate to the Roman Iron Age or are
somewhat younger (cf. Behre and Kucan 1994).
Palynology, macro-remains and charcoal
The Holschkenfehn core was sampled continuously using
1 cm-thick samples in the interval relating to the Neolithic
(each sample is estimated to represent *40 years). Else-
where, the sampling interval was 2–5 cm. The uppermost
15 cm was highly decomposed and was therefore not
analysed. In total, 66 samples were analysed. The Bock-
holter Dose core was sampled at 2–5 cm intervals giving a
total of 39 samples and a time resolution of 40–100 years.
The pollen samples (1 cm3) were processed using the
standard protocol that includes treatment with KOH
(10 %), HCl (10 %), sieving using a 200 lm mesh, HF
(40 %) and acetolysis (2.5 min) (Fægri and Iversen 1989).
Lycopodium tablets (Batch No. 177745, Lund University)
were added at the start of the preparation procedure to
enable subsequent calculation of sporomorph and charcoal
(size range: 10–200 lm) concentrations (Stockmarr 1971).
The samples were mounted in glycerine and analysed using
a light microscope at 9400 magnification. A magnification
of 91,000 and phase contrast was used where it was
desirable to see the detailed structure and surface pattern of
specific pollen grains. Identification and nomenclature of
pollen types mainly follow Beug 2004, and in a few
instances (Moore et al. 1991). Non-pollen palynomorphs
(NPPs) are after van Geel (1978), algae and fungi follow
van Geel et al. (1980/1981) and for testate amoebae the
publication by Charman et al. (2000) was used.
At least 800 arboreal pollen grains (AP; Corylus
excluded) per sample were counted in the Holschkenfehn
profile. In samples with abundant Alnus pollen, 1,000 AP
grains were counted. Large amounts of plant debris made
counting of the Bockholter Dose samples very difficult so
that, in general, three slides (24 9 32 mm2) were required
to achieve an AP count of 600.
Pollen and NPP percentages were calculated using a
total terrestrial pollen sum (TTP). Calluna was excluded
from the pollen sum as it was regarded as a local element
of the bog vegetation. This is supported by frequent
macro-remains of Calluna recorded from both profiles. The
data are presented as percentages which are regarded as
preferable to concentration or pollen accumulation rate
(influx) data (cf. Waller et al. 2012). Other considerations
influencing this choice include the possible effects of cli-
mate on pollen productivity (van der Knaap et al. 2010)
and uncertainties relating to dating uncertainties.
The pollen diagrams were drawn using Tilia ver. 2.0.2.
(Grimm 2004). Pollen zone boundaries were placed after
careful visual inspection of the profiles.
Macro-botanical remains ([200 lm) obtained by siev-
ing during pollen preparation were examined using a ste-
reo-microscope. Charcoal concentrations are given as
particles/cm3.
Ordinations
Pollen percentage data from each profile were analysed
using principal components analysis (PCA), as the data sets
showed a linear response to theoretical gradients during
detrended correspondence analysis (DCA) (gradient
length \2; DCA results not shown).
The analyses were run to objectively investigate pollen
taxa responses to theoretical gradients (axes of the ordi-
nation plot) and to identify taxa that react similarly within
the data set. Other points of interest included checking for
sample clustering and validation of the pollen zones.
For the analyses, all terrestrial pollen types that were
present in at least three samples throughout the profile or
showed values [0.5 % in one sample were included. Per-
centage data were square-root transformed to improve
comparability. For these analyses, CANOCO ver. 4.5 and
CanoDraw ver. 4.5 were used (ter Braak and Smilauer 2002).
A detrended canonical correspondence analysis (DCCA)
was applied to the AP data from the Bockholter Dose to get
a semi-quantitative measure of regional forest changes
during the Neolithic. Herb pollen was excluded from the
analysis as it is under-represented in forested areas (Hicks
1971) and may derive mainly from local vegetation. By
introducing the age-depth relationship as a constrained
gradient in the analysis, the DCCA enables the detection of
total compositional species turnover through time (Birks
2007). Detrending was done by segments and percentages
were square-root transformed and scaled non-linearly. The
species scores are given as direct standard deviation units
(SD) of the compositional turnover (Birks 2007; Hill and
Gauche Hill and Gauch 1980).
Results
Age-depth modelling
The results of the radiocarbon dating are presented in
Table 1 for both profiles, and age-depth models are shown
in Figs. 2 and 3 (Bockholter Dose and Holschkenfehn,
respectively). The radiocarbon dates for Holschkenfehn
were partly published in Nosler et al. (2011) and Kramer
et al. (2012); the radiocarbon date for depth 45 cm
was, however, incorrect and has since been re-dated.
Veget Hist Archaeobot
123
For construction of the age-depth model for Holschkenfehn,
the results from the lowermost sample, 114–115 cm (KIA-
42180), were excluded from the analysis. The leached res-
idue contained insufficient carbon for reliable dating
(0.1 mg). The humic fraction, on the other hand, returned an
unexpectedly young date. This is probably attributable to
humic matter deriving from the overlying peat. The cali-
brated and modelled ages are in good agreement. As regards
Bockholter Dose, all dates were used to construct the age-
depth model and the modelled ages are in good agreement
with the calibrated dates. Ages in the following text are
given as the arithmetic mean of the 95.4 significance
interval (2 r range).
Pollen diagrams
Pollen diagrams from Bockholter Dose and Holschkenfehn
are presented in Figs. 4 and 5, respectively. In Fig. 6,
curves for the wetland taxa and NPP in the Holschkenfehn
profile are presented. The PAZs and the macrofossil data
relating to Bockholter Dose and Holschkenfehn are sum-
marised in Tables 2, 3, respectively.
Ordination
The ordination diagrams (Figs. 7, 8) show the first two axes
of the PCA. Only taxa that fit 25 % or more to the axes are
displayed in the ordination plot. Samples are shown using
symbols that indicate the PAZs as distinguished by visual
inspection of the pollen diagram. The ordination of the
terrestrial pollen data from Holschkenfehn is shown in
Fig. 7. The first and second axes account for 51 % of the
total variance. Both axes seem to reflect vegetation dis-
turbance. The first axis seems to reflect the transition from
woodland to human-altered vegetation while the second
axis appears to reflect a primary/secondary woodland gra-
dient. The clustering of samples serves to confirm
the validity of the zoning, and the transitional character
of the vegetation and a trend towards more disturbances in
PAZ 2.
Results of the PCA analysis of Bockholter Dose pollen
spectra are shown in Fig. 8. The first axis (eigenvalue
0.35), i.e. the main gradient, separates pollen taxa that
reflect woodland from taxa indicative of open vegetation.
This axis is therefore used to assess openness of the veg-
etation (Fig. 9). Quercus separates from Corylus and
Table 1 Dating results
Lab code (KIA-) Depth (cm) Material dated 14C dates
(B.P.)
d13C
(%)
Age, 2r-range
(cal. B.C.)
Modelled
ages (cal. B.C.)
Holschkenfehn
Pollen event 21–20 Secale pollen [1 % A.D. 50–140
42993 36–35 Peat* 3,520 ± 30 -24.27 1920–1750 1920–1750
47126 39–40 Peat* 3,760 ± 30 -23.32 2290–2040 2270–2040
42994 46–45 Peat* 3,960 ± 30 -27.25 2570–2350 2570–2350
47127 49–50 Peat* 4,110 ± 25 -25.88 2860–2580 2750–2580
42995 56–55 Peat* 4,300 ± 30 -26.26 3010–2880 3010–2880
42996 66–65 Peat* 4,430 ± 35 -26.76 3330–2920 3340–3160
47129 70–71 Peat* 4,720 ± 25 -26.62 3630–3380 3630–3380
42997 76–75 Peat* 5,060 ± 35 -24.35 3960–3770 3880–3710
42998 86–85 Peat* 5,210 ± 30 -24.52 4150–3960 4230–3970
42179 96–95 Charred Erica tetralix branch 5,650 ± 45 -27.00 4580–4360 4590–4370
42180 114–115 Sedges (humic acid fraction) 5,630 ± 40 -28.17 4540–4370 Not included
42180 114–115 Sedges (leached residue) 5,800 ± 250 -27.41 5210–4270 Not included
Bockholter Dose
42984 80–79 Peat* 3,760 ± 30 -25.09 2290–2040 2290–2050
42982 110–109 Peat* 4,160 ± 30 -26.97 2880–2630 2870–2630
42983 138–137 Peat* 4,490 ± 35 -26.93 3350–3030 3330–3100
42985 155–154 Peat* 4,710 ± 30 -24.17 3630–3370 3630–3380
42173 180–179 Ericaceae remains, Sphagnum leaves 5,210 ± 35 -29.64 4220–3960 4050–3950
42174 220–219 Ericaceae remains, Sphagnum leaves 5,390 ± 90 -31.50 4440–3990 4690–4400
42176 251–250 Ericaceae remains, Sphagnum leaves 6,270 ± 55 -32.09 5370–5060 5320–5020
* Peat was sieved [200 lm and root material removed
Veget Hist Archaeobot
123
Betula on the second axis, which suggests that the latter
two taxa are indicative of transitional vegetation stages.
The ordination shows poor clustering of spectra which
suggests that the pollen zones are not distinctive and that
changes are gradual.
The DCCA carried out on the AP data from the Bock-
holter Dose profile (Fig. 9) shows a compositional species
turnover of 0.6 (eigenvalue 0.03), measured as standard
deviation units. Total variance of the data sets is given as
0.29 total inertia.
Discussion
Regional vegetation change inferred
from the Bockholter Dose
The pollen spectra from the Bockholter Dose (Fig. 4) are
considered to reflect regional vegetation changes as it
is a large bog complex (Jacobson and Bradshaw 1981).
The pollen record indicates that between 220 and 66 cm
(4500 and 1900 cal. B.C.), the vegetation in the Hummling
area consisted of woodland dominated by Quercus, Bet-
ula, Fraxinus, Tilia and Ulmus. High Alnus values prob-
ably reflect the widespread presence of alder carr that was
favoured by much wet habitat (Bauerochse 2003). Pinus,
which has relatively high percentage pollen values, was
probably largely confined to drier areas. Today, Betulo-
Quercetum roboris communities are regarded as the cli-
max stage of forest development on the sandy soils in the
region (Pott 2002). It seems therefore that the woodland
communities have changed little during the last
6,000 years. Only Tilia, a substantial component of the
Atlantic forests of north-western Germany (Behre and
Kucan 1994), has declined in importance. In the pollen
record, it is greatly underrepresented due to low pollen
productivity and poor pollen dispersal (Andersen 1967).
Tilia was probably distributed on the better soils that are
today occupied by beech or used for agriculture (Behre
and Kucan 1994). Generally, low NAP values underline
Fig. 2 Age-depth model for Bockholter Dose, profile DMB Fig. 3 Age-depth model for Holschkenfehn, profile HSV
Veget Hist Archaeobot
123
Fig. 4 Pollen diagram of main taxa from the Bockholter Dose
(profile DMB). Percentages based on total terrestrial pollen counts
(TTP). Where values are low, silhouettes show the values magni-
fied 910. Analysis: Kramer 2012
Fig. 5 Pollen diagram showing curves for AP, shrub taxa and NAP
from Holschkenfehn (profile HSV). Percentages are based on TTP.
Where values are low, silhouettes show the values magnified 910.
Analysis: Kramer 2012
c
Veget Hist Archaeobot
123
the regional character of the pollen rain at the site but,
mostly wind-pollinated species like Artemisia, Rumex and
members of the Chenopodiaceae and Poaceae are con-
sistently represented. Changes in the woodland
composition are visible throughout the record and the main
trends are indicated by the ordination of samples along
PCA axis 1 (Figs. 8, 9). The vegetation changed in PAZ 1
with Betula decreasing above 210 cm (4400 cal. B.C.),
Table 2 PAZ descriptions and macrobotanical records, Bockholter Dose profile
PAZ Depth
(cm)
Age (B.C.,
model)
Main zone characteristics Macro-botanical remains
6 97–66 2490–1920 Ulmus, Tilia, Fraxinus generally decreasing, Fagus increases,
Carpinus recorded, P. lanceolata increasing
Sphagnum leaves C. vulgaris twigs,
sedge rootlets
5 123–97 2960–2490 Betula and Quercus increase, Ulmus, Tilia, Fraxinus show peaks.
P. lanceolata and R. acetosa only sporadic
Sphagnum leaves, C. vulgaris twigs,
sedge rootlets
4 155–123 3520–2960 Low Ulmus, Tilia, and Fraxinus, high Corylus values, Betula and
Quercus decreasing. P. lanceolata recorded. Poaceae high values,
maximum values for Calluna. High charcoal values
Sphagnum leaves C. vulgaris twigs,
sedge rootlets
3 173–155 3860–3520 Relatively high Ulmus, Tilia and Fraxinus values, Corylus decreases.
P. lanceolata recorded, slightly higher R. acetosa values, cerealia-
type pollen, high charcoal values
Sphagnum leaves, sedge rootlets
E. vaginatum remains, Cenococcum
geophilum
2 200–173 4250–3860 Quercus and Corylus increase at the base of the zone, followed by
Fraxinus, Tilia and Ulmus. First Fagus records
Sphagnum leaves, charred C. vulgaris
twigs, sedge rootlets E. vaginatum
remains
1 220–200 4510–4250 Alnus, Betula, Corylus and Quercus are the dominant taxa. Ulmus
declines at the end of the zone; NAP rather poorly represented
Sphagnum leaves, charred C. vulgaris
twigs, sedge rootlets
Fig. 6 Diagram from Holschkenfehn (profile HSV), showing wetland taxa and non-pollen palynomorphs. Where values are low, silhouettes
show the values magnified 910. Analysis: Kramer 2012
Veget Hist Archaeobot
123
while increasing Corylus values point to openings in the
forest canopy (Kalis et al. 2003). Further opening-up is
suggested by Ulmus which declines at the PAZ 1/2 tran-
sition after 205 cm (4250 cal. B.C.) while Fagus pollen is
recorded for the first time. It has been often discussed
whether opening-up of woodland as a result of human
activity facilitated local establishment of Fagus (Behre and
Kucan 1994; Kuster 1997; Pott 1997; Nielsen et al. 2012).
Our investigations support the idea that beech benefited
from human-induced woodland disturbances in our study
area. The woodland openings seem to have been easily
reversible so that first Betula and then Ulmus soon
recovered again. However, a small change in the forest
composition is visible at the beginning of PAZ 4 (after
155 cm, 3500 cal. B.C.) with declining Tilia and Ulmus in
the mixed-oak woodlands (Figs 4, 9). The high Corylus
values point to woodland clearings that were probably
moderate considering the decline of total tree pollen from
about 75 to 65 %. Settlement indicators like Plantago
lanceolata, Pteridium aquilinum and Poaceae (non-culti-
vated and cerealia-type) in PAZ 3 above 170 cm (after
3800 cal. B.C.) suggest that the changing woodland com-
position is probably related to human activities in the
region. Also increased macroscopic charcoal in samples
from PAZs 3 and 4 (between 160 and 130 cm;
3620–3090 cal. B.C.) points to human activity in the area.
In PAZ 5, 123–97 cm (2960–2490 cal. B.C.) woodland
regenerated and Betula increased when human activities
ceased, indicated by a decrease in Plantago and Pteridium
values. Also, late succession trees like Tilia and Ulmus
recovered for a short period between 2800 and 2600 cal.
B.C. (110 and 100 cm). In PAZ 6 (97–66 cm, 2490–
1920 cal. B.C.), Fagus is at about 1 % which suggests local
presence of beech.
Local environmental changes at Holschkenfehn
Given the small size of the bog, the pollen spectra from
Holschkenfehn (Fig. 5) are assumed to reflect local and
extra-local vegetation changes (Jacobson and Bradshaw
1981; Moore et al. 1991). In PAZ 1 (5190–4180 cal B.C.),
pollen from mixed-oak woodlands dominates and any
changes are small. NAP taxa such as Succisa, Cichorioideae
and Melampyrum are regularly recorded in PAZ 1 which
points to locally present woodland clearings in the late
Atlantic. At the transition PAZ 1/2, Ulmus declined while
Tilia and Quercus increased. Whether higher values for
Quercus and Tilia reflect greater contribution by oak and
lime to the surrounding woodlands, enhanced flowering
within an opened woodland or higher pollen influx from the
regional vegetation is difficult to assess. The overall changes
in PAZs 1 and 2, however, appear to be relatively small. The
results of the PCA ordination also support this view (Fig. 7)
and so it is concluded that there were only minor vegetation
changes during this time.
In PAZ 3 (71–42 cm, 3520–2260 cal. B.C.), increasing
values of human settlement indicators such as Rumex
acetosella-type, Jasione montana-type, Scleranthus annuus
and P. lanceolata as well as a decrease in AP (mainly Alnus,
Betula and Tilia) indicates opening-up of woodland cover
and suggests local presence of farmers. The NAP pollen
types referred to above may be regarded as indicative of
arable land as they are frequently found as segetal flora on
fallow land (Behre 1981). Pollen from cerealia-type is also
regularly present between 70 and 43 cm (3490 and 2310 cal.
B.C.). This supports the idea of crop cultivation, even though
the possibility that some of this pollen may arise from non-
cultivated grasses cannot be excluded (see below). Stron-
ger human impact is particularly evident in the peri-
ods between 3490 and 3230 cal. B.C. (70–65 cm),
3050–2870 cal. B.C. (60–54 cm) and from 2870 to 2310 cal.
B.C. (54–43 cm). The local hydrological changes, as reflec-
ted by the NPP records (Fig. 6), show a shift to more humid
conditions in zone 3 (indicators of dry conditions, including
Assulina muscorum, Callidina angusticollis (Van Geel
1978) and Byssothecium circinalis (Van Geel et al. 1980/81)
decrease). Pollen taxa, such as Hydrocotyle vulgaris,
Utricularia, Scheuchzeria palustris and Sparganium-type,
that are indicative of shallow dystrophic water bodies, point
to increased wetness and probably the formation of an open
water-body during this time. Human-induced woodland
disturbance, which resulted in reduced evapo-transpiration
and increased surface-water run-off, was probably mainly
responsible for this development.
Pteridium aquilinum values increase strongly in PAZ
3b, i.e. at ca. 2870 cal. B.C. This suggests vegetation dis-
turbance. Pteridium is common in mixed woodlands today
but the ability to produce spores is largely restricted to
situations where there are openings in the tree canopy.
Furthermore, rhizomes that are resistant to disturbance and
with high growth potential enable this fern to quickly
expand, given favourable circumstances, by vegetative
reproduction. Tree taxa such as Betula, Ulmus, Fraxinus,
Pinus and Alnus decline sharply between 2610 and
2300 cal. B.C. (49 and 43 cm), which further suggests
severe woodland disturbances. The stable values for
Quercus are interpreted as reflecting a steady input of
Quercus pollen from the wider region, as local vegetation
was cleared. Synchronous with the decline in AP, Corylus
increases in PAZ 3c at 2520–2260 cal. B.C. (47–42 cm)
which supports the idea of further woodland clearance. A
little later at 2420 cal. B.C. (45 cm), this is followed by
maximum Poaceae values. This suggests that grasslands
have replaced closed-canopy woodland, probably as a
result of the impact of grazing animals (Groenman-van
Waateringe 1993; Vera 2000). However, elevated Poaceae
Veget Hist Archaeobot
123
values as a response to locally drier conditions cannot be
excluded from consideration (see below). Woodland
recovered after these disturbances and Betula increased and
achieved maximum values in PAZ 4a between 2260 and
1570 cal. B.C. (42 and 34 cm). As a high pollen producer,
input of Betula pollen probably contributes to the reduction
in Quercus values between 1990 and 1570 cal. B.C. (37 and
34 cm). When Betula decreased after 1570 cal.
B.C. (34 cm), Quercus values increased again implying
presence of well developed mixed-oak woodland.
Since neither human indicators nor high charcoal con-
tent is recorded during the Betula maximum in PAZ 4 we
regard this as a regeneration phase. This phase lasted ca.
690 years and is comparable to regeneration trends recor-
ded in pollen diagrams from the Elbe-Weser area after
human disturbances (Behre and Kucan 1994). The
longevity of the regeneration phase is probably due to the
considerable leaching of the already poor sandy soils as a
result of human disturbance. Betula, a tree that can cope
well with poor, highly leached sandy soils, was favoured.
Increased leaching also resulted in expansion of heathland
during the Neolithic and later periods in this region (Behre
and Kucan 1994; Behre 2000), though this development is
not clearly reflected in the pollen profiles presented here.
Synchronous with the woodland disturbances a shift in
the hydrological regime is suggested by high Sphagnum
spore values after 2520 cal. B.C. (47 cm). According to
Freund (1994) this might be related to drier conditions as
Sphagnum tends to increase spore production under dry
(less favourable) conditions. Bauerochse (2003) argued
that higher Sphagnum values might reflect lowering of the
groundwater table that would favour formation of Sphag-
num peat. Typical open-water taxa (Hydrocotyle vulgaris,
Utricularia, Scheuchzeria palustris, and Sparganium-type)
Table 3 PAZ description of Holschkenfehn pollen spectra and macrobotanical remains
PAZ Depth
(cm)
Age (B.C.,
model)
Main zone characteristics Macro-botanical remains
5 23–16 170 B.C.–
A.D. 390
Ulmus and Tilia present with very low values, Fagus strongly
increases. Poaceae, Secale, and Rumex increase. Sphagnum
values decline, higher fungal spores values (Gelasinospora
and HdV-18)
Sedge rootlets, J. effuses seeds, Calluna vulgaris
twigs
4b 34–23 1570–170 Betula and Alnus decline, Quercus increases. Pteridium,
P. lanceolata and other herb taxa show scattered records.
Poaceae increases, Secale appears
Sedge rootlets, Carex sp. fruits, Sphagnum leaves
4a 42–34 2260–1570 Betula and Alnus are increasing. Quercus, Ulmus, and Tilia
are decreasing. Only sporadic occurrence of Pteridium,
P. lanceolata and other herb taxa, very low Poaceae values.
Local Sphagnum minima, Botryococcus only sporadic, no
charcoal
Sedge rootlets, Carex fruits, Sphagnum leaves
3c 48–42 2570–2260 Betula and Pinus percentages decrease significantly, Ulmus
and Tilia decrease. Corylus shows highest percentages
Fagus is continuously present from this zone on; Carpinus
records begin. Poaceae and Pteridium show highest values,
P. lanceolata and Rumex are decreasing. Sphagnum and
Copepoda spermatophores increase strongly
Sedge rootlets, Carex fruits, Sphagnum leaves
3b 57–48 2950–2570 Ulmus and Fraxinus show higher values than before.
P. lanceolata decreases slightly, Pteridium rising strongly
Sedge rootlets, Carex fruits, Sphagnum leaves,
J. effusus seeds
3a 71–57 3520–2950 Betula, Alnus, Ulmus, Tilia decreases. P. lanceolata,
R. acetosa sharply increase, Jasione montana shows highest
values, cerealia-type pollen continuously present, Poaceae
are rising. Hydrocotyle vulgaris, Utricularia, and
Sparaganium-type show highest values in zone III a/b, algae
are well represented. Charcoal input increasing
Sedge rootlets, Carex fruits, Sphagnum leaves,
J. effusus seeds
2 89–71 4180–3520 Quercus, Tilia and Fraxinus values increase followed by
Ulmus. Betula gradually declines. Pteridium is increasing.
Sphagnum generally increasing, Gelasinospora and HdV-18
increase. Highest Botryococcus values and highest charcoal
values
Sedge rootlets, Sphagnum leaves, J. effusus fruits,
Cenococcum geophilum, unident leaf
fragments, Calluna twigs
1 108–89 5190–4180 Alnus, Betula, Corylus and Quercus are the dominant taxa.
Ulmus declines at the end of the zone. Herbs (Asteraceae,
Fabaceae, Filipendula etc.) are comparably high. Sphagnum
and algae values are comparably low; records of fungal
spores such as Gelasinospora and type HdV-18
Sedge rootlets, Sphagnum leaves, Carex nuts,
E. tetralix twigs
Veget Hist Archaeobot
123
decreased and other dryness indicators like Assulina mus-
corum, Callidina angusticollis and Copepoda spermato-
phores increased so conditions may well have been drier
during this time at Holschkenfehn.
Woodland composition remained stable between 1570 and
170 cal. B.C. (34 and 23 cm) (in PAZ 4b), while occurrences
of Secale, high values of Rumex acetosella and a decrease in
AP, suggest enhanced farming in PAZ 5 (23–16 cm, 170 cal.
B.C.–cal. A.D. 390). Since the major focus of the study was the
Neolithic, the temporal resolution of the record after 1500
B.C. is lower (*500 years) compared with *40 years during
the Neolithic. This part of the Holschkenfehn record is
therefore less detailed but, nevertheless, provides a useful
general indication of developments–increased human impact,
including rye cultivation–during this time.
Considerations regarding the Neolithic transition
First indications of the presence of Neolithic people in
northern Germany are often related to the elm decline. The
elm decline is a widespread phenomenon visible in most
records from north-western Europe shortly after 4000 cal.
B.C. (Tipping et al. 1993; Andersen and Rasmussen 1993;
Peglar 1993; Behre and Kucan 1994; Heider 1995; Kalis
and Meurers-Balke 1998; O’Connell and Molloy 2001;
Molloy and O’Connell 2004; Ghilardi and O’Connell
2013). It is assumed that it originates from the interaction
between the spread of a disease of elm and the impact of
the first Neolithic farmers. A shift to a more continental
climate may have also played a role in the elm decline
(Moe and Rackham 1992; Peglar and Birks 1993; Parker
et al. 2002).
The elm decline does not constitute a distinctive feature
in the pollen profiles presented here as Ulmus does not
exceed 3 %. Elm was probably present in the area (cf.
Huntley and Birks 1983) but the rather infertile, old mor-
ainic soils were hardly favourable for this edaphically
demanding tree. Declining elm values as recorded in the
Hummling profiles (cf. PAZ boundaries 1 and 2) predates
the classical elm decline in northern Europe by about
300 years. This might be attributable to an earlier (human)
impact on elm prior to the spread of an elm disease (cf.
Bakker 2003; Feeser et al. 2012). In the Holschkenfehn
profile, Ulmus declined rather gradually, from ca. 4270 cal.
B.C. (91 cm) onwards. Whether this early decline resulted
from Neolithic activity is difficult to say. The PCA ordi-
nation suggests little change (Fig. 7) and evidence for
pastoral or arable farming is lacking (P. lanceolata is first
recorded at 3520 cal. B.C.–71 cm). The records for cerealia-
type pollen before the declining of elm probably originate
from wild grasses such as Glyceria fluitans (Beug 2004)
that is common in wet environments. A potential indicator
Fig. 7 Plot of results of PCA
analysis of Holschkenfehn
pollen data. Spectra relating to
the various PAZs are enclosed
within envelopes
Veget Hist Archaeobot
123
of human-induced disturbances is Pteridium aquilinum that
expanded as Ulmus declined. This fern is considered to
reflect wood pasture and fire-induced forest clearings
(Behre 1981). The practice of ‘slash and burn’ was prob-
ably important for Neolithic people in Germany (Rosch
et al. 2002) and new results from the TRB North Group
also indicate early use of fire (Feeser et al. 2012). We
assume that the Holschkenfehn charcoal record is more
related to activities at local settlement sites as no direct
correlation between woodland cover, spread of Pteridium
and enhanced charcoal input is visible.
Records of spores that may originate from coprophilous
fungi, such as Sordaria, Cercophora (Van Geel 1978) and
Sporormiella (Davis and Shafer 2006), point to wood pasture
but wild animals cannot be excluded. We favour the latter,
given that these spore types were also recorded prior to
declining elm values and the first occurrence of P. lanceo-
lata, the classic Neolithic settlement indicator (Iversen 1941;
Behre 1981; Behre and Kucan 1994).
In the Bockholter Dose, Ulmus declines around 4250 cal.
B.C. (205 cm) with a quick recovery after 250 years and a
second decline again around 3620 cal. B.C. (160 cm) while
settlement indicators did not register before 3800–3620 cal.
B.C. (170–160 cm). As pollen spectra from the Bockholter
Dose reflect regional woodland vegetation patterns, settle-
ment-indicator pollen taxa and herb pollen in general is
underrepresented (Hicks 1971). However, small decreases in
Betula, Tilia and Alnus and slightly increasing Poaceae
values after 4180 cal. B.C. (195 cm) point to woodland
openings that in turn may indicate human activity. The
DCCA data suggest a distinct change in woodland compo-
sition between 4250 and 4050 cal. B.C., after which there is a
distinct change in the opposite direction which may point to
woodland regeneration. The appearance of P. lanceolata at
3800 cal. B.C lagged declining elm values by 450 years and is
associated with a decrease in Betula and an increase in Po-
aceae and Quercus. All together, there is no general pattern
of vegetation responses to possible human activities. Eval-
uation of single woodland taxa, settlement indicators and the
comparison between pioneer trees (e.g. Corylus, Betula,
Fraxinus) and late succession trees (Ulmus, Tilia, Quercus)
(Kalis et al. 2003) does not suggest any persistent trend in
vegetation dynamics. However, the evidence points to
woodland opening and possibly small-scale clearings in
areas with differing woodland composition as dictated by
successional and other factors.
The Neolithic farming practices that had led to the vege-
tation changes at the beginning of the Neolithic are difficult to
Fig. 8 PCA plot from the
Bockholter Dose. Spectra
relating to the various PAZs are
enclosed within envelopes
Veget Hist Archaeobot
123
assess by palynological investigations. Troels-Smith (1954)
assumed that cutting leaves and twigs (from elm) for fodder
for the stabled livestock and small-scale clearing of the
woodlands for cultivation of cereals were the main techniques.
He excluded wood pasture as Poaceae values were not par-
ticularly elevated. This model was also used by Behre and
Kucan (1994) for the interpretation of several Neolithic pro-
files from the Siedlungskammer Flogeln. They connect the so-
called leaf-fodder period to a ‘pre-megalithic culture’. Com-
parable interpretations have been applied to early Neolithic
developments as documented at in the Alps (Rasmussen 1993;
Akeret et al. 1999) and the Netherlands (Casparie et al. 1977).
A recently published work investigated the stable-isotope
composition of tooth enamel of livestock from the Neolithic
village site Bercy in Paris, France. The results imply that
feeding on leaves took place during winter which probably
prolonged the period of cattle breeding from three to almost
6 months, and thus possibly securing dairy-product supply
throughout the year (Balasse et al. 2012). Seasonal feeding
that included wood pasture during summer and leaf fodder
during winter might have been possible also in a north-west
German context where such practices persisted well into the
historical period (Burrichter and Pott 1983; cf. Bakker 2003).
Pointers in the Bockholter Dose profile suggestive of wood-
land pasture include increases in Poaceae and Pteridium as
Ulmus declines and P. lanceolata records commence.
Increasing Quercus and Corylus values also support this view.
In a review of woodland structure during the Linearbandke-
ramik (LBK) in Central Europe, Kreuz (2008) points out that
high Quercus and Corylus values are probably indicative of
open woodland suitable for wood pasture. A combination of
grazing and leaf-foddering is also assumed by Bakker (2003)
from palynological investigations on the Drenthe Plateau, the
Netherlands. He found a comparable expression and timing of
the elm decline, and sometimes delayed appearance of
P. lanceolata followed by increasing Poaceae values. He
related ‘‘Neolithic Occupation Period I’’ to the Swifterbant
culture that was established in the Netherlands
from *5000 cal. B.C. (Out and Verhoeven 2013; Raemaekers
2013). It seems that the Neolithic transition took place as a
gradual process within the Swifterbant culture from ca.
4500 cal. B.C. when animal husbandry began, while the first
signs of cereal cultivation date to ca. 4200 cal. B.C (Cappers
and Raemearkers 2008; Louwe Kooijmans 2009; Out 2009;
Huisman et al. 2009; Raemaekers 2013). A comparable neo-
lithisation process has also been put forward for areas to the
east of Hummling where comparable palynological data have
been interpreted as indicative of different stages in the Neo-
lithic transition within the Ertebølle-Ellerbek culture and TRB
North Group (Kalis and Meurers-Balke 1998; Hartz et al.
2000; Kirleis et al. 2012).
The pollen data from the Bockholter Dose profile might
also be interpreted as indicative of a series of gradual
developments in the early Neolithic that included scattered
openings in the woodland, these possibly dating to before
the classical Elm Decline recorded in other areas in
northern Europe, and varying vegetation responses to early
farming activities. There are indications of grazing also in
the Holschkenfehn profile before the decline of elm.
Mesolithic activity, involving improving grazing condi-
tions for wild animals or part adoption of a Neolithic
subsistence strategy in the form of livestock breeding, may
be involved. Indeed, grazing of wild animals may have
produced these disturbances.
While clear archaeological evidence is lacking for early
Neolithic in the region, settlement sites such as Hude I in
the Dummer (Schutrumpf 1988; Kampffmeyer 1991), Bo-
berg at the river Elbe (Huser 2009) and Sievern 114, in the
Cuxhaven district (Nosler et al. 2011; Kramer et al. 2012)
point to the presence of early Neolithic settlers in north-
western Germany, although details regarding the nature
and extent of the activity and the chronology have yet to be
resolved. Raemaekers (2013) argues for analogous neo-
lithisation processes in north-western Germany and the
Netherlands, a view that receives support in the interpre-
tation of Mesolithic lithic artefacts from Lower Saxony as
indicative of a more or less closed cultural group in Lower
Saxony and the Netherlands (Mahlstedt 2012).
Human impact ascribable to the TRB West
The vegetation change as recorded in the regional pollen
profile and species turnover (Figs. 4, 9) at Bockholter Dose
between ca. 3600 and 3500 cal. B.C. involves more long-
lasting modifications in woodland composition. Ulmus,
Tilia, Fraxinus and Quercus decrease while Corylus
increases strongly and Fagus is established in the region
(Fig. 4). This indicates that woodlands opened and Corylus
quickly occupied the resulting gaps or experienced
enhanced flowering due to increased availability of light.
Opening-up is also suggested by the PCA ordination plot
(Fig. 8) where the distribution of samples on axis 1, which
is regarded as reflecting a landscape openness gradient,
shows a gradual trend to more open landscape and negative
values after 3500 cal. B.C. (Fig. 9). The pollen spectra from
Bockholter Dose reflect a regional picture of vegetation
changes while spectra from the small site Holschkenfehn
enable direct inference of local Neolithic land use (Behre
and Kucan 1994). Here, changes in wood composition
coincide with sharp increases of human settlement indica-
tors at the site after 3490 cal. B.C. The first period of
woodland disturbance lasted ca. 300 years, i.e. until
3230 cal. B.C. (70–65 cm) when tree pollen taxa (Betula,
Tilia) recovered and P. lanceolata decreased. Between
3050 and 2870 cal. B.C. (60–54 cm), P. lanceolata values
Veget Hist Archaeobot
123
increase again and also AP taxa which suggests that stable
forest vegetation, especially Ulmus and Tilia, declined.
Recently, several pedological and pollen analytical
studies have been carried out in the vicinity of Neolithic
and bronze age burials to investigate the degree of local
landscape openings (Dreibrodt et al. 2009; Demnick et al.
2011; Fyfe 2012; Sadovnik et al. 2012). Holschkenfehn is
located close to nine megalithic tombs, seven of which lie
within 100–250 m (Fig. 1c). It is to be expected therefore
that the impact of the construction and use of the tombs
on the local environment might be inferable from the
Holschkenfehn pollen data. The phases of landscape open-
ness described above probably largely reflect the influence
of local settlements. P. lanceolata, Scleranthus annuus,
Rumex acetosella-type and also cerealia-type records are
presumed to reflect local farming and, in particular, arable
activity (Behre 1981). To date, two possible TRB settlement
sites have been identified, based on find scatters to the south
and west of the Holschkenfehn site (Fig. 1c).
The vegetation changes appear to mirror the pattern of
Neolithic land use as reflected by pollen profiles in the
Elbe-Weser area, and referred to as Landnam sensu Iversen
(1941) and relating to the TRB culture (Dorfler 1989;
Behre and Kucan 1994; Heider 1995). Substantial opening-
up of the woodland cover indicates greater land use than
before, and is probably related to more extensive wood
pasture and cereal cultivation with barley, emmer and
einkorn as major crops, the latter supported by macro-
botanical records (Behre and Kucan 1994; Kirleis et al.
2012). Cerealia records, i.e. Hordeum-type and unidentified
cereal-type pollen but excluding Secale which is identifi-
able with a high degree of certainty, probably also reflect
cereal growing. Further differentiation of the cerealia-type
pollen as regards size etc. may help in distinguishing pollen
of cereal origin from large cereal-like pollen produced by
non-cultivated grasses (cf. Ghilardi and O’Connell 2012).
Why the major impact on vegetation took place at ca.
3500 cal. B.C has been much discussed. In north-western
Germany and the Netherlands the timing coincides with the
emergence of the TRB West Group (Brindley 1986; Muller
et al. 2010) as a clearly defined Neolithic culture. Similar
changes are visible in Schleswig–Holstein. There, the first
evidence for the Neolithic more or less coincides with changes
in woodland composition at 4100 cal. B.C. (Muller et al. 2010;
Feeser et al. 2012). The much later strong increase in human
impact has been interpreted as a change from subsistence,
mainly based on husbandry, to a farming economy with
increased emphasis on arable activity (Behre and Kucan
1994). Other theories involve technological developments
coinciding with the introduction of the ard (Wiethold 1998).
The earliest dated plough marks in Lower Saxony, recorded
close to Wittenwater (Tegtmeier 1993), point to the intro-
duction of ploughing between the TRB and the early bronze
age. However, the ard might be less efficient than hoes for
cultivation (Bogaard 2004) and tillage in areas recently under
woodland would not have been without difficulty due to
persistent root systems (Rosch et al. 2002). In Ireland, changes
in Neolithic subsistence strategies have been ascribed to cli-
mate change (precipitation and also temperature; Stolze et al.
2012) but it remains uncertain if climate had such a dominant
influence (cf. Ghilardi and O’Connell 2012). Evidence
gleaned from plant macroremains from Neolithic sites in
northern Germany, and especially the shift in emphasis away
from plant material collected in the wild to crops, imply a
change from a surplus strategy to a full Neolithic lifestyle
(Kirleis et al. 2012). Taken in conjunction with the emergence
Fig. 9 Summary of vegetation changes and settlement activity as
derived from openness of vegetation, compositional species turnover
and human indicator pollen curves; main settlement phases are
indicated by shading
Veget Hist Archaeobot
123
of the TRB West Group in north-western Europe, this appears
to be an acceptable explanation also for the intensification of
land use at ca. 3500 cal. B.C. recorded in the pollen profiles
presented here.
Neolithic land-use pattern
The close proximity of the Holschkenfehn pollen profile to
the archaeological evidence facilitates the reconstruction of
different phases of Neolithic land use in the area. Small
scale agriculture and wood pastures are assumed for the
period between 3520 and 2870 cal. B.C. (71–54 cm) with a
decrease in settlement activities between 3230 and
3050 cal. B.C. (65–60 cm) and lower impact between 3050
and 2870 cal. B.C. (60–54 cm). This is different to the sit-
uation at Flogeln during the Neolithic where Behre and
Kucan (1994) assumed uninterrupted arable activity and
continuous settlement. Given the poor sandy soils, highly
susceptible to leaching, fertility would have rapidly
declined if no manuring took place. This, in turn, would
lead to new clearances. Further clarification of various
aspects of the Neolithic economy, such as shifting culti-
vation and/or use of permanent plots, requires further
research including archaeobotanical investigations (cf.
Bogaard 2002; Kreuz and Schafer 2011).
Between 2870 and 2260 cal. B.C. (54–42 cm), the type of
human impact changed. Considering the strong representation
of Pteridium and sharp declines in Betula, Ulmus, Pinus and
Fraxinus, intense grazing in the context of cleared woodland
is assumed. This is also supported by a sharp increase in Po-
aceae between 2420 and 2310 cal. B.C. (45–43 cm) which is
regarded as indicative of grassland expansion. These changes
seem to parallel the emergence of the single grave culture
(SGC) in northern Europe (Muller et al. 2010). Unfortunately,
there is little archaeological evidence of settlement pattern and
subsistence practice during the SGC period in the study area
(Strahl 1990; Nosler et al. 2011). The results from Bockholter
Dose show only minor changes which might imply intensifi-
cation of land use on a very local level which is also favoured
by Bakker (2003) for the Drenthe Plateau. That the soil
became exhausted due to crop cultivation has often been cited
as an argument for the abandonment of grazed and arable land
at the end of the TRB (Behre and Kucan 1994; Wiethold
1998). This may also explain the developments seen in the
Holschkenfehn profile at this time. The longevity of the sub-
sequent regeneration phase, involving first birch and then oak,
supports the idea of soil exhaustion and ultimately the aban-
donment of settlement at Holschkenfehn.
The character of the human activity cannot be so easily be
reconstructed on the basis of the evidence provided by the
Bockholter Dose profile. However, changes in woodland
composition and a trend to more open vegetation are in syn-
chrony with phases of TRB settlement intensification and
reduction as reconstructed by pollen indicator taxa from
Holschkenfehn (Fig. 9). This suggests that settlement patterns
are regional in character. There appears to have been high
variability in climate in northern Europe at the beginning of
the Neolithic (Charman 2010). Investigation of lake sediments
and tree-ring data suggest warm summers with mild winters
and increasing wetness in the early Neolithic (after ca.
4000–3800 cal. B.C.) (Leuschner et al. 2002; Dreibrodt et al.
2012). After that, the climate became progressively drier until
2600 cal. B.C. while temperatures declined considerably at
3350 cal. B.C. (Leuschner et al. 2002; Dreibrodt et al. 2012).
Based on the data obtained so far, no direct connection
between regional settlement activities and changing climatic
conditions can be made for the Hummling region.
Conclusions
High percentages of AP pollen indicate that, prior to and
during the early Neolithic, closed canopy woodland dom-
inated in Hummling. Woodland composition changed
during the Neolithic but early opening-up of the woodlands
may be under-estimated due to shifts in tree composition
from low to high pollen producers (cf. Rasmussen 2005)
and also increased overall AP production due to increased
light availability as a consequence of human disturbance.
The mid Holocene decline in Ulmus pollen representa-
tion in Hummling predates the classical elm decline in
northern Europe by ca. 300 years and is only weakly
expressed in both profiles. The decline as expressed in
these profiles may be attributable to an early farming
impact, independent of any disease factor which may not
have been important anyhow given the sparse elm popu-
lation (cf. Bakker 2003).
The timing of the overall vegetation changes as reflected
in the shift in compositional turnover of AP taxa and the PCA
data from Bockholter Dose are in good agreement with the
chronology of Neolithic cultures in north-western Europe.
Initially, farming was probably small scale and the responses
of the vegetation to farming were accordingly varied.
On the basis of the pollen data, there appears to be little or
no change in type of human impact in the region during the
early Neolithic (prior to 3520 cal. B.C.), but human impact
increased considerably after 3520 cal. B.C. and continued until
2260 cal. B.C. A change from a subsistence-type farming
economy with little wood pasture and arable farming to more
intensive wood pasture coincides with the emergence of the
SGC. The underlying factors that influenced these settlement
dynamics have still to be clarified, especially the possibility
that they are connected with changing climatic conditions (cf.
Berglund 2003, Kalis et al. 2003; Schulting 2010).
The palynological evidence from prior to the emergence
of the TRB does not unambiguously prove the presence of
Veget Hist Archaeobot
123
Neolithic people in the study area. On the other hand, the
vegetation changes parallel developments seen in pollen
profiles from neighbouring regions such as the Netherlands
and Schleswig–Holstein where the Neolithic transition is
regarded as a gradual process that included the adoption of
various Neolithic subsistence strategies while essentially a
Mesolithic way of life was maintained. It is assumed that
vegetation at a regional level was relatively resilient to low
human impact but that the influence of the TRB led to
compositional change in the woodlands, and to strong
leaching and ultimately exhaustion of soils that is visible,
especially at a local scale during the early SGC.
Apart from questions concerning vegetation dynamics
and settlement history, insights into regional vegetation
developments may contribute to a better understanding of
early greenhouse effects and climate change by assessing
land-cover changes in northern Europe during the mid
Holocene (Ganopolski et al. 1998; Ruddiman 2003; Kaplan
et al. 2011) and hence deserve further careful attention.
Acknowledgments We thank Gerfried Caspers for providing
insights into archived maps from the Hummling, Stefanie Muller for
introduction to analysis of testate amoebae, Ingo Feeser for an
introduction to the OxCal program, Steffen Wolters for helpful dis-
cussions and, together with Jens Luhmann, for help during fieldwork.
An earlier version of the manuscript benefited from constructive
comments by Michael O’Connell, Walter Dorfler and an anonymous
reviewer. This research was supported by the Deutsche Forschungs-
gemeinschaft (DFG Jo 304/3).
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