J Paleolimnol (2013) 49:663678
DOI 10.1007/s10933-013-9683-0
ORIGINAL PAPER
Natural and human-induced environmental changes
in Eastern Europe during the Holocene: a multi-proxy
palaeolimnological study of a small Latvian lake in a humid
temperate zone
Jaanus Terasmaa Liisa Puusepp
Agata Marzecova Egert Vandel
Tiit Vaasma Tiiu Koff
Received: 26 January 2011 / Accepted: 21 January 2013 / Published online: 15 February 2013
Springer Science+Business Media Dordrecht 2013
Abstract This study uses the Holocene lake sed-
iment of Lake K uzi (Latvia, Vidzeme Heights) for
environmental reconstruction with multi-proxy
records including lithology, computerised axial
tomography scan, grain-size analysis, geochemistry,
diatoms and macrofossils, supported by AMS radio-
carbon dating. Numerical analyses (PCA; CONISS)
reveal three main phases in the development of the
lake. Response to the LateglacialHolocene transi-
tion in Lake K uzi took place around 11,300 cal. BP.
Organogenic sedimentation started with distinctive
5-cm-thick peat layer and was followed by lacustrine
sedimentation of carbonaceous gyttja. Several find-
ings of the peat layer with similar dated age and
position at different absolute altitudes indicate that
lake basin was formed by glaciokarstic processes. In
the Early Holocene (until around 8,500 cal. BP), the
lake was shallow and holomictic, surrounded by
unstable catchment with erosion and inflow events.
Predominance of diatom species of Cyclotella and
Tabellaria, large numbers of respiratory horns of
phantom midge pupae (Chaoboridae), high Fe/Mn
ratio, as well as the presence of laminated sediments
J. Terasmaa (&)  L. Puusepp  A. Marzecova 
E. Vandel  T. Vaasma  T. Koff
Institute of Ecology, Tallinn University, Uus-Sadama 5,
10120 Tallinn, Estonia
e-mail: jaanus.terasmaa@tlu.ee
indicates the transition to a dimictic and oligo-
mesotrophic lake conditions with high water level,
anoxia in the near-bottom and stable catchment in
the Middle Holocene (8,5002,000 cal. BP). This
contrasts with many hydrologically sensitive lakes in
Northern and Eastern Europe in which the water
level fell several meters during this period. During
the Late Holocene (from 2,000 cal. BP to the
present), the lithological and biotic variables reveal
major changes, such as the increase in erosion
(coarser grain-size fraction) and eutrophication [dia-
toms Aulacoseira ambigua (Grun.) Sim., Stephano-
discus spp., Cyclostephanos dubius (Fricke) Round].
Characteristics of lake-catchment system during the
Late Holocene reflect anthropogenic signal superim-
posed on the natural forcing factors. To date, the
Late Quaternary palaeolimnological reconstructions
using lake sediment has been limited in the Baltic
region. Therefore, findings from Lake K uzi provide
important information about environmental and
climatic changes that took place in this part of
Eastern Europe. This study shows that the relative
importance of climate and local factors has varied
over the time and it is essential to consider the lake
basin topography, catchment size and land cover as
potential dominant forcing factors for changes in
sedimentary signal.
Keywords Diatoms  Macrofossils  Grain size 
Geochemistry  CAT scan  Holocene  Lake level
reconstruction
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Introduction
Up to now, little use has been made of sediments of
lakes in the Baltic region for climate reconstruction. It
is important to understand processes in the region as it
is a transitional area from maritime to continental
climate, characterized by a strong westeast gradient
in the continentality of climate and the distribution of
precipitation (Jaagus et al. 2010).
Evidence for occurrence of a number of distinct
climatic oscillations in Europe has been widely
studied (Korhola et al. 2002; Lotter and Birks 2003;
Ilyashuk et al. 2005; Brooks 2006; Seppa et al. 2009;
Enters et al. 2010). Quantitative reconstructions of the
climate in Northern Europe and the Baltic region
during the Holocene are primarily based on pollen
(Seppa et al. 2009; Heikkila and Seppa 2010) or
chironomids (Korhola et al. 2002; Ilyashuk et al.
2005), yet the reconstructions exhibit several signif-
icant dissimilarities. Contrasts exist between the rather
high Early Holocene July temperatures in Kola
Peninsula reconstructed by Ilyashuk et al. (2005)
using chironomids and relatively low Early Holocene
summer temperatures in Latvia inferred from pollen
(Heikkila and Seppa 2010). Therefore, a question
arises whether the differences in interpretation are due
to different proxies or variations in regional climate. In
regional climate reconstructions, one of the key factors
is precipitation, which is not fully considered in
pollen-based studies. At temperate latitudes with
moist climate, the composition and formation of
sediments generally is closely connected with hydro-
logical balance as reflected in contemporaneous
changes in water level (Punning et al. 2005).
Lake-level records can provide a means to deter-
mine regional effective moisture because they reflect
the water table. However, studies of the water levels in
the lakes of the Baltic region (Saarse and Harrison
1992; Koff et al. 2005; Punning et al. 2005; Novik
et al. 2010) have shown variable patterns. It is still
unclear whether the fluctuations reflect regional
hydrological trends or local variations, because most
fluctuations were not as synchronous as they would be
if connected to climate change. This asynchrony
cannot be always explained by dating uncertainties.
There is some evidence that the role of climate
lessened with time, and that geochemical and litho-
logical variations were caused mainly by other local
factors (e.g., morphological structure of the catchment
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J Paleolimnol (2013) 49:663678
area, lithology of the bed rocks, genesis and shape of
the lake depressions, intensity of the water cycle,
position of the subsoil water level; Fritz 2003).
In recent palaeolimnological studies (Birks and Birks
2006), application of multiple indicators is considered
especially important for a reliable reconstruction. More
high-resolution, multi-proxy palaeolimnological stud-
ies are required until a more detailed picture of moisture
conditions in various regions during the Holocene is
achieved. The use of comparable zoning by a number of
different proxies may help to distinguish phases of
stability or change in lakes. If boundaries associated
with different proxies coincide, this indicate substantial
changes in one or more environmental factors, but if
boundaries do not coincide, this might be a result of
events that are important only for one particular variable
(Lotter and Birks 2003).
The main aim of the current study was to find new
palaeolimnological evidence of major environmental
changes in Eastern Europe, specifically in the Baltic
region. Based on analyses of sediment lithology,
geochemistry and biotic remains (diatoms and macro-
fossils), we determined major phases (natural and
human-induced) in the development of Lake K uzi, a
small lake in humid temperate zone of Latvia. We also
distinguished changes in climatic conditions at a
regional level from those related to local changes in
the development of the lake basin and its catchment.
Study site
Lake K uzi (Fig. 1a) is located in Central Latvia in the
western part of the Vidzeme Heights (5720 N,
25200 E; absolute height 191.5 m a.s.l.). The bedrock
of the area consists of Upper Devonian sedimentary
rocks covered by thick (100140 m) Quaternary
deposits, mainly last glacial (Weichselian) tills. The
age of the Last Glacial Maximum moraine is between
14,500 and 15,500 cal. BP (Kalm 2006). The topog-
raphy of the area is hilly, with elevation range
180240 m, and it is dominated by subglacial land-
forms (Zelcs and Markots 2004). The Vidzeme
Heights have a moist climate with a mean annual
precipitation of about 800 mm. The mean monthly
temperature varies from -7.5 C in January to
16.5 C in June (Jaagus et al. 2010). The growing
season lasts for 175185 days and the area is covered
with snow from November to April.
J Paleolimnol (2013) 49:663678 665

Fig. 1 a Location of the study site, b topography and the location of the lake, c bathymetry and coring sites and d cross section of the
lake with lithological descriptions and radiocarbon dates of the sediment

123
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Lake K uzi has a surface area of 6.3 ha (maximum
length 380 m, width 210 m, maximum depth 8 m,
average depth 3.7 m and average slope 5.8 %) and
total water volume of 240,000 m3. The mean resi-
dence time is 0.6 years. The lake was closed, but
presently there is a limited flow through artificially
created ditches. In the relatively large 1.55 km2
catchment, spruce [Picea abies (L.) Karst.] forest
alternates with meadows and agricultural land. The
NW part of the lake is surrounded by a 100-m-wide
mire (Fig. 1b). Absolute height of the slopes in the
south of the lake is low. The geomorphological
features of Lake K uzi (a closed lake with proportion-
ally large catchment area and predisposition for
overflows) indicate that the lake hydrology is sensitive
only to the large-scale climatic changes in the region.
There is no archaeological evidence of human
habitation in the lake proximity. In Latvia, archeolog-
ical evidence and pollen analyses date the emergence
of crop cultivation to about 6,000 years ago (Ozola
et al. 2010). Farming became the predominant form of
economy in Latvia in the middle of the Bronze Age (ca
3,000 cal. BP; Vasks et al. 1999).
Methods
Sediment coring and chronology
Coring was carried out on a longitudinal transect
across the lake (Fig. 1c, d). Two sediment cores
(KS089, KS0827) were retrieved with a small Belarus
(Russian) sampler for the initial litostratigraphical
survey. The main core (KC081) from the central part
of the lake and marginal core (KC082) were obtained
from a raft in May 2008. Another marginal core from
the adjacent mire (KM071) was obtained in 2007. The
Table 1 Radiocarbon dates obtained from Lake K uzi, showing depths for core KC081, dated material, laboratory number, 14C age
BP and 14C-calibrated age BP. The dates were calibrated using IntCal09 (Reimer et al. 2009) in OxCal v4.1.3 (Bronk Ramsey 2008)
Depth, Dated material Laboratory
cm number
260265 Picea seeds Poz-26181
365370 Picea needle, Betula fruits, Salix Poz-26182
bud scales
460465 Betula fruits, Picea seeds Poz-26184
660665 Betula fruits, Alnus fruits Poz-26185
860865 Wood remains Poz-26186
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J Paleolimnol (2013) 49:663678
upper unconsolidated sediments (80 cm) were sam-
pled using a modified LivingstoneVallentyne piston
corer (Vallentyne 1955) with a diameter of 7 cm. The
cores were divided into 2-cm sections and packed into
PVC boxes. The single lower sediment core was
sampled using a Belarus (Russian) peat sampler
(diameter 10 cm) with casing. The core was described
in the field, and then packed into bisected tubes for
transportation. The marginal cores (KC082, KM071)
are described in detail elsewhere (Kangur et al. 2009;
Puusepp and Kangur 2010). The sediment from main
core KC081 was used for this multi-proxy study.
The chronology of the sediment core KC081 was
determined using five AMS 14C dates obtained from
terrestrial macrofossils (Table 1). The samples were
dated in the Poznan Radiocarbon dating laboratory,
Poland. The age-depth model was constructed using
the depositional model option in OxCal v4.1.3 (Bronk
Ramsey 2008). Samples were calibrated to calendar
years using the IntCal09 calibration curve (Reimer
et al. 2009). Rates of accumulation were calculated
using the dry mass of the sediment samples, together
with five AMS dates obtained from core KC081.
Lithology and geochemistry
Core KC081 was subsampled contiguously at 2-cm
intervals for loss-on-ignition (LOI). Dry matter con-
tent in sediment was determined by drying the samples
at 105 C to constant weight. Dried samples were
ignited in two-step process, at 550 C for 3.5 h to
determine organic matter content, followed by an
ignition of LOI550 residues at 950 C for 2.5 h for
estimation of carbonates. Organic matter, CaCO3 and
siliciclastic mineral matter were calculated accord-
ing to standard methods (Boyle 2001; Heiri et al.
2001).
14C date, Calibrated age (cal. BP, 2r Weighted average (l)
year BP interval) (cal. BP)
2,508 30 2,7462,680 2,714
4,553 35 5,1124,877 5,034
5,257 40 6,1745,928 6,030
7,123 60 8,1817,973 8,123
9,891 60 11,38511,193 11,268
J Paleolimnol (2013) 49:663678
Grain-size distribution was estimated in 90 sedi-
ment samples using a Fritsch Laser Particle Sizer
ANALYSETTE 22 (measuring range 0.3300 lm).
Samples were chemically pre-treated with 30 % H2O2
to remove organic matter and 10 % HCl to remove
carbonates (Konert and Vandenberghe 1997; Vaasma
2008). Prior to analysis, dispersing agent [1 %
(NaPO3)6] and ultrasonic agitation were used to
prevent flocculation. Grain-size spectra are presented
according to the UddenWentworth scale.
Sediment density was measured with a Somatom
Definition (Siemens) at the Tallinn Diagnostic Centre
Ltd. Computerised axial tomography (CAT) scan
results are expressed in CT numbers in Hounsfield
units (HU) (Hounsfield 1973). Tomographic intensity
was estimated with the software eFilm Light 3.0 and
OSIRIS. CT values were calculated using an average
of at least five sections. A spatial resolution of scans
was 2 mm, but it was reduced by smoothing to 2 cm.
Bulk geochemical analysis was carried out on 48
dried and homogenized samples by X-ray fluorescence
spectroscopy (Boyle 2000; Boyle 2001) using an ED-
XRF analyser (Bruker Axs S2 Ranger with DECONV
software, Department of Geography, Liverpool Uni-
versity, UK). Total organic carbon (TOC) and nitrogen
(N) contents were measured by a CHNS/O Elemental
Analyser (Perkin-Elmer 2400 Series II, University of
Tartu, Estonia). The values were calculated as per-
centages of dry weight.
Diatom and macrofossil analysis
Samples for diatom slides were pretreated with 33 %
H2O2 and 10 % HCl (Battarbee et al. 2001). The
suspension was mounted on glass slides with Naphrax.
At least 500 valves were counted per sample using a
10009 oil immersion objective (N.A. 1.5) on an
Olympus BX41 microscope equipped with DIC optics.
Relative abundance of diatom species was expressed
as percentage of total diatom counts. The concentra-
tion of diatoms was calculated by adding a known
concentration of synthetic microspheres to the sam-
ples and expressed as valves per dry sediment mass
(Battarbee and Kneen 1982). Diatom identification
and nomenclature were based on taxonomic refer-
ences by Krammer and Lange-Bertalot (19881991,
19992004).
For macrofossil analysis (remains of plants, inver-
tebrates and macroscopic charcoal), samples with a
667
volume 100 cm3 from 5-cm-thick sediment sections
were washed with water through a 250-lm sieve, after
which the residues were analysed under a stereomi-
croscope. Macrofossils were counted and identified
according to the reference collections and descriptive
manuals (Birks 1980; Cappers et al. 2006). The results
were expressed as number of macrofossils per
100 cm3.
Lake levels
Evidence from marginal cores can provide additional
information in the documentation of lake level
(Digerfeldt 1986; Punning et al. 2005; Koff and
Vandel 2008). Therefore, lithology, sediment texture
and plant macrofossil remains, especially the distri-
bution pattern of emergent and aquatic species from
marginal cores (KM071 and KC082) were used to
support the lake levels reconstruction from the central
core KC081.
Numerical methods
The numerical database from core KC081 of Lake
K uzi contains total of 18,628 data points (CAT-scan
3,782, lithology 3,680, grain size 2,700, diatoms
4,838, macrofossils 1,624, geochemistry 2,004).
Because of different numerical properties of the data
sets, the data were individually transformed to
approach a normal distribution and reduce the effect
of dominant variables within data-sets (Birks 1995;
Jongman et al. 2002; ter Braak and Smilauer 2002).
A square-root transformation was applied to the
diatom and grain-size percentages, and a log transfor-
mation was applied for the macrofossil data. Geo-
chemical data were mean-centred and standardized by
the standard deviation value. Stratigraphic zones for
each data set were determined using CONISS (Grimm
1987) in the program TILIA (Grimm 1991). To
explore the major patterns of change, the individual
datasets (diatoms, macrofossil, geochemistry and
grain size) were analysed using principal component
analysis (PCA) in the program Canoco 4.5 (ter Braak
and Smilauer 2002). Choice of linear response model
was based on the preliminary detrended correspon-
dence analysis (DCA) in which the lengths of gradient
of each dataset fell within 2 S.D (ter Braak and
Smilauer 2002).
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Results
Sediment chronology and stratigraphy
The age-depth model (Fig. 2) for sediment core
KC081 was constructed using five calibrated AMS
radiocarbon dates (Table 1) and the deposition model
was implemented in program OxCal v4.1.3 (Bronk
Ramsey 2008), assuming sedimentation rates between
modeled dates were linear. All dated material was
terrestrial plant remains (Table 1) to avoid old carbon
effect and to achieve potentially more reliable results.
The ages were not anomalous and they fit with respect
to agedepth stratigraphy. The age of the oldest
sample (11,268 cal. BP; Table 1, Poz-26186) from
peat matched closely with the age of the analogous
peat layer (11,230 cal. BP) from marginal core
KM071 (Kangur et al. 2009).
The mass accumulation rate (MAR) from the
sediment core KC081 was calculated for each date.
The values vary from 87 to 207 g m-2 year-1. The
MAR was highest during the interval 655365 cm
(8,0005,000 cal. BP) and the trends are visualised in
Fig. 2.
Lithology
Based on visual description and photographs, lithol-
ogy of the sediment core KC081 can be divided into
seven lithological units (Fig. 2). A simple correlation
Fig. 2 Age-depth
curve based on five
AMS 14C dates,
lithological units and
calculated mass
accumulation rate
(g m-2 year-1) of
Lake K uzi sediment
core KC081. Error
bars represent
95.4 % confidence
intervals
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J Paleolimnol (2013) 49:663678
analysis of LOI and CAT scan data showed strongest
association (r = 0.93, p \ 0.001) between CT values
and mineral matter content (in g g-1 of wet sample).
This implies that CT values are closely related to the
bulk density of the sediment. Accordingly, the values
can serve as a high-resolution generalised indicator for
the sedimentological changes (Fig. 3a).
Between 920 and 865 cm (before 11,300 cal. BP;
Fig. 3a), the sediment was minerogenically rich
([69 %) and the water content was low. The propor-
tion of organic matter in the sediment began to
increase at around 865 cm (11,300 cal. BP), whereas
below this level it was less than 2 % on average. From
865 to 730 cm (up to 9,200 cal. BP), the lithological
composition of the sediment is very variable. The
CaCO3 content varied from 2 to 29 %. In the grain-
size distribution, there was a peak in the sand fraction
(17.9 %) at around 840 cm (10,900 cal. BP), and after
that the sand fraction disappears until 770 cm
(9,800 cal. BP). Content of medium silt was stable
during same period. The median grain size showed a
similar trend, i.e. a peak around 840 cm (10,900 cal.
BP), followed by a decrease.
Between 740 and 690 cm (9,300 and 8,500 cal.
BP), significant changes in the sedimentary regime
took place. There are two separate peaksthe first at a
depth of 730 cm (9,200 cal. BP) in grain size (median
value 18.4 lm; coarse silt is the dominant material)
and the second at a depth of 710 cm (8,900 cal. BP) in
lithology (91 % organic matter).
J Paleolimnol (2013) 49:663678 669

Fig. 3 Lake Kuzi sediment core KC081. a Lithology and grain- seedwings, needles; Betula spp.seeds, catkin scales; Menyanthes
size distribution, b total concentrations of selected chemical trifoliataseeds; Carex spp.seeds; Nymphaea albaseed
elements and elemental ratios and c the relevant diatom taxa and fragments; Potamogeton spp.seeds; Cristatella mucedostato-
macrofossils. Horizontal white lines denote depth at 100-cm blasts; Chaoboridaerespiratory horns. Because of some very high
intervals, dotted lines show calibrated age (BP) at intervals of values in the lower part of the core, the x-axis is adjusted for some
1,000 years. Types of counted macrofossils: Picea abiesseeds, variables (CaCO3, CT value, mineral matter/organic matter ratio)

123
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At approximately 640 cm (7,900 cal. BP), most of
the lithological indicators showed a degree of stabi-
lisation, and remained relatively constant up to
220 cm (2,300 cal. BP) with some fluctuation in grain
size. Sediment grain size was characterised by
enrichment in the clay fraction (up to 29.5 %) and
almost a disappearance of the sand fraction. A peak at
the clay fraction around 4,700 cal. BP was contem-
poraneous with the peak in mineral/organic matter
ratio and in sediment density.
Between 360 and 198 cm (5,0002,000 cal. BP)
regular laminations were observed. Since their pattern
had lower frequency than is expected for annual
laminations, their seasonal character is not plausible.
Possible genesis and preservation is explainable by
high input of clay fraction from catchment, and in-lake
turbidity currents caused by basin topography and
stable hypolimnetic conditions.
Between 220 and 100 cm (2,3001,000 cal. BP),
the sediment was less minerogenic, but coarser (clay
fraction \9 %), and the ratio of mineral to organic
matter decreased. Between 100 and 0 cm, the content
of mineral matter increased (up to 64 %) and fine-
grained sediment was dominant. The CaCO3 content
showed a clear peak at a depth of 170 cm (around
1,700 cal. BP).
Geochemical stratigraphy
Selected profiles of chemical elements and elemental
ratios are given in Fig. 3b. The major lithogenic
elements were closely related throughout the whole
core. Changes in the lithogenic component are repre-
sented by Ti and Zr/Ti profile. Comparison with
grain-size distribution (Fig. 3a) confirmed that Ti is
predominantly supplied in the clay fraction. Enrich-
ment in Zr relative to Ti was used as an indicator for
the coarsening of the sediment (Boyle 2001). Because
of its affinity to bind to terrestrial organic matter,
bromine (Br), expressed as Br/LOI550 (Fig. 3b), was
used to identify enrichment in organic matter from
terrestrial sources.
In the basal sequence of the record, the high Zr/Ti
ratio alongside with high presence of alkali and
alkaline-earth elements such as Ca, Mg (data not
shown) indicates clastic silicates mechanically eroded
from undeveloped soils. The sharp decrease in the
lithogenic component at a depth of 865 cm was
followed by a period (865690 cm, 11,3008,500 cal.
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J Paleolimnol (2013) 49:663678
BP) of significant fluctuations in the geochemical
record, particularly in S, P, Fe and Mn profiles. In this
interval, Br/LOI550 indicated higher proportion of
terrestrial organic matter. Ti concentrations gradually
decreased to the minimum values around 720 cm
(9,000 cal. BP). In contrast to the low Ti values, the
increase in Zr/Ti ratio showed a temporal coarsening
of the sediment.
After 690 cm (8,500 cal. BP), gradual increases in
Ti indicated transition to more minerogenic and fine-
grained sediments. The elemental profiles remained
relatively stable, with low-amplitude peaks in Ti and
high Fe/Mn ratios until 250 cm (2,500 cal. BP).
The sediment interval between 200 and 100 cm
(2,0001,000 cal. BP) was characterized by a partic-
ular peak in Si/Al ratio indicative of rise in biogenic Si,
and a moderate peak in Zr/Ti ratio. The Ti values were
temporarily low. Based on calculations of enrichment
factor for trace metals, only Pb showed clear,
individual enrichment with two distinctive peaks at
200 and 100 cm (2,000 and 1,000 cal. BP).
The C/N ratio was relatively stable, with values
around 13 throughout most of the record. The highest
value, 17.5, was detected in the lowermost sample at
850 cm (11,100 cal. BP). In the uppermost part of the
core the C/N ratio was low, around 10 (Fig. 3c).
Diatom stratigraphy
In contrast to samples from lowermost sediment
(920860 cm, before 11,200 cal. BP), which con-
tained only few broken diatom valves, the overall
sedimentary record was generally rich in well-
preserved diatoms. Between 850 and 750 cm
(11,1009,500 cal. BP), diatoms were already abun-
dant and the lake was dominated by periphytic taxa
(over 80 %), mainly small species of Fragilaria and
Achnanthes. There were no diatoms in the samples
obtained from depths of 740 and 730 cm
(9,3009,200 cal. BP), but they reappeared in the
sample at 720 cm (9,000 cal. BP).
The interval between 720 and 195 cm (9,000
2,000 cal. BP) was characterised by the significant
occurrence of Cyclotella spp. [C. stelligera Cleve and
Grunow, C. ocellata Pantocsek, C. radiosa (Grunow)
Lemmermann] and Tabellaria flocculosa (Roth)
Kutzing, and the percentage of planktonic taxa then
rose to 80 %. At 410 cm (5,500 cal. BP) the propor-
tion of T. flocculosa reached 50 % and the percentage
J Paleolimnol (2013) 49:663678
of acidophilic and periphytic Eunotia species
increased.
The proportion of the meso-oligotrophic Cyclotella
spp. and T. flocculosa and acidophilic Eunotia taxa
began to decrease at 195 cm (2,000 cal. BP), and the
more eutrophic Aulacoseira ambigua (Grunow)
Simonsen dominated, its percentage rose to 30 %.
Between 135 and 0 cm (from 1,400 cal. BP to
present), the eutrophic Cyclostephanos dubius
(Fricke) Round became more common and Stephan-
odiscus spp. (S. hantzschii Grunow, S. medius
Hakansson, S. parvus Stoermer and Hakansson)
appeared but their percentage never rose above
10 %. Furthermore, the percentage of Cyclotella
radiosa was relatively high during this period (12 %).
The concentration of diatom valves per gram of dry
sediment was quite stable between 800 and 195 cm
(10,3002,000 cal. BP). It increased between 195 and
135 cm (2,0001,400 cal. BP), reached a maximum at
the depth of 150 cm (1,500 cal. BP), and decreased
from 135 cm (Fig. 3c).
Macrofossil data
Although the macrofossil record revealed few species,
it showed several major changes in the composition
and abundance during the Holocene (Fig. 3c). Below
860 cm (11,200 cal. BP), only a few macrofossils
were found. The zone between 860 and 840 cm
(11,20010,900 cal. BP) was dominated by terrestrial
plant remains, including Betula spp. seeds and catkin
scales, Carex spp. seeds, and to a lesser extent by the
limnophytic Potamogeton natans L.
In the interval 840740 cm (10,9009,300 cal.
BP), previously abundant macrofossils gradually
decreased. In the same interval, there was an increase
in the number of seed fragments of Nymphaea alba L.
Beside plant macrofossils, the bryozoan Cristatella
mucedo statoblasts also first appeared at the depth of
840 cm. Their maximum number was detected in the
interval between 740 and 690 cm (9,3008,500 cal.
BP).
Between 690 and 195 cm (8,5002,000 cal. BP),
the only plant macrofossils were a few Picea spp.,
Betula spp. and Salix spp. remains. This zone,
especially the interval from 630 to 254 cm
(7,8002,600 cal. BP), was dominated by a high
number of respiratory horns of Chaoboridae (phantom
midges) that have been only rarely studied from the
671
sediments (Wedmann and Richter 2007). Above the
depth of 195 cm (2,000 cal. BP), the number of
macrofossils, including Chaoboridae, sharply
declined, and several charcoal-rich layers were
present.
Discussion
Zonation scheme
Three general phases were detected in the develop-
ment of Lake K uzi, namely rapid changes in the Early
Holocene (before ca 8,500 cal. BP), a switch to more
stable conditions in the Middle Holocene
(8,5002,000 cal. BP), and effects of human impact
in the Late Holocene (ca 2,000 cal. BP to present).
Major variations in the individual datasets were
expressed through the PCA axes 1 and CONISS-
derived zonation (Fig. 4).
In a number of cases, the zone boundaries closely
coincided, confirming substantial changes in environ-
mental factors, and three main stages of lake devel-
opment. Similar trends were obtained from diatoms
and pollen analysis in other studies of Lake K uzi
(Kangur et al. 2009; Puusepp and Kangur 2010) and
pollen-inferred mean summer temperature reconstruc-
tion from Lake Kurjanovas (Heikkila and Seppa
2010). The synthesis of evidence for Holocene envi-
ronmental change from sedimentary and biological
variables allowed reconstruction of lake development,
water circulation, water level changes and lake-
catchment dynamics (Fig. 5).
Period of rapid changes during the Early Holocene
(11,7008,500 cal. BP)
During the Early Holocene, the water level in Lake
K uzi fluctuated markedly. The lake basin was formed
by minerogenic sedimentation of clay-rich material
with high sediment density. High mineral content of
the sediment reflects erosion from catchment bedrock
and unstabilised soils (high Zr/Ti and Ca values), in
which vegetation was sparse (Engstrom and Wright
1984). Organogenic sedimentation started around
11,300 cal. BP with a clearly distinctive 5-cm-thick
peat layer, followed by lacustrine sedimentation of
carbonaceous gyttja with high content of organic
matter (50 %). Analogous peat layers were detected at
123
672
Fig. 4 Results of principal component analysis (PCA) showing
Axis 1 sample scores for diatoms, macrofossils, geochemistry
and grain size throughout the Holocene and stratigraphic zones
(main zonesdark gray; sub-zoneslight gray) determined by
CONISS (constrained incremental sum of square cluster
analysis). Variance explained by axis 1 and cumulative of axis
1 and 2 are (respectively) for diatoms 30.3 and 54.2 %, for
macrofossils 41.5 and 58.4 %, for geochemistry 43.9 % and
68.2, for grain size 72.4 and 89.1 %. Variables which strongly
the mineral-organic boundary in other cores (Fig. 1c,
d). Formations of peat layers of similar age positioned
at different absolute altitudes usually indicate lake
basins formed by melting of the dead ice-block
(glaciokarstic origin). Given hilly morainic features
of the site, it is plausible that the subglacial inter-hill
depression isolated an ice-block, which became cov-
ered with sediment and meltwater, allowing the
growth of mosses and herbaceous plants forming a
peat layer (Fig. 1d). The melting of the ice, triggered
by the Early Holocene warming, gradually formed a
depression in which sediments began to accumulate
and the lake was formed. As other studies have shown
(Henriksen et al. 2003; Andreev et al. 2005;
Kabailien_e 2006), glaciokarstic lakes are formed
gradually over a long period and the process may be
bidirectional. The depth of the lake basin might be a
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J Paleolimnol (2013) 49:663678
influence scores for axis 1 are for diatoms: (positive) Cyclotella
radiosa, C. ocellata and periphytic Fragilaria spp. and
(negative) Tabellaria flocculosa, Cyclotella stelligera, Eunotia
spp; for macrofossils: (positive) Chaoboridae, Cladocera, Salix
and (negative) Carex seeds, Nymphaea seeds, Potamogeton
spp.; for geochemistry: (positive) S, P, Mn and (negative) Ti, Al,
Ce; for grain size: (positive) very fine sand, very coarse silt and
(negative) very fine silt, clay
function of the melting of the ice buried under the
sediment, with a coincident rise in water level in the
lake basin. It can be expected that both a delayed onset
of organic sedimentation and an unstable shallow
sedimentary environment were influenced by the ice
block. Timing of these clear depositional changes
indicates that in the area of Lake K uzi, response to the
LateglacialHolocene transition took place later than
commonly defined (Walker et al. 2009). This shows
the importance of the local and regional peculiarities,
which lead to the spatial and temporal lag in the
sedimentary records.
Between 11,200 and 10,500 cal. BP, the large
number and variety of macrofossils combined with a
input of coarse sediments reflect a nearshore unstable
sedimentary environment, characterised by shallow
water. The high percentages of dry and mineral matter
J Paleolimnol (2013) 49:663678
Fig. 5 Summary of reconstructions of lake level changes and
maximum depth in coring site KC081, lake productivity, water
circulation pattern and catchment erosion. Landscape dynamics
in the sediment, reflected also in high Zr/Ti ratio, show
a significant degree of surface runoff from the
surrounding drainage area. The diatom flora was
dominated by periphytic taxa, mainly Fragilaria spp.
and Achnanthes spp. These species represent pioneer-
ing communities in the post-glacial period and indi-
cate unstable environmental conditions associated
with strong erosion from catchment and alkaline
waters (Lotter and Bigler 2000; Bigler et al. 2002;
Puusepp and Kangur 2010). At the same time,
simultaneous increases in Fe, S and of Fe/Mn ratio
suggest deposition of iron-sulphides, which are typi-
cally formed under limited water circulation and
hypolimnetic anoxia. Given the otherwise unstable
traits of the lake system, and the fact that organic
matter content was high (50 %), oxygen depletion
would most likely develop during the long winter
period through the degradation of organic matter
(Engstrom and Wright 1984; Enters et al. 2010). The
dominance of periphytic diatoms together with chem-
istry indicative of winter hypolimnetic anoxia imply
that until at approximately 10,500 cal. BP, the lake
was controlled by periods of prolonged ice cover.
Sedimentation processes in glaciokarstic basin can be
considerably affected by a collapse of buried dead ice
block (Henriksen et al. 2003), but the absence of sand
and helophyte macrofossils confirms that Lake K uzi
achieved a higher water level (*5 m) around
10,000 cal. BP.
Between 9,300 and 9,200 cal. BP, an abrupt
temporal disappearance of diatom valves, contempo-
raneous with coarse-grained sedimentation (peak in
673
and characteristic vegetation interpretations are from Puusepp
and Kangur (2010), regional climatic interpretations are from
Seppa et al. (2009) and Heikkila and Seppa (2010)
Zr/Ti, and peak in sand fraction; Fig. 3), suggest a
strong physical degradation and breakage of diatom
valves in the sedimentation process. Hence, consider-
ing the topographical features of the catchment
(Fig. 1b), it is probable that a smaller dammed water
body, located about 800 m north from the basin of
Lake K uzi, gradually filled with meltwater, began to
overflow and eventually broke through, causing water
and material discharge into Lake K uzi. Signs that
could be attributed to breaching of the external water
body and its drainage to the main lake were also
individually detected in the sediment outcrop found on
the adjoining slope (Koff and Terasmaa 2011). At this
site, the Early Holocene organic sedimentation was
interrupted by a sudden deposition of older deluvial
coarse sediments. Such water overflow or flow through
the slope already undergoing soil-forming processes
would cause water-logging and reduction of available
Fe and Mn oxides into their mobile forms (Engstrom
and Wright 1984; Boyle 2001). The mobility and
transport of Fe and Mn is likely to cause remobilisa-
tion of P sorbed on the iron oxides. Thus coupled
signal of Fe, Mn and P in the sediment around
9,000 cal. BP could represent a culmination of the
soil-derived particles from the slope and it supports
the scenario of the strong in-wash. After inflow event
the dominance of periphytic taxa was replaced with
planktonic taxa indicating deeper water conditions.
From 9,100 cal. BP, the switch of diatom commu-
nity to planktonic, smoothing of the lithological
records and substantial increase in temperature sensi-
tive bryozoa Cristatella mucedo (Fig. 5) suggest
123
674
stable aquatic conditions and warmer water tempera-
tures (kland and kland 2000; Bennike et al. 2004).
Based on pollen data (Puusepp and Kangur 2010) the
vegetation around the lake was dominated by alder,
elm, lime and hazel, indicating stabilisation of soils
and warm, moist climate. This is consistent with the
regional climate interpretation which indicated period
of warming from 9,000 cal. BP that lead to the period
of the Holocene Thermal Maximum (HTM) at
8,000 cal. BP.
Period of stable lake level in the Middle Holocene
(8,5002,000 cal. BP)
The first stage of this transition, between 8,500 and
8,000 cal. BP was characterised by a coarser sedi-
mentation. According to the marginal cores macro-
fossil analysis the water level in the lake rose
continuously until ca. 7,000 cal. BP, causing signif-
icant erosion from the surrounding slopes, as also
indicated by an initial peak in the sand fraction and a
gradual increase in clay and Ti values.
Heikkila and Seppa (2010) describe the interval
from 8,500 to 8,000 cal. BP as a period of gradual
progressive warming with a particularly clear nega-
tive short-term punctuation by 8.2 ka of cooling.
Erosional activity might have been intensified under
the cold and wet conditions, however resolution of
this study does not allow decoupling of the long-term
stabilising trend and potential short-time climate
perturbation.
In many hydrologically sensitive lakes in Northern
and Eastern Europe, water level fell several meters
between 8,000 and 4,000 cal. BP (Punning et al. 2005;
Seppa et al. 2009), but this was not detected in our
record. Between 7,000 and 2,000 cal. BP, the lake
water depth was up to 11 m in the KC081 site and for
the whole period the water level was stable. In a
temperate humid zone, this discrepancy can be caused
by decrease in the general role of the climate (mainly
precipitation and humidity) and/or a rise in importance
of indirect or local factors (lake basin topography,
catchment size or land cover).
Lake sedimentary signals reflect stability of the
catchment since 8,000 cal. BP. The Middle Holocene
sediment record (8,0002,000 cal. BP) is character-
ized by a series of well-preserved laminations in the
sediment and high Fe/Mn ratios. Formation of lami-
nations require stable conditions, typically connected
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J Paleolimnol (2013) 49:663678
with anoxic bottom waters, and generally a stable
lake-catchment system with low perturbation by
erosional fluxes. Striking features of the same interval
were high numbers of respiratory horns of phantom
midge pupae (Chaoboridae). The palaeoecological
significance of remains of Chaoboridae is unclear, but
Luoto (2009) has concluded that their remains can be
used quantitatively to infer past water-level changes.
Sweetman and Smol (2006) connected respiratory
horns of phantom midge pupae with severe anoxic
conditions at the bottom of the lake. Close association
of Chaoboridae remains with laminations and Fe/Mn
ratio in the Lake K uzi sediment record support their
indicative potential for a seasonally anoxic hypolim-
nion, as occurs in deeper stratified lakes. From
8,000 cal. BP, the shift from coarse sediment to fine
grained clay (high Ti values) indicated stabilisation of
the lake level. The diatom community was dominated
by planktonic, oligo-mesotrophic or mesotrophic taxa
(Fig. 3c).
Generally, sediment variables indicated stable
conditions over time, but gradual changes around
5,000 cal. BP were noted in several variables. These
included more than two-fold decrease in sedimenta-
tion rate (Fig. 2), a low-amplitude fluctuation in
sediment density, a slight increase in Fe and P, and a
clear increase in acidophilous diatom taxa (Eunotia
spp.) (Fig. 3c). In Lake K uzi, two possible causes
(regional or local) might have occurred: (1) lower
primary production associated with climatic cooling
or (2) the steep sloped sub-basin became filled with
sediments. It has been shown that during the infilling
of a lake, deposits are distributed over a continuously
increasing area (Terasmaa 2005), and if we assume a
constant yearly rate of in-lake production, it implies
the deposition of a thinner layer of sediment each year
(Davis and Ford 1982). The configuration of the Lake
K uzi basin (Fig. 1d) indicates that the rapid broaden-
ing of the accumulation area could have started after
5,000 cal. BP.
Spruce became dominant around Lake K uzi after
ca. 5,000 cal. BP (Puusepp and Kangur 2010) causing
progressive leaching of catchment soils. Higher pro-
duction of humic acids and lowering of the pH would
lead to the progressive leaching of the pH sensitive
elements, presented in the sediment chemistry by
higher P and Fe (Fig. 3b). The increase of acidophilic
Eunotia species suggest that spruce expansion can
lead to lake water acidification and changes in
J Paleolimnol (2013) 49:663678
terrestrial vegetation may have considerable influence
on aquatic ecosystem.
Parallel to the biological and sedimentation change,
the low-amplitude but high-frequency peaks in sedi-
ment density and mineral/organic matter ratio denote
changes in erosion patterns and the appearance of
short-lived events of minerogenic inwash or reworked
sediments after 5,000 cal. BP. The regularity of the
fluctuations, stable water levels and continual signs of
the stability of the lake and catchment indicate that
these changes do not represent a response to the
destabilisation of the catchment. Taking into account
the relative dominance of spruce in the catchment
(Puusepp and Kangur 2010) and onset of regional
cooling (Heikkila and Seppa 2010), a possible hypoth-
esis for these fluctuations could be precipitation
changes, particularly more frequent heavy rainfall.
The available evidence thus suggests that subtle
changes in lake properties after 5,000 cal. BP reflect
shift in climatic conditions mediated through the
vegetation changes in the catchment.
Between 3,000 and 2,000 cal. BP, the accumulation
rate increased (Fig. 2) and the sediment record started
to display a series of changes, which included a
gradual decrease in Ti and in lithology (increase of
grain-size median and decrease of sediment density)
since 2,500 cal. BP, followed by a disappearance of
laminations around 2,000 cal. BP. The declining trend
in Ti was likely caused by a coarsening of the
sediment. As Seppa et al. (2009) describes a warmer
and dryer period that peaked at around 2,000 cal. BP,
these changes might have been triggered by climate.
However it has been shown that agrarian activity
started in Latvia around 2,500 cal. BP (Heikkila and
Seppa 2010), therefore we cannot rule out the early
human influence.
Period of human impact during the Late Holocene
(2,000 cal. BP to present)
From 2,000 cal. BP sedimentary indicators show
simultaneous changes attributed to a human perturba-
tion, most notably increase in lake productivity,
sediment disturbance and fluctuation of water level.
At this time, the lake oligotrophication trend was
reversed by a clear transition to the diatom taxa
(Aulacoseira ambigua, Stephanodiscus spp., Cyclo-
stephanos dubius) indicative for eutrophic conditions.
Simultaneously, absolute diatom productivity rapidly
675
rose more than 3.5 times (reflected also in high Si/Al
ratio). An increase in the abundance of A. ambigua
among diatoms also indicates higher turbidity in the
lake and their presence has been noted in lakes
affected by deforestation (Bradbury et al. 2002).
Charcoal fragments found in the upper part of the
sediment sequence, paralleled by a clear rise in grass
and rye (Secale) pollen (Puusepp and Kangur 2010),
represent a clear sign of forest clearance by fire, with
the aim of increasing the amount of arable land and/or
of opening up the landscape to grazing the cattle.
Higher primary production (higher content of organic
matter with lower C/N ratio), more turbid conditions
due to open landscape and lower water level is the
most probable reason for the disappearance of lami-
nations. Greater exposure to wind (related to the
deforestation) can lead to an increased input of sand
and silt into the lake and to a generally higher median
grain size (Fig. 3a). Similar trends have been noted by
other authors (Enters et al. 2008, 2010), and have been
associated with an increased aeolian minerogenic
input to the lake.
In Lake K uzi distinctive water level drop, up to
1.6 m, was clearly detected in marginal mire core
KM071 around 700 cal. BP (Kangur et al. 2009).
Water-level drop is also reflected in the lithology of
the central core with some differences in timing,
however, this is likely an artifact of lower resolution of
dating in uppermost part of the core. At present, the
lake is controlled by an artificial inflow and outflow
(Fig. 1b). It is probable that the ditches for drainage of
agricultural land are old, and for the past several
hundred years were used to regulate lake level. As a
result of a smaller area of accumulation and of the
reworking of previously accumulated material, both
the clay and Ti curves show a recent rapid increase.
Besides the changes caused by erosion, the chem-
istry of uppermost sediment shows separate enrich-
ment in Pb with two distinct peaks around 2,000 and
1,000 cal. BP. Similar trajectories of Pb enrichment at
other sites in the Northern Europe were connected to
the early anthropogenic airborne Pb pollution from
Roman and Medieval utilisation of Pb ore and
smelting processes (Renberg et al. 2001). Analogous
shape of the Pb profile suggests that Lake K uzi
sediments also bear the signal of this regional
phenomena.
These findings show that the last 2,000 years of
changes in lake hydrology and the ecosystem were
123
676
strongly influenced by continuous human activities,
mainly farming. Since 2,000 cal. BP, human impact
became the dominant driver of lake changes obscuring
the potential influence of natural forcing factors.
Conclusions
The multi-proxy study of sediments from Lake K uzi in
Latvia provides continuous environmental and cli-
matic reconstruction of the Holocene for the temperate
humid zone in the poorly studied region of Eastern
Europe and creates a better prerequisite for its
comparison with well-investigated Northern and Cen-
tral Europe. The Holocene lake development and
inferred environmental change has been summarised
in Fig. 5. The main findings are as follows:
1. According to the sedimentological, geochemical
and biological evidence, Lake K uzi and its
catchment developed during the Holocene
through three main phases-rapid changes in the
Early Holocene (before ca. 8,500 cal. BP), a
period with stable lake level (8,5002,000 cal.
BP), and a period of clear human influence since
ca. 2,000 cal. BP. The LateglacialHolocene
transition at the site was around 11,300 cal. BP,
later than the dates previously reported in the
region, probably due to local glaciokarstic
processes.
2. The Lake K uzi water level changed most mark-
edly in the beginning of the Holocene. Between
11,200 and 10,900 cal. BP, the lake was shallow
and lake level fluctuated. Around 9,000 cal. BP
after possible melting of the covered dead ice
block, the lake basin achieved its current shape
and lake level started to rise. The lake reached its
maximum water depth of *11 m between 8,000
and 7,000 cal. BP and lake level became stable
until 5,000 cal. BP. After the lake level reached its
maximum stage, water depth started to decrease
because of a gradual infilling of the basin by the
sediment. Unlike from many lakes in Northern
and Eastern Europe, in which the water level fell
several meters between 8,000 and 4,000 cal. BP,
the Lake K uzis water level was stable until
2,000 cal. BP. Around 700 cal. BP, lake level
temporarily dropped and was 1.6 m lower than
present water level.
123
J Paleolimnol (2013) 49:663678
3. A warm and humid climate at the beginning of the
Holocene was followed by fluctuations between
cooler and warmer periods with varying precip-
itation patterns. Our study shows that the cooler
period between 8,500 cal. BP and 8,000 cal. BP
in the transition between the Early and the Middle
Holocene was followed by the period of the
Holocene Thermal Maximum that ended shortly
after 5,000 cal. BP. The effect of climate forcing
on lake development could be detected more
explicitly in the first half of the Holocene. Later,
the changes in the sediment properties seem to be
invoked by lake-catchment interactions and
anthropogenic factors.
4. Significant changes connected with the effect of
human activity (e.g., increases in erosion, eutro-
phication and lake level drop) in Lake K uzi
became clearly evident since 2,000 cal. BP.
Certain shifts in sedimentary signals (decrease
of Ti, increase of grain-size median, disappear-
ance of Chaoboridae) were detectable since
2,500 cal. BP. It is possible, that these sediment
changes represent the indirect evidence of the
onset of human activity in the catchment.
5. Comparison of several indicators in this study
show that respiratory horns of phantom midge
pupae (Chaoboridae) serve as indicators for high
water level and near-bottom anoxic conditions.
Acknowledgments This study was supported by the Estonian
target-financed project SF0280016s07 and the Estonian Science
Foundation under grant numbers 6857, 8189 and MTT3. We
hereby express our sincere thanks to our colleagues from the
Institute of Ecology for their participation in fieldwork and
discussions. We would also like to thank Prof. Andy Lotter,
Shinya Sugita, John F. Boyle and Prof. James B. Cotner for their
helpful comments and suggestions, and Prof. Bent Vad Odgaard
for helping to identify the respiratory horns of Chaoboridae.
Three anonymous reviewers are acknowledged for their helpful
comments and suggestions.
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