Transition from shallow lake to a wetland: a multi-proxy case study in Zalavári Pond, Lake Balaton, Hungary

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ISSN 0018-8158, Volume 641, Number 1

PRIMARY RESEARCH PAPER

Transition from shallow lake to a wetland: a multi-proxycase study in Zalavari Pond, Lake Balaton, Hungary

Janos Korponai • Mihaly Braun • Krisztina Buczko •

Istvan Gyulai • Laszlo Forro • Judit Nedli •

Istvan Papp

Received: 10 July 2009 / Revised: 18 December 2009 / Accepted: 30 December 2009 / Published online: 16 January 2010

� Springer Science+Business Media B.V. 2010

Abstract Lake Balaton, the largest shallow lake in

Central Europe, has no natural outlet, therefore,

underwent water level changes during its 15,000–

17,000 years of history. The lake is very sensitive to

both climate changes and human impacts. Surround-

ings have been inhabited since the Stone Age;

however, heavy human impact can be recognized

during the past 6000 years. In this study, we estab-

lished three different stages for and reconstructed

water level changes of Lake Balaton by geochemical

data, subfossil Cladocera and diatom remains in the

sediments of the Zalavari Pond, a part of the Kis-

Balaton wetland. In 9900–8600 cal. year BP, climate

was dry, water level was low, and there was a

wetland in this area. Although organic matter content

was low in the sediment, the ratio of Fe/Mn was high.

Between 5600 and 5000 cal. year BP, water level

increased, Fe/Mn ratio shows that oxygen conditions

of sediments was improved in agreement with the

relatively low number of diatom remains and dense

chydorid remains. About 5000 cal. year BP, water

level of Lake Balaton decreased as indicated by high

organic content with low carbonate and high Fe/Mn

ratio in the sediments (oxygen depletion). At the

bottom of this section, high Fe and S concentrations

showed accumulation of pyrite (FeS2) that is common

in wetlands with very low redox potential. Low

abundance of Cladocera remains together with rich

and diverse diatom flora confirm the low water level

hypothesis. Our data support that the water level of

Handling editor: J. Padisak

J. Korponai (&)

West-Transdanubian District Water Authority,

Csik F. str. 4, 8360 Keszthely, Hungary

e-mail: [email protected]

J. Korponai

Department of Chemistry and Environmental Sciences,

University of West Hungary, Szombathely, Hungary

M. Braun

Department of Inorganic Chemistry, University of

Debrecen, 4010 Debrecen, Hungary

K. Buczko

Department of Botany, Hungarian Natural History

Museum, 1087 Budapest, Hungary

I. Gyulai

Department of Applied Ecology, University of Debrecen,

4010 Debrecen, Hungary

L. Forro � J. Nedli

Department of Zoology, Hungarian Natural History

Museum, 1088 Budapest, Hungary

I. Papp

Department of Mineralogy and Geology, University of

Debrecen, 4010 Debrecen, Hungary

123

Hydrobiologia (2010) 641:225–244

DOI 10.1007/s10750-009-0087-0

Author's personal copy

Lake Balaton was higher between 8600 and 5000 cal.

year BP than it is at present.

Keywords Lake Balaton � Shallow lake �Cladocera remains � Chydorids � Diatoms �Trace metals

Introduction

Studying the paleoenvironment of small closed basins

has a long tradition all over the World. Lacustrine

sediments of deep lakes can preserve almost undis-

turbed evidences of changes in the past; therefore,

they provide ideal objects for paleolimnological

studies, including taxonomical, evolutional, and pa-

leoenvironmental reconstruction. Datasets delivered

from deep lakes are also used frequently for making

‘‘climate models’’. The so-called shallow lakes (Pad-

isak & Reynolds, 2003) have great economic and

social importance. Their development and historical

ecological status are in the focus of many studies.

Paleolimnological experts fully appreciate the fact

that the sedimentation of the extended, shallow water

bodies is far from ideal; however, their investigations

are urgent tasks of limnology.

Sediments of shallow lakes show greater mixing

than those in deep waters due to wind effect or

activity of benthic organisms (Kearns et al., 1996).

Shallow lakes and wetlands are more sensitive to

climate and human impact (Whitmore et al., 1996;

Kenney et al., 2002; Moss et al., 2003; Valiranta

et al., 2005) since small water level fluctuations in

absolute terms are considerable changes in relative

water depth. It is especially true for such a large and

shallow lake as Lake Balaton is. The estimated age of

the lake is cca. 15,000 years (Cserny & Nagy-Bodor,

2000) and from then, Lake Balaton was almost

endorheic while sandbars interfered with the lake

water to flush toward Danube.

Starting at the end of the nineteenth century, the

lake and its surroundings have intensively been

studied by a great number of experts (see the

comprehensive studies by Virag, 1998, 2005). Pan-

tocsek (in Loczy, 1916), Zolyomi (1952, 1995), and

Sebestyen (1969a, b, 1970, 1971) conducted first

paleolimnological studies in Lake Balaton sediments.

Zolyomi took sediment cores from several places in

lakebed for pollen analysis while Sebestyen analyzed

subfossil Cladocera remains in large intervals and

characterized the Cladocera fauna of Lake Balaton as

Alona dominant. Sagi (1968) and Bendefy (1968)

reconstructed former water levels by historic maps and

archeological remains while the Geological Institute of

Hungary has performed geological survey by several

proxies as palynology, siliceous algae, trace elements,

and stable-isotope (d13C, d 18O) analysis of carbonates

since 1989 (Cserny & Nagy-Bodor, 2000; Tullner &

Cserny, 2003). These studies revealed that Lake

Balaton is 15,000–17,000 years old, and its develop-

ment started with formation of several shallow, cold

water lakes. Between 5800 and 2500 cal. year. BP,

warming climate and increased precipitation caused

rise of water level as a consequence of which a uniform

lake was formed (Cserny & Nagy-Bodor, 2000). In the

late Holocene, the water level varied approximately

between 104.6 and 112.5 m aAsl (above Adriatic sea

level) as a function of changing climate (Virag, 1998;

Cserny & Nagy-Bodor, 2000).

The sediment of Lake Balaton is rich in carbonate

and its content varies between 50 and 70% depending

on the basins. Carbonate content increases from South-

West (Keszthely) to North-East (Siofok) (Cserny &

Nagy-Bodor, 2000; Tullner & Cserny, 2003).

When water level decreased, Lake Balaton lost

large amount of its surface and the shallowest basins

became wetlands, while when water table lifted up,

area of open water extended and lacustrine conditions

settled. Studying the series of wetlands and lacustrine

deposits in the shallowest area of lakes is a good tool

to reconstruct the past water level changes of large

lake systems such as Lake Balaton. In this study, we

aim to perform reconstruction of changes in water

level of Lake Balaton by geochemistry and subfossils

of Cladocera and diatom remains in the sediment of

the Zalavari Pond as a part of Kis-Balaton wetland.

Our proxies (trace elements, cladocera, and diatom

remains) have been widely used as proxy data to study

history of aquatic ecosystems. Biological proxies as

diatoms (Cameron et al., 1999; Battarbee et al., 2001;

Koster et al., 2004) and cladocerans (Korhola &

Rautio, 2001; Jeppesen et al., 2002) have been shown

to respond sensitively to changes in the past lake water

levels (Korhola & Rautio, 2001; Korhola et al., 2005),

water temperature (Bloom et al., 2003), and eutrophi-

cation (Siver, 1999; Wessels et al., 1999); thus,

subfossil assemblages have become widely used in

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Europe (Buczko et al., 2009) and North America (e.g.,

Harmsworth, 1968; Hofmann, 2001, 2003) for the

reconstruction of paleoclimate.

Materials and methods

Study site

Lake Balaton is the largest shallow lake in Central

Europe. Its catchment area is 5,175 km2, and the major

inflow is the Zala River that discharges about half of the

total Balaton watershed. It has an average discharge of

about 5–10 m3s-1. The only outflow is the artificial Sio

canal that was constructed in the 1860s and connects

the lake to the Danube River. Zala River enters Lake

Balaton through the Kis-Balaton Wetland (KBW), a

large area which acts as a filter for suspended material

and nutrients that would otherwise be deposited in the

lake (Tatrai et al., 2000). The KBW comprises two

parallel lowland basins running north to south, divided

by a low ridge of hills (Figs. 1, 2).

Formerly, Kis-Balaton was the fifth basin of Lake

Balaton until the beginning of the nineteenth century.

From then, it has been called Kis-Balaton. Its open

water area decreased to 8% by the end of eighteenth

century and to 4% by the end of nineteenth century

(Domotorfy et al., 2003). River Zala flushed over the

wetland at high flooding. From the middle of the

nineteenth century, intense canalization was carried

out to form large farmlands in the area of Kis-

Balaton. As farming was not economically efficient,

canalization was abandoned in the 1960s. This

canalization resulted in two major changes:

(1) Only two small water bodies Zalavari (Table 1)

and Vorsi Ponds escaped from desiccation and had

open water areas. (2) The Zala river flew directly into

Lake Balaton instead of filtering through an extended

delta. As a result of increased phosphorus load, the

originally mesotrophic lake underwent rapid eutrophi-

cation: Keszthely basin became hypertrophic during

the 1970s. In the 1980s, large-scale eutrophication

management measures were initiated involving con-

struction of a large reservoir to improve water quality.

The reservoir occupies the former Kis-Balaton delta

area and is, in fact, an extended wetland including open

water and swamp habitats. Water quality of the lake has

been improved due to wetland reconstruction and

investments of drainage and sewage treatment around

the lake (Istvanovics et al., 2007; Hajnal & Padisak,

2008; Tatrai et al., 2008).

Field and laboratory methods

Soil and sediment were sampled at 30 sampling sites

in the Kis-Balaton wetland. Three random samples

Fig. 1 Lake Balaton and its

possible expansion at

several altitude (min. 104.8

recent shore line; max.

113 m aAsl)

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were taken at each sampling point and the uppermost

15 cm layers were used for determination of loss-on-

ignition (LOI550) (Heiri et al., 2001) and carbonate

(Scheibler method) content.

Two parallel, 165 and 148 cm long sediment cores

were taken with a Livingstone corer (Fig. 2). First

core was used for geochemical analysis and diatoms,

the other for identification of Cladocera remains.

Cores were matched stratigraphically by lithography,

organic matter (LOI550), and carbonate (LOI950)

content. Cores were subsampled (1 cm3) at every

5 cm for chemical, diatom, and Cladocera analyses

using a volumetric subsampler.

For loss-on-ignition analysis, samples were weighted

and ignited at 550�C for 4 h (Heiri et al., 2001) and at

950�C for 2 h. Trace element content was determined by

ICP-OES (Thermo Iris Intrepid II XSP). Macrophyte

and charred plant remains were selected from the core

and three radiocarbon dates (Table 3) were determined

at the Laboratory of Environmental Studies, Institute of

Nuclear Research, Hungarian Academy of Sciences in

Debrecen. Dates were calibrated using the software

CALIB REV5.0.0 (Stuiver et al., 2005).

Subsamples (1 cm3) for Cladocera analysis were

treated with 250 ml 10% KOH solution and heated at

80�C for at least 1 h. Subsamples were gently stirred

by glass-rod to deflocculate all organic matter

without damaging the remains and sieved through a

65-lm mesh. Quantitative slides were prepared by

pipetting 100 ll of each subsample on a microscope

slide and were mounted with glycerol jelly stained with

safranine. All slides were then examined under a

microscope at magnifications 100 and 400 and about

200–250 Cladocera remains were counted from each

sample (Korhola & Rautio, 2001). All Cladocera

remains were encountered (carapaces, head-shields,

post abdomen, ephippia, and post-abdominal claws)

separately, but only the most frequent body parts for

each taxon were used for estimating the species

abundance. Poorly fragmented remains were counted

if they contained a clear diagnostic feature as proposed

by Korhola & Rautio (2001). Total concentration per

cm3 was calculated and identification was carried out

by using Frey (1950, 1962, 1988, 1991), Goulden &

Frey (1963), Gulyas & Forro (1999), Sebestyen

(1969b, 1970, 1971), Whiteside et al. (1978).

Fig. 2 The location of

Zalavari Pond and sampling

site. The solid line is the

boundary of the watershed

of Lake Balaton (basins of

the lake: 1 Keszthely, 2Szigliget; 3 Szemes, 4Siofok)

Table 1 Main features of Zalavari Pond

Latitude (N) 46.6669�Longitude (E) 17.2184�Elevation (m aAsl) 106.5 m

Max. depth (m) 1.2

Mean depth (m) 1.1

Surface area (m2) 45,000

Catchments area (km2) 2,125

Mean pH 7.81

Conductivity 622 lS

Total phosphorus 1.95 mg/l

Total nitrogen 0.18 mg/l

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For diatom analysis, 1 cm3 subsamples were

treated with hot hydrogen-peroxide, nitric acid and

the cleaned material was mounted in Zorax (refrac-

tive index is 1.7). At least 300 diatom valves were

counted at each level. For light microscope analysis,

LEICA DM LB2 was used (100 X HCX PLAN

APO). Diatom identification followed the nomencla-

ture of Krammer and Lange-Bertalot (1986–1991),

and the volumes of Diatoms of Europe (Krammer,

2000; Lange-Bertalot, 2001), as well as some other

manuals (Lange-Bertalot & Krammer, 1989; Lange-

Bertalot, 1993; Lange-Bertalot & Metzeltin, 1996;

Houk, 2003; Kobayasi et al., 2006).

Major and trace element concentrations were

assessed by multivariate methods [cluster analysis,

principal component analysis (PCA) with SPSS

(SPSS Inc., 1998)]. Further multivariate analyses

[detrended correspondence analysis (DCA) and

redundancy analysis (RDA)] were carried out with R

(R Development Core Team, 2008), vegan (Oksanen

et al., 2007), and BiodiversityR (Kindt & Coe, 2005)

packages. Stratigraphic zones were determined by

psimpoll (CONISS clustering, Bennet, 2005). Con-

centrations and species data were log-transformed and

standardized for PCA, DCA, and RDA, and PCA

scores were used in cluster analysis (Euclidean

distance with Ward method).

Significance of each environmental variable in

explaining the variance in taxon data was determined

by testing the significance of the first canonical axis

by means of a Monte Carlo permutation test (1,000

random permutations).

Results

Soil, lithology, ages, major and trace elements

stratigraphy

Carbonate content of soil of the lower part of Kis-

Balaton Wetland reflects sedimentation conditions in

Lake Balaton. It varies within a wide range (1–40%)

and is the highest in the former fifth basin of Lake

Balaton (Fig. 3).

Fine gravel and sand were found at the bottom of

the sequence between 150 and 165 cm. Organic

matter content was very low (1–2%), and carbonate

was \10% in this layer. Highest carbonate concen-

tration (ca. 30%) was measured between 70 and

145 cm. Concentration of organic matter showed

Fig. 3 Distribution of

CaCO3 content of soil and

sediment in Kis-Balaton

wetland (others: residue on

ignition)

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maxima (50–60%) and the lowest carbonate concen-

trations (6–7%) between 25 and 70 cm. The upper-

most sediment contained 20–30% organics and

20–25% carbonate, which was similar to the values

measured in surface soil samples in the area of the

fifth basin of Balaton. The core can be divided into 12

lithological units (Table 2) according to Troel-Smith

(1955). The high carbonate content and low organic

yield complicated the radiocarbon dating of the core.

No dates were obtained for the calcareous silts

Table 2 Lithological description of the core from Zalavari Pond

Depth (cm) Troel-Smith

categories

Description

0–5 Th3, Dh1 Dark, blackish-brown peat, containing macroscopically recognisable

plant remains. Organic content high

5–20 Th3, Lc1 Blackish-brown peat with high organic content. The amount of plant

remains is smaller, the amount of calcareous Mollusc shells is large

20–30 Th1, Lc1, Dg1, Dh1 Gray-black sediment, containing wood, Phragmites and Typha remains

30–55 Th2, Dl1, Dg1 Dark brown peat with wooden and herbaceous remains

55–70 Dl3, Dg1 Blackish sediment dominated by wood remains accompanied by

herbaceous remains

70–75 Dl1, Dg2, Ld1 Brownish peat containing charcoal grains

75–85 Lc2, Dg2 Calcareous gray silt with mollusc shells, containing visible plant remains

85–130 Lc3, Dg1 Calcareous gray silt containing less plant remains and numerous mollusc

shells

130–135 Dg1, Ld2, Lc1 Grayish-brown calcareous silt with mollusc shells

135–145 Dl1, Dg2, Ld1 Blackish peat with mollusc remains. Small amount of coarse sand and

small gravel appeared

145–150 Ga1, Ld1, Dl1, Blackish sediment with plant remains. The number of gravel significant

150–165 Ga1, Gg1, Ag2 Gray with black patches. Containing sand and gravel.

The number of 2–5 mm gravels large. Small amount of fibrous plant

remains

Table 3 Conventional,

calibrated radiocarbon ages

and d 13C for Zalavari Pond

Lab-code sample (cm) d 13C (PDB)

± 0.2 (%)

14C year BP Cal. year

BP (2r)

deb-13986 Zalavar 45–50 cm -27.82 1660 ± 35 1569 ± 69

deb-13996 Zalavar 65–70 cm -28.74 4135 ± 50 4680 ± 148

deb-13994 Zalavar 140–148 cm -28.24 7935 ± 60 8801 ± 184

Fig. 4 Age-depth model for Zalavari Pond

Table 4 Redundancy analysis of Cladocera remains and

major and trace elements partitioning of variance

Inertia Proportion

Total 14.145 1.0000

Constrained 7.792 0.5509

Unconstrained 6.535 0.4491

Axis RDA1 RDA2 RDA3 RDA4

Lambda 1.3123 1.1512 0.8356 0.7018

Variance explained % 20.6576 18.1211 13.1533 11.047

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Fig. 5 Major and trace

elements stratigraphy in

Zalavari Pond. The

lithology is described in

detail in Table 2

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between 70 and 140 cm. Thus, the chronology of this

sequence is not ideal. The results of the three

radiocarbon dates are presented in Table 3. Relation-

ship between age and depth is not linear (Fig. 4).

Highest sedimentation rate (0.3 mm year-1) was

found in the youngest part (0–47.5 cm) of the core,

whereas the lowest (0.1 mm year-1) was between

47.5 and 67.5 cm.

Three principal components and five stratigraphic

zones were distinguished by the multivariate analysis

(Fig. 5). The first three principal components were

explained 89.6% of total variance. The trace elements

of sediment of the Zalavari Pond could be classified

into three groups. Elements used as indicators of

erosion (Al, K, Mg, Cr, Co, and Ni) and human

impact (Pb and Zn) were loaded into the first

principal component (PCA1). The redox indicator,

Fe was also loaded into the first principal component.

Each of these elements showed positive correlations,

and their concentrations are higher in the ZVGC-2

(140–155 cm) and the ZVGC-4 and ZVGC-5 (0–

70 cm) zones than they were in the others. The

organic matter (LOI550), sulfur, sodium, and copper

loaded into the second group (PCA2), refers to

wetland development. Concentration of these com-

ponents is higher in the upper part of the sequence

(20–70 cm). The third group (PCA3) indicates

lacustrine environment with higher concentrations

of Ca, Mn, and carbonate content (LOI950).

ZVGC-1 (9900–9100 cal. year BP; 165–150 cm)

Concentration of elements indicating erosion (PCA1

group) is high. The sediment is poor in carbonates

and organic matter. Appearance of fine gravels and

coarse sand confirms higher level of erosion. Low Ca/

Mg ratio shows that water level might be lower with

poor vegetation.

ZVGC-2 zone (9100–8600 cal. year BP;

150–140 cm)

Color of the sediment in this zone is darker than

either above or below. Carbonate concentration

remained low and Ca/Mg ratio decreased. Concen-

trations of Fe, Mn, S, Cu, P, and organic matter have

a local maximum at 9000 cal. year BP. The increase

of Fe is higher than that of Mn, which causes a high

Fe/Mn ratio. Sulfur concentration increased from 1 to

13 g kg-1. These phenomena may indicate shallow

water and reductive conditions and increase in the

trophic state of the lake. Intense erosion is indicated

by gravel, coarse sand, and high concentration of

aluminum and other elements loaded in PCA1.

ZVGC-3 zone (8600–4800 cal. year BP; 140–70 cm)

Color is grayish with high concentration of calcium,

manganese, barium, and carbonate. Amount of Mg is

also significant. Ca and carbonate concentrations

gradually increase toward the middle of the zone, and

then they decrease indicating water level changes.

Concentration of Al and all others in PCA1 group is

very low. Water level was high in the middle of the

zone. Later, organic matter content increased upward

the sequences and wetland phase started to be

established.


Organic content varies between 50 and 60%. The

high peak of Fe/Mn ratio indicates that redox

conditions turned to anoxic again. The average

concentrations of Al, K, Pb, Zn, Ni, Co, Cr, S, Na,

Cu, and P are higher than in ZVGC-3 zone, while the

amount of Ca, Mn, Ba, and carbonate decreases.


The concentration of Fe, Al, Ni, Co, Cr, S, and Na is

similar to that of ZVGC-4. Organic content is lower,

while the concentration of Pb is slightly higher in this

zone. Ca, Mn, Ba, and carbonate content show a

strong increase. In the topmost 5 cm, the concentra-

tion of Ni, Co, and Cr slightly increases.

Cladocera

There were remains of 27 Cladocera species along the

core of Zalavari Pond. Species numbers varied

between 11 and 19 in the samples. These species are

very common in littoral habitats of eutrophic shallow

lakes with dense macrophytes, and in Lake Balaton as

well. Only ten species achieved dominant status

([10% in at least one subsample), of which nine were

chydorids, and only three long-term dominants were

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found throughout the profile: Alona rectangula, Alona

quadrangularis, and Chydorus sphaericus. By far the

most prominent species was Alona rectangula,

accounting for over 30% of all the Cladocera abun-

dances in the majority of the subsamples. Three zones

were distinguished on the basis of abundance of

Cladocera remains. Noteworthy, the frequency of

well-preserved remains was cc. ten-folds higher in

the lowermost than in the uppermost layer (Fig. 6).

ZVCL-1 (8600–5300 cal. year BP; 146–82 cm)

This zone is characterized by rich and abundant

Cladocera fauna (Fig. 6). The estimated total amount

of Cladocera varied between 10,000 and 40,000 of

individuals per cm3; the peak occurred at 124 cm.

Besides the dominant species Chydorus sphaericus,

Alona rectangula, A. quadrangularis, and Pleuroxus

uncinatus, planktonic Bosmina longirostris, Leydigia

Fig. 6 Cladocera stratigraphy for Zalavar Pond. The lithology is described in detail in Table 2

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leydigi, L. acanthocercoides, Alona affinis, Graptol-

eberis testudinaria, A. guttata, and Disparalona

rostrata were numerous.


Frequency of remains of dominant Cladocera species

decreased in numbers in this zone. Total abundance

of cladocerans decreased; it varied between 700 and

2500 ind. cm-3. The pelagic Bosmina longirostris

were found in very low numbers. Besides the

dominant species, Alonella exigua, Pleuroxus trigo-

nellus, Camptocercus rectirostris, Graptoleberis

testudinaria, Pleuroxus uncinatus, and Acropterus

harpae were found in higher numbers. These species

occur in large number in shallow and densely

vegetated habitats. Increased frequency of these

species and Camptocercus rectirostris reflects the

expansion of vegetated bed. Some Daphnia ephippia

and Simocephalus claws were only found in the

bottom of this zone.


Number of Cladocera remains was low (500–4200 ind.

cm-3) but slightly increased toward the uppermost

layers, and reached maximum in (4200 ind. cm-3)

the topmost subsample. Number of the pelagic species

such as Bosmina longirostris was higher than in the

previous zone. Abundance of Pleuroxus uncinatus

slightly increased in this zone. Kurzia latissima and

Sida crystallina were found in the lower part of the

core (Fig. 6). Besides that the species composition of

this zone is similar to the previous one indicating

wetland phase, increasing number of remains of

Bosmina longirostris shows increased trophic levels.

Diatoms

The diatom flora of the core from Zalavari Pond was

diverse and variable with more than 150 taxa. Some of

the most frequent species were present through the

sequence like Amphora ovalis, Aulacoseira granulata,

A. ambigua, Pseudostaurosira brevistriata, and Staur-

oneis construens varying in abundances. Amphora

pediculus and Cocconeis placentula were also con-

stant members of the flora and they were missing

only from one sample. Summarizing the abundance,

Pseudostaurosira brevistriata, Stauroneis construens,

S. construens var. venter, and Staurosira pinnata were

the most characteristic members of the diatom assem-

blages of Zalavari pond. All of them are members of

the sand diatom flora; they are common benthic taxa

(Fig. 7). Mainly alkalophilous or indifferent diatom

species were recorded along the sequence, indicating

that there was no significant change in pH.

ZVD-1 (9900–7700 cal. year BP; 165–126 cm)

Staurosira construens–Staurosirella pinnata zone

Low diatom diversity in every aspect was the main

feature of this zone. The number of species was low

(10–20), uniformity of forms and dimensions were

characteristic, and mainly small sized (\20 lm)

diatoms were present. Some dissolved fragments of

larger diatoms were also recorded. Staurosira

construens and Staurosirella pinnata dominated the

assemblages. The extremely low abundance of

planktonic species indicated the lack of deep water.

Wind-disturbed shallow water without macrophytes

can be reconstructed on the basis of the mud diatom

flora.

ZVD-2 (77000–1900 cal. year BP; 64–50 cm)

Aulacoseira ambigua–A. granulata zone Between

126 and 64 cm, there was poor diatom preservation.

The diatom flora of this zone is characterized by high

diversity (the number of taxa in the samples varied

between 46 and 72). Planktonic forms, such as

Aulacoseira ambigua, A. granulata, A. granulata

var. angustissima, and Stephanodiscus species were

the most frequent in this zone. The ratio of planktonic

forms was[50%, indicating the possible existence of

extended open water. Cymbella neocistula was found

only in this zone.

ZVD-3 (1900–1300 cal. year BP; 50–38 cm)

Navicula tenelloides–Fallacia tenera zone: Abundant

diatom flora was of great diversity (Shannon–Weaver

diversity was between 4.81 and 5.04) with a decrease

in Aulacoseira spp. and an increase in benthic taxa.

Apart from Aulacoseira species, the ratio of other

planktonic taxa remained relatively high indicating a

gradual decrease in the depth of water. The nominal

species of this zone (Navicula tenelloides) attained a

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definite peak exactly in the middle of the zone (44 cm).

In this sample, 20% of the total diatom floras belong to

this species. Its abundance gradually decreased above

and below this level. The presence of Fallacia tenera

was characteristic for this zone. In addition,

Achnanthes delicatula and Platessa conspicua

Fig. 7 Diatom stratigraphy in Zalavari Pond. The lithology is described in detail in Table 2

Hydrobiologia (2010) 641:225–244 235

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showed peaks in this zone further supporting the

shallowness of the lake.

ZVD-4 (1200–400 cal. year BP; 38–14 cm)

Staurosirella pinnata–Gyrosigma attenuatum zone: The

ratio of planktonic species further decreased and

benthic forms became dominant. Staurosirella

pinnata attained a peak at 32 cm. Amphora species

(Amphora ovalis and A. pediculus) were the most

frequent in this zone. Ulnaria ulna and Fragilariforma

virsecens also reached maximum abundances in this

zone. Diatom preservation was extremely poor

between 16 and 22 cm. This is indicated in Fig. 7 by

white shading. Some Gyrosigma fraction, broken

Epithemia valves, strongly silicified Aulacoseira

valves, and some small Fragilaria (sensu lato) were

found in these samples. These sporadic diatom remains

suggest that the flora and the composition of diatom

assemblages did not change; only the preservation was

unsatisfactory.

ZVD-5; (400–0 cal. year BP; 14–0 cm)

Pseudostaurosira brevistiata–Cocconeis placentula

zone: The diatom flora of this zone is characterized

by great taxonomic diversity and abundant

occurrence of benthic taxa (up to 90%). However,

the abundance of the planktonic forms gradually

decreased. Pseudostaurosira brevistiata was the most

frequent and abundant diatom in the upper part of the

zone. Staurosira construens and its varieties are also

frequent and abundant here. Increasing abundance of

the epiphytic Cocconeis placentula is detected in this

zone.

Redundancy analysis

As the ordination analyses applied to this study are

sensitive to rare species, only cladoceran taxa occurred

with Hill’s number [10. Of the initial 27 taxa, 18

fulfilled these criteria. Cladocera data yielded DCA

gradient lengths of 1.42372 and 1.04327 SD for axes 1

and 2, favoring the use of redundancy analysis (RDA).

Geochemical variables with high multicollinearity

[Variance Inflation Factor (VIFs [20)] were elimi-

nated one at time beginning with the variety having the

highest VIF, and the procedure was repeated until all

environmental VIFs were \20. Result of analysis on

the reduced datasets shows that 55% of total variance

of Cladoceran data were accounted by chemical data

and 45% remained unexplained. The eigenvalue of the

first two RDA axes, 5.38 and 0.91 captured 44% of the

cumulative variance in the Cladocera data, a slightly

larger proportion of variance than is explained by PCA

(38.77%) (Table 4). Monte Carlo permutation test

revealed that Na, Pb, and Fe contents were the most

explanatory environmental variables in the distribution

of Cladocera remains (Table 5). Since Fe has real

ecological value on the dispersion of cladoceran data,

as a redox indicator, and the PCA analysis revealed that

chemical variables were explained by PCA factors,

repeated RDA analysis were done on the cladoceran

dispersion with PCA scores of chemical data.

Result of the RDA is illustrated in Fig. 8 as a

Cladocera—chemical data biplot. On the biplot, PCA

factors are shown as arrows and the length of the

arrows indicates their relative importance in explain-

ing the species variation; whereas their orientation is

proportional to their approximate correlation to the

RDA axes. In this RDA, axis RDA1 is most clearly

related to those groups of elements that were found in

high concentration in wetland phase of Zalavari

Pond. This axis effectively separates the top sediment

layers. Cladocerans were found with low abundance

in these layers. RDA2 axis is related to those groups

of chemicals that contain carbonate, Ca, Mn, and P.

RDA2 axis also correlates with those groups of

elements that is explained by PC2 factor, of which

concentration is higher in the ZVGC-4 zone.

Table 5 Result of permutation test (1000 Monte Carlo per-

mutation) in constrained RDA of Cladocera data

Df Var F N. Perm Pr ([F)

Na 1 3.305 10.9259 100.000 0.01 **

Pb 1 1.214 4.0143 100.000 0.01 **

Zn 1 0.214 0.7080 100.000 0.72

Ba 1 0.644 2.1285 100.000 0.08 .

Fe 1 0.875 2.8940 100.000 0.04 *

LOI550 1 0.239 0.7904 100.000 0.60

P 1 0.495 1.6364 100.000 0.08 .

Cu 1 0.332 1.0961 100.000 0.30

Cr 1 0.473 1.5640 100.000 0.15

Residual 21 6.353

Signif. codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’ 0.1 ‘’ 1

236 Hydrobiologia (2010) 641:225–244

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Discussion

Due to its geographical location, Lake Balaton is

exposed to wind. Shallowness and large fetch of the

wind cause frequent sediment re-suspensions (Istva-

novics et al., 2004). Besides re-suspension, carbonate

cycle is also important in carbonaceous lakes (Smol,

2008). The Pannonian Sea rippled in the area of

Carpathian basin in the Pliocene epoch and filled with

fluvial and lacustrine sediments as well as Pleistocene

loess. Base rock and soil of the catchment area are

rich in carbonates, confirmed by high Ca/Mg ratio

(about 2.2) of the water of the Zala River (Korponai

unpubl.). Since catchment area of the Zala River is

about half that of Lake Balaton, majority of the total

Ca and Mg loads are transported to the lake by Zala

River.

Carbonate content of sediments depends on pH of

lake water. At low pH, carbonate dissolves in a water

column while at high pH it precipitates into the

sediment. Intense carbonate precipitation has caused

the decreasing ratio of Ca/Mg along the longitudinal

axis of Lake Balaton. Recent data show that Mg

content of the sediment in the Keszthely basin is lower

than that in the eastern and deeper basins, similar to the

lake water (Korponai unpubl.). Precipitation of bio-

genic carbonate varies annually depending on the

trophic state of lake water but interannual trends can be

recognized. Heinsalu et al. (2008) also found increas-

ing carbonate content as a sign of eutrophication in the

sediment of Lake Vorstjarv. When water level of the

lake was so high that it extended to the area of Zalavari

Pond, in other words this pond was the fifth basin of

Lake Balaton, CaCO3 content of the sediment was as

high in the pond as it is now in recent sediments of the

lake (Fig. 3) due to intense precipitation of carbonate

as a result of high eutrophic level. According to this

situation, we raised the conclusion that carbonate

content of sediment core and soil of Kis-Balaton

wetland indicates ancient open water of Zalavari Pond.

Since Mg concentration increases in a water body

with decreasing water level and forms Mg-rich

Fig. 8 Ordination diagram based on redundancy analysis

(RDA) (Samples given as black dots, PCA scores are arrowsand cladoceran species are black squares. Abbreviations:

Acro_harp Acroperus harpae, Alon_affi Alona affinis,

Alon_gutt Alona guttata, Alon_quad Alona quadrangularis,

Alon_rect Alona rectangula, Alon_exig Alonella exigua,

Camp_rect Camptocercus rectirostris, Chyd_spha Chydorus

sphaericus, Disp_rost Disparalona rostrata, Eury_lame Eury-

cercus lamellatus, Grap_test Graptoleberis testudinaria,

Leyd_acan Leydigia acanthocercoides, Leyd_leyd Leydigia

leydigii, Mono_disp Monospilus dispar, Pleu_adun Pleuroxus

aduncus, Pleu_trig Pleuroxus trigonellus, Pleu_unci Pleuroxus

uncinatus, Bosm_long Bosmina longirostris)

Hydrobiologia (2010) 641:225–244 237

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calcite in the sediments (Cserny & Nagy-Bodor,

2000), high Mg concentration of sediments reflects

low water level in the given basin.

Changes in the plankton:periphyton ratio of dia-

tom assemblages indicate water level fluctuation

(Heinsalu et al., 2008). Different preservations of

diatom frustules take a bias into reconstruction of

paleoenvironment (Battarbee et al., 2005). Poorly

preserved diatom frustules in this region should

reflect to shallow water also, since temperature, pH,

salinity, grazing or bioturbation, and water depth

affect diatom preservation (Flower, 1993). Smol &

Boucherle (1985) and Newberry & Schelske (1986)

found poor preservation of frustules in early Holo-

cene, and speculated that enhanced dissolution during

this period was linked to the higher lake water pH at

the time of carbonate deposition from the core of the

Little Round Lake, Canada. Diatoms were poorly

preserved in calcareous sediments from Lake Malawi

(Gasse et al., 1995), and from ephemeral lakes

(Flower, 1993). Barker et al. (1994) found that the

pH is the most significant factor affecting silica

dissolution, and the dissolution increases exponen-

tially when pH is above 9.

Water pH of Lake Balaton is high, commonly

exceeding 8.5 in summer and this condition is

coinciding with high carbonate content (50–55%).

Under such conditions, Cserny et al. (1991) found

poorly preserved diatoms in the sediment of Lake

Balaton.

In the sediment of Zalavari Pond, high Ca/Mg was

found in the zone ZVGC-3 suggesting the water level

was relatively high (Fig. 9).

Ratio of planktonic (e.g., bosminids and daphnids)

to littoral (e.g., chydorids) Cladocera species is widely

used for paleolimnological reconstruction of lake

Fig. 9 Ca/Mg and Fe/Mn

ratio in sediment of Zalavari

Pond and zones of studied

proxies. The lithology is

described in detail in

Table 2

238 Hydrobiologia (2010) 641:225–244

123


water level changes (Sarmaja-Korjonen & Alhonen,

1999; Korhola et al., 2000; Sarmaja-Korjonen, 2001).

Different chitinisation of exosceletons of microcrust-

acens, comparing bosminids and daphnids, and ecol-

ogy could biased in reconstructed community.

Hofmann (1998) warned that using planktonic/littoral

ratio was not suitable for all cases. He found that

Bosmina longirostris was present at shallow periods,

and B. coregoni occurred in deep layers in Belauer See.

High carbonate content in water body has strong

effects on cladoceran community. Re-suspension and

intense carbonate precipitation make Lake Balaton a

stressful habitat for cladocerans especially for daph-

nids (G.-Toth, 1992). As a result, daphnids occur in

low numbers among recent zooplankton communities

and in the sediment as well. In the recent plankton, B.

longirostris and B. coregoni, Diaphanosoma brachiu-

rum are the dominant cladoceran species. Bosminids

are considered as planktonic species, but they occur

both in open water and in the littoral planktonic

habitats (Korhola & Rautio, 2001). Thus, amount of

bosminid remains may reflect extension of open

water (Gasiorowski & Kupryjanowicz, 2009).

Although Bosmina longirostris indicates rather eutro-

phic conditions than water level changing (Szeroc-

zynska pers. comm), we can consider this species as

good indicator of water level changes in shallow and

eutrophic ecosystems. Whenever water table of the

lake was elevated during the lake’s history, large

areas were inundated in Kis-Balaton and Nagy-Berek

areas; thus littoral zone increased in the western and

southern parts of Lake Balaton (Fig. 1). Since B.

longirostris prefers shallow habitats, its abundances

will increase with extension of littoral zone. We

found B. longirostris remains along the core both in

wetland and lacustrine sediments, but their density

was the highest in the 146–82 cm deep core section;

therefore, high abundances of B. longirostris indi-

cated higher water level in shallow environment.

Monospilus dispar is most abundant in the deepest,

Siofok basin while Bosmina longirostris is most

abundant in the largest Szigliget and the Keszthely

basins of Lake Balaton (Ponyi, 1971, Gyulai,

unpubl.).

Besides carbonate content, we found that redox

potential indicates water level changes, precisely, the

transition from lake to wetland as well. In the core of

Zalavari Pond, deviating trends are observed in

parameters often considered as indicators of redox

changes (Fe, Mn, and Fe/Mn ratio, Figs. 5, 9). Fe and

Mn exhibit very low solubility in oxidative environ-

ment, but Mn is more soluble than Fe; thus it is

mobilized at first when oxygen content decreases

(Engstrom & Wright, 1984). Concurrently, low Ca/

Mg and high Fe/Mn ratios indicate low water level

and wetland environment (Fig. 9).

Sedimentation was similar to that observed in

F}onyed and Zalavar (Medzihradszky, 2005; Juhasz

et al., 2007) and in the Keszthely basin of Lake

Balaton (Cserny and Nagy-Bodor, 2000). Cores from

F}onyed and Zalavar were taken close to the shoreline

of Kis-Balaton wetland and authors did not recognize

any sign of hiatus in sediment profiles.

At 10000–8600 cal. year BP years, the climate

was cold and dry and the water level of Lake Balaton

was low (Cserny & Nagy-Bodor, 2000; Juhasz et al.,

2007). Decreased runoff of rivers and creeks results

in the decrease of ionic concentrations in the lake

water. We found high Mg and trace metal concen-

tration in this zone, and low Ca/Mg ratio. These

phenomena indicate that water level was low and the

high Fe/Mg ratio reflects low redox environment

(Koinig et al., 2003), which infer the pond turned to

wetland (Fig. 9). Vegetation was sparse in the ancient

pond as shown by low organic matter content in the

sediment. The absence of planktonic diatom species

indicated the lack of deep water. Cladocera commu-

nity was characterized by phytophylous species, but

occurrence of Monospilus dispar proved large bare

lake bottom; macrophytes could be localized at

shoreline.

Between 8500 and about 5000 cal. year BP zone,

climate became warmer and wetter thus the water

level of the lake elevated (Cserny & Nagy-Bodor,

2000; Juhasz et al., 2007). In this period, dense

planktonic and littoral community developed simi-

larly to the recent Keszthely basin. This environment

was well oxygenated by activity of algae and

macrophytes and biogenic carbonate precipitation

increased Ca/Mg ratio and decreased Fe/Mg ratio in

the sediment.

The absence of diatoms between 8000 and 4000 cal.

year BP coincides with the high carbonate content in

the sediment due to high water level. Zalavari pond

must have been as large and deep as the recent

Keszthely basin as indicated by the poorly preserved

diatom zones; dissolution of silica frustules was

substantial in sediment in relation to strongly alkaline

Hydrobiologia (2010) 641:225–244 239

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water with pH[ 9.0 or to the high carbonate content

(Barker et al., 1994; Reed, 1998). Water alkalinity also

depends on trophic level. Within eutrophic conditions,

pH can exceed 9 with high carbonate precipitation.

High amount of Cladocera remains and large number

of remains of Bosmina longirostris and the occurrence

of mud living Monospilus dispar also indicate deeper

water. Since chydorid fauna was dense and rich, we

could conclude that water level was high accomplished

with dense submerged vegetation along the shore in

large open water like the recent Keszthely basin is.

About 5000 cal. year BP, climate changed, con-

firmed intensive terrestrial erosion caused by human

deforestation (Sumegi et al., 2007). Human population

occupied the region in the pre-Neolithic, but the

agrarian population arrived around 7500 cal. year BP

as indicated by the relative increase of hazel pollen

(Juhasz et al., 2007). Strong human impact can be

observed from Middle Copper Age (Banffy, 2007)

when dominance of dock plantain and nettle pollen

indicated deforestation (Juhasz et al., 2007). Increasing

trace element concentrations from 5000 cal. year BP

coincide with intense deforestation while population

of Balaton-Lasinja culture had pastoral economy

(Banffy, 2007).

High organic matter content with low carbonate

and high Fe/Mn ratio (oxygen depletion) indicates the

shallowness and transition of the pond system to

wetland. At the bottom of ZVGC-4 section, high Fe

and S concentrations show bounding of pyrite (FeS2)

that is common in wetland with extremely low redox

environment. Pyrite can be found in the Holocene

sediment of Lake Balaton (Cserny et al., 1991).

In shallow environments, carbonate precipitation

is less intense due to decreased importance of

planktonic primary production. Among the cladoc-

erans, abundance of phytophylous species as Gra-

ptoleberis testudinaria, Camptocercus rectirostris,

Pleuroxus laevis increased. Water level in this period

had no effects on diatom communities, among which

the planktonic/tychoplanktonic diatoms were domi-

nant. We are able to solve the discrepancy between

diatoms and the other proxies if we assume the pelagic

habitat was so large among also extended littoral

(larger than in the historic eras) that pelagic diatoms

could develop but large emerged macrophyte i.e.,

reeds protected this basin from disturbance by wind.

Low carbonate content and cladoceran remains show

us the possible water level decrease. The gradually

decreasing but still relatively high water level per-

mitted the preservation of diatoms valves from

4000 cal. year BP.

Low concentration of Cladocera remains, rich and

divers diatom flora confirm the low water level

hypothesis. From about 1000 cal. year BP up to

recently, water level slightly increased as indicated

by higher carbonate content coinciding with lower

organic matter and slightly increasing Ca/Mg and

decreasing Fe/Mn ratios. We could reconstruct low

water level with dense macrophyte and high P

content since strong correlation exists between

submerged macrophyte and P accumulation (Rooney

et al., 2003), in ZVGC-2 and ZVGC-4 zones (Fig. 5).

Several studies proved uneven and mosaic-like

hydrological changes in the area of historical Lake

Balaton (e.g., Cserny & Nagy-Bodor, 2000; Jakab

et al., 2005). Jakab et al. (2005) found lacustrine

layers in 420–390 cm depth in the sediment of

Szigliget bay in the northern wetland of the lake, and

concluded that the water level was the highest around

5200 cal. year BP. It is consistent with the result of

Cserny & Nagy-Bodor (2000) who stated there was

humid climate, water level increased and modern lake

formed between 5800 and 2500 cal. year BP. Our

data, however, contradict these reconstructions since

lacustrine sediments were found in 140–70 cm depth;

it means that there was high water level between

8600 and 5000 cal. year BP.

This paleoenvironment of the pond is comparable

with the recent phase of Lake Balaton. In that time,

the pond was connected to the lake and it was

composed of a well-developed wetland with large

open water. In the Lake Balaton, waves induced by

weak wind (4 m s-1) are able to reach the bottom of

the lake and their energy dissipates in the surface of

the sediment disturbing it parallel. In that time, the

long wind fetch expanded into the Zalavari Pond. The

wind in the Lake Balaton results in almost permanent

mixing of the whole water column. Due to the

frequent re-suspension of highly calcareous (50–60%

carbonates) sediments and slow sedimentation of

precipitated carbonates, the water is always turbid

and phytoplankton and submerged vegetation become

light-limited. Although reduction of water level

increases light penetration and improve light climate

for submerged macrophytes at canopy level (Scheffer,

1998), vegetation remains weak at low water level of

the Lake Balaton due to the strong wind effect.

240 Hydrobiologia (2010) 641:225–244

123


According to Istvanovics et al. (2008), 12% of the

surface area is available for plant growth, and this

situation has been characteristic of Lake Balaton.

Virag (1998) concluded that the area of emergent

plants (mainly reed) has shown little variation

(11–18 km2, \3% of the surface area) from the end

of the eighteenth century, but the expansion was the

most striking in Keszthely Basin. Although the

formation of a macrophyte-dominated stable state

can be excluded in Lake Balaton irrespective of the

trophic conditions, patches that represent this state can

develop locally. Entz & Sebestyen (1946) demon-

strated that species richness of algae and invertebrate

communities of patches of Potamogeton perfoliatus

and Myriophyllum spicatum is much higher in all the

seasonal succession than the plant-free littoral. Bıro &

Gulyas (1974) found Alona affinis and Chydorus

sphaericus the most common taxa in Lake Balaton and

Alona rectangula, Disparalona rostrata, and Macro-

thrix laticornis were restricted to the reed-grass stand

of Keszthely basin. They also found Sida crystallina,

that occurred commonly in Lake Balaton. Reed-grass

vegetation of the Keszthely Bay at the inflow of Zala

River was characterized by the highest number of

species, both planktonic and littoral. In comparison,

the Cladocera community between 8600 and 5300 cal.

year BP in the Zalavari Pond and the recent fauna in

the Keszthely Basin of Lake Balaton we concluded

that they correspond to similar water level.

Water level has been recorded since 1803 and was

regularized since 1863. Datum level of the official

water gauge was established at 104.09 m aAsl. Since

water table is regulated between ?70 and ?110 cm

and the bottom of Keszthely basin is between 101.7

and 102.2 m aAsl, the maximum depth is cc. 3 m in

the Keszthely basin at the moderate water level.

Recently, the water table of Zalavari Pond stabilized

around 106.1 m aAsl. According to our measures, the

maximum depth of Zalavari Pond is 1.2 m; thus

bottom of Zalavari Pond is at higher altitude

(104.9 m aAsl). We found similar Cladocera com-

munity in Zalavari Pond between 8600 and 5300 BP

to recent Keszthely Basin. In this sense, water level

of this period in Zalavari Pond must be similar to the

recent level of Keszthely Basin, suggesting that it has

to be min. 3 m higher than the regulated level.

Result of new archeological excavations in the

southern shore of the Lake Balaton revealed the

archeological remains from Copper Age to Migration

period (7200–1500 cal. year BP) were found between

107.7 and 112.7 m aAsl in southern shore (Serlegi,

2007). This reconstructed water level changes are not

presented in the sediment records due to the short

duration of high water level stages. Our result

confirms that the water level of Lake Balaton was

higher between 8600 and 5000 cal. year BP than the

present.

Acknowledgments The authors thank their colleagues from

Laboratory of West-Transdanubian District Water Authority for

their help with data collection and collaboration. We would like

to express our special thanks for Kaarina Sarmaja-Korjonen for

her valuable comment. This study was financially supported by

Hungarian National Science Foundation, OTKA-T 049098 and

Hungarian National Research and Development Program

BALOKO 3B022/04.

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