CLADOCERA

Cladocera and geochemical evidence from sediment coresshow trophic changes in Polish dystrophic lakes

Izabela Zawiska • Edyta Zawisza •

Michał Woszczyk • Krystyna Szeroczynska •

Waldemar Spychalski • Alexander Correa-Metrio

Received: 24 April 2012 / Accepted: 23 February 2013 / Published online: 13 March 2013

� Springer Science+Business Media Dordrecht 2013

Abstract Change in the trophic state of lakes is a

topic of primary interest for limnologists and paleo-

limnologists, but also for governments in many

countries. These changes can be the result of the

natural evolution of lake ecosystems, but nowadays

are most often connected with human activity influ-

encing water bodies. In this article, we reconstruct

changes in the lake productivity and trophic state in

three dystrophic (humic) lakes located in Northern

Poland. Sediments from these lakes, which are part of

a national park, were submitted to Cladocera and

chemical composition analyses. Currently, the trophic

state of these lakes has been described based on the

water’s chemical composition, and they have been

classified as undisturbed ecosystems with a stable

trophic state. The main objective of this study was to

evaluate whether these lakes have been stable and

undisturbed ecosystems during the past centuries and

therefore whether they can be classified as natural and

pristine. The results of subfossil Cladocera analysis

and sedimentary geochemical analysis confirmed the

specific nature of studied lakes. However, our results

were surprising and showed that during the last

200 years two of the three lakes have undergone

distinct trophic changes, while one of them has barely

changed at all.

Keywords Dystrophic lakes � Cladocera �Eutrophication � Geochemical analysis �Wigierski

National Park

Electronic supplementary material The online version ofthis article (doi:10.1007/s10750-013-1482-0) containssupplementary material, which is available to authorized users.

Guest editors: Marina Manca & Piet Spaak / Cladocera:

Proceedings of the 9th International Symposium on Cladocera

I. Zawiska (&)

Institute of Geography and Spatial Organization, Polish

Academy of Sciences, Twarda 51/55, 00818 Warsaw,

Poland

e-mail: ikrajewska@twarda.pan.pl

E. Zawisza � K. Szeroczynska

Institute of Geological Sciences, Research Centre

in Warsaw, Polish Academy of Sciences,

Twarda 51/55, 00818 Warsaw, Poland

E. Zawisza

Instituto de Geofisica, Universidad Nacional Autonoma de

Mexico, Ciudad Universitaria, Mexico, DF, Mexico

M. Woszczyk

Department of Quaternary Geology and Palaeogeography,

Adam Mickiewicz University, Dziegielowa 27, 61680

Poznan, Poland

W. Spychalski

Department of Soil Science, University of Natural

Sciences, Szydłowska 50, 60656 Poznan, Poland

A. Correa-Metrio

Instituto de Geologıa, Universidad Nacional Autonoma de

Mexico, Ciudad Universitaria, 04510 Mexico, D.F.,

Mexico

123

Hydrobiologia (2013) 715:181–193

DOI 10.1007/s10750-013-1482-0

Introduction

Studies on sediments of lakes with a varying trophy are

of special importance when changes in the natural

environment are considered. Dystrophic lakes, also

called humic lakes, are of special interest as they are

sensitive ecosystems, very vulnerable to environmental

changes and characterized by unique faunal and floral

species composition (Gorniak et al., 2003). Dystrophic

lakes have unique features, the most noticeable of them

being the brown color of their waters, caused by the

inflow of humic substances (Gorniak et al., 2003, 2006;

Holopainen et al., 2003; Willen, 2003). As a result,

waters of dystrophic lakes contain a high content of

dissolved organic forms of carbon (DOC) in relation to

its dissolved inorganic forms (DIC) (Jones, 1992;

Gorniak et al., 2003). Thus, the Secchi disk visibility

in these kinds of lakes can reach values lower than 1 m,

with acidic water and very low conductivity. In order to

determine the level of the lakes’ dystrophy, the HDI

index (Hydrochemical Dystrophy Index) has been

proposed (see Gorniak, 1996), taking into account the

water conductivity, DOC:DIC ratio and water pH.

Lakes with a summer HDI higher than 50 are considered

dystrophic (Gorniak, 1996, 2004; Keskitalo & Eloranta,

1999). Dystrophic lakes are typical of Scandinavia,

Northern Canada and boreal Russia, but they are quite

rare in Middle European Lowlands. In Poland, they can

be found in the lake district located in the northern part

of the country, but they are very rare and are considered

boreal climate relicts. One of the biggest groups of these

lakes is located in northeast Poland in Wigierski

National Park (Fig. 1). All dystrophic lakes in this area

have been protected as part of a National Park since

1989 and have been part of a strictly protected area since

1985. Since 2000, they also have been protected by

European law in the scope of the Natura 2000 network

(Council Directive 92/43/EEC, 1992). Their protection

status describes them as natural and immutable

(unchanged) ecosystems on the basis of their chemical

water properties. However, up to now there have been

no studies on their natural history and development.

In this article, we present the results of a study on

the surface sediments of three of the lakes located in

Wigierski National Park. Our study shows the

subfossil Cladocera fauna and chemical composition

of sedimentary sequences derived from these lakes.

The main objective of the study was to evaluate

whether these lakes have been undisturbed ecosystems

and therefore natural and pristine. The results are

especially relevant in light of the conservation status

attained by the area. Our analyses were based on

subfossil Cladocera, which are known to be a useful

tool for reconstructing changes in lake productivity

and trophic state (Whiteside, 1970; Alhonen, 1972,

1985; Birks et al., 1976; Boucherle & Zullig, 1983;

Korhola 1990; Szeroczynska, 1991; Hofmann, 1996;

Chen et al., 2010). Analyses of the sediment’s

chemical composition, known to reflect productivity

changes in the lake (Boyle, 2001), provided further

support to our results. The lakes chosen for this study

were Suchar Wielki, Suchar IV and Suchar III, which

show different distophy levels as reflected in their HDI

values. The Cladocera fauna of these lakes has not

been studied in detail, although different studies have

noted the presence of 1 (Tunowski, 1992) to 14 species

(Karabin, 1999; Gorniak & Dobrzyn, 1999). Their fish

stock has not been well studied either, but from

unpublished information, it is known that there are

three, two and one fish species in Suchar Wielki,

Suchar IV and Suchar III, respectively. Up to now,

nothing was known about their subfossil Cladocera

fauna composition. Given the differences among lakes

in terms of their chemical and physical properties, size

and depth, it is of special interest to investigate

whether these differences are also reflected in the

composition and evolution of their subfossil Cladocera

fauna.

Fig. 1 Localization map. Wigry National Park is marked by an

asterisk

182 Hydrobiologia (2013) 715:181–193

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Materials and methods

The lakes selected for the purposes of this study are

located in northeast Poland in Wigierski National

Park. The entire catchment of the studied lakes is

occupied by coniferous forest, and the majority of the

area of is covered by Vaccinio uliginosi-Pinetum and

Sphagno girgensohnii-Piceetum. The near-shore parts

of the lakes are overgrown by floating vegetation mats.

With a mean annual temperature of 6.4�C, this area

represents the coldest part of Poland.

Around 20 lakes located in this area are the only

ones known, being a group of dystrophic lakes in

Poland (Gorniak, 2004). The national park area

includes lakes with HDI values of more than 120,

but also lakes that hardly exceed an HDI of 50, which

is considered the threshold value for dystrophy. The

study was conducted on three lakes with widely

varying HDI values: Lakes Suchar III (SIII), Suchar

IV (SIV), and Suchar Wielki (SW). The highest HDIs

among all lakes in this region ([100) are found in two

of the studied lakes (Suchar III and Suchar IV), and the

lowest HDI value slightly exceeds 50 at Lake Suchar

Wielki. Chemical and physical water parameters of

the studied lakes were described by Gorniak (2006).

Sediment cores of these three lakes were recovered

using a KC-Denmark Kajak-type gravity corer in

summer 2010. The material was described in the field

and cut into 1-cm-thick sections. Following standard

procedures, samples for subfossil Cladocera analysis,

geochemical analysis and 210Pb dating were taken

from each section and stored in a refrigerator until

analysis (Frey, 1986).

Subfossil Cladocera analysis

Material for Cladocera analysis was prepared accord-

ing to standard procedures (Frey, 1986). Samples

(1 cm3 of fresh sediment each) were treated with hot

10% KOH for 20 min using a magnetic stirrer in order

to deflocculate the material, sieved through 33-lm

mesh and diluted in 10 cm3 distilled water. Slides were

prepared using 0.1 ml of each sample and examined

under a microscope (100, 200 and 400 magnifications).

Two to six slides from each sample were scanned.

Cladoceran remains were counted (head shields, shells,

ephippia, postabdomens). Identification of cladoceran

remains was based on keys by Flossner (2000) and

Szeroczynska & Sarmaja-Korjonen (2007). Results

were plotted in an absolute abundance diagram by

means of C2 software (Juggins, 2005, 2007).

Dating methods

Sediments were dated using 210Pb analysis. An amount

of 3 cm3 of fresh homogenized material was taken

from each section. The 210Pb activity of sediments was

determined indirectly by alpha spectrometry measure-

ment of 210Po (Ea = 5.31 MeV, T1/2 = 138 days)

activity (Flynn, 1968). The analysis was conducted at

the Laboratory of the Institute of Geological Sciences,

Polish Academy of Sciences, Warsaw. The activity of210Pb and 208Pb was measured by means of an

OCTETE PC alpha spectrometer produced by

EG&G ORTEC. The constant rate of the supply

(CRS) model was used to calculate sediment age

(Appleby, 2001). Age-depth models were derived for

sediments of Lakes Suchar III and Suchar Wielki. In

Lake Suchar III, 200, 100 and 50 years old corre-

sponded to depths of 30, 18 and 12 cm, respectively

(Supplementary Material 1). In Lake Suchar Wielki,

200, 100 and 50 years old were derived from 40, 23

and 14 cm of depth, respectively (Supplementary

Material 2). Activity measured for the sediments of

Lake Suchar IV did not allow the development of an

age-depth model (Supplementary Material 3).

Geochemical analyses

Several geochemical analyses of the sediments were

carried out for the purpose of the study: SiO2ter,

SiO2biog, TOC, TN, TS, P, Fe, Mn, K, Cu, Zn and Al.

The analyses of the chemical composition of sediment

deposits provide an alternative proxy of biological

productivity in the studied lakes (TOC, N, SiO2biog)

and allow linking the Cladocera-based reconstructed

changes with natural and anthropogenic processes in

the catchment area. The former include physical and

chemical denudation, indicated by the contents of

SiO2ter, Al and alkali/alkaline earth metals, and the

latter are reflected in the contents of heavy metals.

Samples at 1-cm resolution were lyophilized and

homogenized in an Fritsch agate grinder, model

Pulverizette 2. Contents of terrigenous silica (SiO2ter)

and biogenic silica (SiO2biog) were determined in

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organic matter-free material. Organic matter was

removed by combustion at 550�C for 4 h (Heiri

et al., 2001). Acid- and base-soluble fractions of the

sediments were dissolved in HCl (12 M, room tem-

perature/12 h and t = 100�C/2 h/water bath), and

subsequently in 0.5n NaOH (t = 100�C/2 h/water

bath) (Woszczyk et al., 2011). After each stage of

the extraction, residues were transferred to a paper

filter, flushed with distilled water, combusted and

weighed. The residue after acid treatment was mainly

composed of total silica (SiO2tot = SiO2ter ? SiO2-

biog), and after alkali digestion of only SiO2ter. The

difference between SiO2tot and SiO2ter was represented

by the content of SiO2biog. The analyses of total carbon

(TOC), total nitrogen (TN) and total sulfur (TS) were

carried out with the application of the elemental

analyzer VarioMax CNS (Elementar). Due to the non-

carbonate character of the sediments (evidenced by

carbon determinations in 15 randomly selected and

acidified samples), TC was presumed equivalent to

total organic carbon (TOC). The solutions for the

analyses of P, Fe, Mn, K, Cu and Zn were prepared by

dissolving 0.5–1.0 g of sediment and pre-combusted

at 550�C for 4 h in 12 M HCl. The phosphorus content

was determined colorimetrically by means of ammo-

nium molybdate. Contents of Al, Fe, Mn, K, Cu and Zn

were determined by the FAAS method using a

spectrometer SpectrAA 220 (Varian). N2O was used

to enhance atomization of Al. The quality of the

measurements was controlled by certified reference

materials (Sulfadiazine, Metals in sewage sludge

SQC01S, and Nutrients in Soil SQC014S).

Statistical methods

Cladocera zones were established through stratigraph-

ically constrained clusters (CONISS), which consist of

grouping based on Euclidean distances among samples

of a same core, conditioned to each sample, preserving

its position within the sedimentary sequence (Grimm,

1987). Two independent detrended correspondence

analyses (DCA) (Hill & Gauch, 1980) were carried out

using species and geochemical data, respectively. In

both cases DCA was performed pooling together the

data from the three lakes. Consequently, samples were

ordinated in two bidimensional coordinated spaces

(biplots), one defined by the biological data (Clado-

cera) and the other defined the abitoic proxies. DCA

was preferred over other ordination techniques

because it avoids arch effects derived from linearity

assumptions, and the units of the ordination (standard

deviations) are directly interpretable as ecological

turnover (Gauch, 1982). Axis 1 and 2 scores of species

DCA were plotted stratigraphically along depth to

easily identify points of inflection in terms of ecolog-

ical change. Additionally, Euclidean distance from

each sample to the modern sample was calculated

using the first four axes of the ordination, with

resulting units being standard deviations of ecological

turnover (Gauch, 1982). As the space defined by DCA

axes represents the ecological and environmental

envelope defined by the changes revealed by the

samples analyzed, distance analysis offers insights

into the ecological similarity between modern condi-

tions and each analyzed time slice of the past (Restrepo

et al., 2012).

Results

Subfossil Cladocera analysis

Remains of 27 Cladocera species belonging to five

families (Daphniidae, Bosminidae, Leptodoridae,

Chydoridae and Sididae) were found in the sediments

of the three lakes. Results of the analysis are shown in

absolute frequency diagrams (Figs. 4, 5, 6), allowing

the identification of Cladocera phases (CLAD), which

in turn reflect lake-development stages.

Lake Suchar III

Remains of 14 Cladocera species were found in the

sediments of Lake Suchar III, with only two of them

being pelagic: Bosmina (E.) coregoni and Bosmina

(E.) longispina. Littoral Cladocera, particularly Alona

affinis, Acroperus harpae and Alonella excisa, were

dominant through the sedimentary sequence. Simi-

larly, Alonella exigua and Graptoleberis testudinaria

occurred continuously. Contrastingly, species living in

association with plants were not very abundant.

Species inhabiting sediments were almost absent with

remains of Disparalona rostrata being found in one

sample (Fig. 2). Three cladocera phases were identi-

fied. In phase CLAD I (48–24 cm, sediments older

184 Hydrobiologia (2013) 715:181–193

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than 150 years ago), frequencies of Cladocera remains

were low, whereas littoral Cladocera predominated,

with Alonella excisa as the most abundant species.

Additionally, the lower part of this phase was char-

acterized by the continuous presence of pelagic

Cladocera Bosmina (E.) coregoni. CLAD II

(24–14 cm, 150–70 years ago) was distinguished by

an increase in total Cladocera density to 8,000

individuals in 1 cm3. The dominant species were

Acroperus harpae, Alonella excisa and Alonella

exigua living in association with plants. The rare

species Kurzia lattisima appeared again and was

continuously present. In CLAD III (14–1 cm, 70 years

ago to present), the total frequency of almost all

species decreased, except for Bosmina longirostris,

which was more numerous than in previous phases.

Lake Suchar IV

Remains of 23 Cladocera species were recorded in the

sediments of Lake Suchar IV, including pelagic

Bosmina (E.) coregoni, Bosmina (E.) longispina,

Daphnia longispina group and Ceriodaphnia spp.

Through the sedimentary sequence, littoral species

were dominant, particularly Acroperus harpae, Alo-

nella nana, Alonella excisa and Camptocercus recti-

rostris. In this lake, Cladocera living in association

with plants were numerous. Besides Camptocercus

rectirostris, this group was dominated by Alonella

exigua, Graptoleberis testudinaria and Eurycercus

lamellatus. Cladocera living in the sediment were

represented by only one species, Alona quadrangu-

laris, which reached very low frequencies (Fig. 3).

Three cladocera phases were identified. Phase CLAD I

(69–36 cm) was distinguished by high frequencies of

Alonella excisa, Alonella nana and Acroperus harpae.

Species living in association with plants were very

numerous and dominated by Graptoleberis testudina-

ria and Alonella exigua. Frequency of those two

species slowly increased during the time, whereas the

frequency of the other littoral species Camptocercus

rectirostris decreased. The pelagic Cladocera Bos-

mina (E.) coregoni reached the highest frequency in

the core. In phase CLAD II (36–20 cm), the high

frequency of Alonella excisa, Alonella nana and

Acroperus harpae, dominant in the previous phase,

Fig. 2 Absolute frequency of Cladocera remains in the sediment core from Lake Suchar III (axis Y indicates sediment depth cm)

Hydrobiologia (2013) 715:181–193 185

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was maintained. The abundance of species living in

association with plants slowly decreased. The fre-

quency of pelagic species Bosmina (E.) coregoni

decreased, whereas other open-water Cladocera

appeared, namely Bosmina (E.) longispina and Cer-

iodaphnia spp. The frequency of Bosmina longirostris

slowly increased in this phase. In phase CLAD III

(20–1 cm), the total frequency of all littoral species

decreased, whereas the abundance of Cladocera living

among plants was the lowest. The frequency of

Bosmina longirostris increased significantly, and the

species became predominant. Cladocera living in the

open-water zone, namely Bosmina (E.) longispina and

Ceriodaphnia spp., were also the most numerous in

the core.

Lake Suchar Wielki

Remains of 19 species were found in the sediments of

Lake Suchar Wielki, with only two of them being

pelagic: Bosmina (E.) coregoni and Bosmina (E.)

longispina. Littoral Cladocera were dominant, includ-

ing Alona affinis and Alonella excisa, whereas species

living in association with plants were not very

numerous. Two plant-associated species were contin-

uously present: Camptocercus rectirostris and Eury-

cercus lamellatus. Cladocera living in the sediment

were represented by only one species, namely Alona

quadrangularis, which was present in almost all

samples, but at low frequencies (Fig. 4). Two clado-

cera phases were identified. The dominant species in

phase CLAD I (69–22 cm, older than 100 years ago)

were Alona affinis and Alonella excisa. Among species

living in association with plants, Camptocercus rec-

tirostris and Eurycercus lamellatus were continuously

present. Remains of Alona quadrangularis living in

the sediment were found. Pelagic species were also

quite abundant. In phase CLAD II (22–1 cm,

100 years ago to present), the frequency of Alonella

nana increased substantially, and it became the

dominant species, while the number of Alona affinis

and Alonella excisa decreased. Pelagic species Bos-

mina (E.) coregoni and Bosmina (E.) longispina

became more numerous, with the increase being more

notable for Bosmina longirostris.

The composition of subfossil Cladocera fauna of

the three lakes under study showed a number of

similarities. In all lakes, dominant species were living

in the littoral (Acroperus harpae, Alonella nana,

Alonella excisa and Alona affinis). Species living in

association with plants, such as Alonella exigua,

Camptocercus rectirosris and Graptoleberis testudi-

naria, were quite numerous. The very significant

feature of these lakes was the almost entire lack of

Fig. 3 Absolute frequency of Cladocera remains in the sediment core from Lake Suchar IV (axis Y indicates sediment depth cm)

186 Hydrobiologia (2013) 715:181–193

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species living in the sediment. Whereas Alona quad-

rangularis reached very low frequencies in Lakes

Suchar IV and Suchar Wielki, Disparalona rostrata

was recorded in only in one sample of Lake Suchar III.

In general, within the last 200 years in the studied

lakes, littoral species were dominant, and pelagic

species, mostly represented by Bosminidae, were the

minority.

Results of geochemical analysis

The analyzed cores exhibited similar vertical changes

in chemical composition (Fig. 5). Sediments were rich

in TOC, but its contents distinctly decreased in the

upper 25–30-cm-thick layer in favor of SiO2ter and

SiO2biog. The contents of terrigenous and biogenic

silica were rather low (up to 22 and 6.5%, respec-

tively). The carbonate content was very low, and on

the basis of Ca as well as single measurements of TIC,

was estimated as\3% except at the upper part of the

Suchar IV profile.

In the upper part of the studied cores, the sediments

displayed enrichment in sulfur, metals and phospho-

rus. The Fe/Mn ratio was higher (between 35 and 45)

in the upper layer than in underlying sediments. The

sediments of Lake Suchar Wielki showed a relatively

high and vertically invariant ratio of TOC/N. Along

the Lake Suchar III core, molar TOC/N steadily, albeit

slightly, increases; however, its values were lower

than in the Lake Suchar Wielki site. In Lake Suchar

IV, the proportion of Fe/Mn displayed vertical fluc-

tuations between 14 and 12 in the middle and upper

sections, respectively.

Results of statistical analysis

DCA analysis based on Cladocera species and chem-

ical components of the sediment (Fig. 6 and Table 1)

showed a trend of samples from within each lake to be

ordinated in clusters when plotted in the space defined

by axes 1 and 2. Axis 1 clearly set apart the three

studied lakes (Fig. 6b, d), which shows that there is

more consistency in Cladocera fauna composition

within each lake in time than among lakes. DCA

shows that the environmental gradient associated with

axis 1 was the most important element differentiating

these lakes during last 300 years (Fig. 7a). However,

it was not possible to straightforwardly interpret the

environmental variable explaining axis 1, although we

can clearly state that these lakes have been different

Fig. 4 Absolute frequency of Cladocera remains in the sediment core from Lake Suchar Wielki (axis Y indicates sediment depth cm)

Hydrobiologia (2013) 715:181–193 187

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throughout time. DCA axis 2 also strongly separated

the studied lakes (Figs. 6b and d, 7b). This axis seems

to be associated with changes in lake productivity.

Cladocera that were ordinated in the most positive end

of axis 2 are mostly open-water species (Holopedium

gibberum, Bosmina longispina, Daphnia longispina

group) and Bosmina longirostris, species living in the

pelagic zone, but which also appear in the littoral zone.

Most of them (specially Bosmina longirostris) are

indicative of a nutrient presence in open-water zones

(Davidson et al., 2011). The youngest samples of

Lakes Suchar Wielki and Suchar IV scored the highest

DCA axis 2 (Fig. 7b). DCA based on chemicals

(Fig. 6c) supports the idea that axis 2 represents

productivity. Positively associated with axis 2 are

SiO2biog and phosphorus and negatively Al and K.

Maximum distances to modern samples were around

0.5, 1.0 and 1.5 SD for Suchar III, Suchar IV and

Fig. 5 Chemical composition of the sediments of Lakes Suchar Wielki, Suchar III and Suchar IV. Asterisk indicates concentrations of

TIC in 18 samples. Shaded areas indicate missing geochemical data

188 Hydrobiologia (2013) 715:181–193

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Suchar Wielki, respectively, and all of them occurred

at the bottom of the sequences (Fig. 7c), implying that

at this point species composition was most dissimilar.

Distances to the modern sample also showed a pattern

toward lower values in time for Lakes Suchar Wielki

and Suchar IV, while Lake Suchar III showed a

U-shaped pattern with minimum values around the

mid-depths of the sedimentary sequence.

Discussion and conclusions

Results of subfossil Cladocera and geochemical

analyses of the sediments of lakes Suchar III, IV and

Wielki reflect the very specific nature of dystrophic

lakes. The number of Cladocera species found varied

between 14 and 23, which is relatively low compared

to typical Polish lowland lakes where more than 30

species are usually found in sediment cores. All

studied lakes were dominated by Acroperus harpae,

Alonella nana, Alona affinis and Alonella excisa,

species known for their resistance to unfavorable

environmental conditions, especially low nutrients

and low pH (Fryer, 1968, 1991; Whiteside, 1970;

Rautio, 1998; Flossner, 2000; Bjerring et al., 2009).

Additionally, these species have low calcium (Ca?)

demands (Shapiera et al., 2011). The most dominant

species in the studied lakes were those living among

plants, which seems to be characteristic of dystrophic

lakes. In the studied sediments, there were almost no

Cladocera species living in association with the

sediment, suggesting a lack of a typical littoral zone

of dystrophic lakes, which has probably been replaced

by floating sphagnum.

The specific nature of these lakes was also

confirmed by the results of sedimentary geochemi-

cal analyses. Very high contents of TOC indicate

enhanced production of organic compounds in the

water column and/or delivery of organic matter from

terrestrial sources. TOC/N values between 12 and 16

show that both mechanisms played a significant role in

the environmental dynamics of these dystrophic lakes

(Meyers & Teranes, 2001). Low abundance of Ca can

be regarded as an indication of a very low content of

carbonates in the sediments, which is in turn associ-

ated with acidic pH (4.5–5.8). In low pH, environ-

mental calcite hardly precipitates, and sedimentary

calcium preferentially occurs associated with organic

matter. Low contents of SiO2biog were probably the

result of a low frequency of diatoms in the phyto-

plankton of dystrophic lakes (Woszczyk, 2011).

The main aim of this study was to evaluate whether

dystrophic lakes protected by several laws are undis-

turbed ecosystems and therefore natural and pristine, at

least during the last 200 years. We expected the two

most dystrophic lakes (Suchar III and Suchar IV) to be

undisturbed, contrary to the less dystrophic Lake

Suchar Wielki. However, our analyses show that lake

environment in all three studied lakes was not stable

during the last 200 years. Species composition of the

oldest phases in all lakes (Clad I) confirms that these

lakes were dystrophic and their shores were thickly

overgrown by plant mats. Cladocera results and DCA-

based distances to modern samples show that there

were important changes in the aquatic ecosystem

around 100 and 70 years ago in Suchar Wielki and

Suchar IV, and in Suchar III, respectively (Figs. 2, 3, 4,

6b). Our results suggest that ecological turnover of

species was closely associated with increasing pro-

ductivity through time (Figs. 6b, 7b, c). The increase of

productivity is also reflected in the chemical compo-

sition of the sediment. The most dystrophic lake

(Suchar III) was the most stable in time in terms of

subfossil Cladocera fauna composition (Fig. 7c),

implying that the productivity of that lake changed

very little during the studied time period. Contrasting-

ly, the sediments of the second most dystrophic lake in

the region, Suchar IV, revealed important changes in

the Cladocera community. The most visible ecological

turnover happened in the Cladocera community of

Suchar Wielki, where, as suggested by the DCA-based

distance between fossil and modern samples, over 50%

of subfossil Cladocera composition changed in terms

of species composition. Noted changes in both lakes

(Suchar IV and Suchar Wielki) are probably evidence

of simultaneously increased productivity (Fig. 7b).

The reason for increased productivity in the studied

lakes, especially in Suchar IV and Suchar Wileki,

seems to be edged uploading of nutrients caused be the

erosion from the catchment and changes in land use of

the region. An increased share of terrestrial compo-

nents in the sediments is well documented in the results

of geochemical analysis of the sediment. In the upper

part of the cores, a distinctive drop of TOC occurred

accompanied by enrichment in terrigenous silica. This

provides evidence for the enhancement of erosion in

the lakes’ surroundings in the recent past 100 years in

Suchar Wielki and 70 years in Suchar III. Intensified

Hydrobiologia (2013) 715:181–193 189

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catchment erosion has often been attributed to defor-

estation and produces an import of both suspended

matter and nutrients to lakes (Cundy & Croudace,

1995; Eusterhues et al., 2005). Increased loading of

nutrients, shown by the enrichment in P in the top part

of the cores, resulted in a change to less dystrophic/

Fig. 6 DCA analysis performed for the sediments of Lake Suchar

III, Suchar IV and Suchar Wielki based on Cladocera species (a,

b) and chemical components of the sediment (c, d). Explanation of

the Cladocera species acronyms used in the DCA diagram: Aaffi—

Alona affinis, Acost—Alona costata, Agutt—Alona guttata, Ain-

ter—Alona intermedia, Arect—Alona rectangula, Aquar—Alona

quadrangularis, Aexcisa—Alonella excisa, Aexigua—Alonella

exigua, Anana—Alonella nana, Acharp—Acroperus harpae, Al-

elon—Alonopsis elongta, Bcore—Bosmina (E.) coregoni, Blgir—

Bosmina longirostris, Blgis—Bosmina (E.) longispina, Chsphae—

Chydorus sphaericus, Crectir—Camptocercus rectirostris, Dlon-

gis—Daphnia longispina group, Elamell—Eurycercus lamellatus,

Gtestu—Graptoleberis testudianria, Kurzia—Kurzia lattissima,

Ptrigo—Pleuroxus trigonellus, Scryst—Sida crystallina

190 Hydrobiologia (2013) 715:181–193

123

more eutrophic conditions in which diatoms could

develop (Lepisto & Rosenstrom, 1998). Increasing

importance of diatoms in the phytoplankton is suggested

by the enrichment in SiO2biog (Boyle, 2001). On the

other hand, a slight increase in lake water pH due to

eutrophication provides a plausible explanation for the

increase in Ca, specifically in Lake Suchar IV. The more

eutrophic condition in the lakes during the last 100 years

is also reflected in the Fe/Mn ratio, which is used to

reconstruct past red-ox conditions in the lakes (Boyle,

2001). The detrital origin of both metals in the lake

sediments was concluded based on the highly positive

correlations. An upward increase in Fe/Mn, combined

with the enrichment in sulfur, points to a slight

worsening of the oxygenation of the bottom waters in

the lakes.

In the most dystrophic lake, Suchar III, the first

changes in Cladocera species composition started ca.

Table 1 Detrended correspondence analysis (DCA) results for

cladocera and chemical analyses

Proxy DCA axis

1

DCA axis

2

DCA axis

3

DCA axis

4

Cladocera

Eigenvalue 0.216 0.133 0.093 0.063

Axis length 1.954 1.608 1.338 1.285

Geochemistry

Eigenvalue 0.028 0.006 0.002 0.005

Axis length 0.664 0.274 0.187

Fig. 7 DCA and derived analyses on Cladocera from lakes

Suchar Wielki, Suchar III and Suchar IV. a, b DCA axis 1 and 2

scores plotted in depth, respectively. c Euclidean distance to the

modern sample calculated using the first four DCA axes as an

indicator of past ecological divergence from modern conditions

Hydrobiologia (2013) 715:181–193 191

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150 years ago. At that time, planktonic species almost

disappeared, and there was a substantial increase of

species living in association with plants (phase Clad II,

Fig. 2). The reason for this change was probably

lowering of the water level. This change was probably

triggered by the building of a melioration channel in

the close vicinity of lake. The increased frequency of

most littoral species reflects the widening of the littoral

zone by channeled water by a drainage canal. Later on,

as the canal became overgrown by plants, this caused

the frequency of littoral species to slowly decrease and

be replaced by planktonic species. In the youngest

sediment, ca. 70 f years ago, a slight increase in the

trophic status of Lake Suchar III was apparent (phase

Clad III, Figs. 2, 7b). As in Suchar Wielki and Suchar

IV, this change could be indicative of land use changes

in the lake catchment. However, it should be empha-

sized that, as evidenced by DCA, changes in the

trophic level in Lake Suchar III are much slighter than

those in Lakes Suchar IV and Wielki.

In summary, the results of subfossil Cladocera and

geochemical analyses of the sediments of these three

dystrophic lakes were surprising. It turned out that

they are not as pristine and stable as had been

considered and that there are important differences

in their trophic states despite their very similar HDI

values (Lakes Suchar III and Suchar IV). It seems that

human activity mainly expressed in deforestation of

the catchment area and changes in land use strongly

influenced the ecosystem of two of the three studied

lakes (Suchar IV and Suchar Wielki). As a result, their

productivity has significantly increased during the last

200 years. This process was clearly reflected in the

subfossil Cladocera and geochemical analysis results.

Suchar III, the most dystrophic lake in the region,

although influenced by increased transport of terres-

trial matter to the lake reflected in chemical analysis

results, still maintained its trophic status. As suggested

by DCA-based distances to modern conditions

(Fig. 7c), this particular lake seems to have been very

stable during the last 200 years.

Paleolimnological methods are very useful for

assessing lake productivity and trophic changes

through time. The subfossil Cladocera and chemical

compositions of the sediments of the three studied

lakes showed that modern chemical and physical

water properties do not reflect real trophic states well.

Our research highlights the importance of studying the

history of natural systems for assessing conservation

priorities and strategies. Although protecting these

environments is crucial, especially under ongoing

climate change, our data demonstrate that sometimes

these conservation tasks are undertaken based on

assumptions that may not be true.

Acknowledgments This study was founded by the Polish

Ministry of Science (Grant no. N306 228039). This research was

possible only with the support of the Institute of Geography and

Spatial Organization and Institute of Geological Sciences,

Polish Academy of Sciences, and employees of Wigierski

National Park, especially Lech Krzysztofiak.

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