Late Glacial–early Holocene environmental changes in Charzykowskie Lake (northern Poland) based on...

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Quaternary International xxx (2013) 1e11

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Quaternary International

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Late Glacialeearly Holocene environmental changes in CharzykowskieLake (northern Poland) based on oxygen and carbon isotopes andCladocera data

Joanna Miros1aw-Grabowska*, Edyta ZawiszaInstitute of Geological Sciences, Polish Academy of Sciences, Research Centre in Warsaw, INGPAN, Twarda St. 51/55, PL-00818 Warsaw, Poland

a r t i c l e i n f o

Article history:Available online xxx

* Corresponding author.E-mail address: [email protected] (J. Miros1a

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a b s t r a c t

Based on isotopic and cladoceran investigations of a 6-m-long sediment core, a reconstruction of pale-oenvironmental dynamics in Charzykowskie Lake (northern Poland) is proposed. The sediments consistof sandy silts at the bottom, followed (in upward succession) by gyttja characterized by increasing CaCO3

content. The measured d18O values oscillate from �9.3 to �5& and d13C from e 5.7 to þ0.4&. Weidentify 24 taxa of subfossil Cladocera and six phases (CAZ) of faunal development. Isotopic andcladoceran data together with detrended correspondence analyses (DCA) allow reconstruction of theenvironmental conditions, particularly changes in water level, temperature, and trophic status, duringthe Late Glacial and early Holocene. The lowest water temperature, connected with the inflow of coldmelt water, occurred in the Late Glacial and Preboreal period. Since the Boreal period, gradual warming isobserved and expressed through a positive trend in both d18O and d13C values and changes in Cladoceraspecies assemblages. The lake was deepest at the beginning of the Holocene as a result of dead icemelting (supply of melt water; lowering of lake bottom) and/or precipitation increase (change in at-mospheric circulation). During the Boreal period, the fall in water level takes place despite the oppositetrend observed at nearby sites. In addition, the cladoceran data indicate an initial decrease in trophicconditions from a-mesotrophic to oligotrophic and then an increase to the b-mesotrophic state.

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1. Introduction

The Late Glacial and early Holocene are the subject of manypaleolimnological studies (e.g., Moscariello et al., 1998; Schwanderet al., 2000; Hammarlund et al., 2003; Magny et al., 2006, 2007;Bohncke and Hoek, 2007; Bos et al., 2007; Boettger et al., 2009;Lauterbach et al., 2011; Apolinarska et al., 2012). At that time,rapid climate changes occurred causing environmental modifica-tions, such as changes in vegetation (terrestrial and aquatic) andlake conditions (temperature, trophic status, primary production,zooplankton development, water level).

This study presents new results and interpretations of isotopicand cladoceran investigations of the Late Glacial and Holocenesediments of Charzykowskie Lake (northern Poland). The aimwas areconstruction of the lake conditions in northern Poland (TucholaForest), particularly the changes in water level, trophic state and

w-Grabowska).

nd INQUA.

-Grabowska, J., Zawisza, E., Lad carbon isotopes and Cladoc

water temperature during the transition period from the LateGlacial to the early Holocene.

A goal was to establish the order and nature of Late Glacial andearly Holocene environmental alterations. Comparison of the re-sults with the data from nearby Skrzynka Lake (Apolinarska et al.,2012) can illustrate local variations in the evolution of the lakeecosystem resulting from the different sizes of these lakes.Charzykowskie Lake is much larger and deeper than SkrzynkaLake and features continuous carbonate sedimentation from theLate Glacial to the present day, which enables a detailed analysisof the record of environmental changes. An additional goal was tosee whether there are indications of an increase in the water levelduring the Boreal Period (Milecka and Tobolski, 2008).

2. Study site and materials

Charzykowskie Lake is located at an elevation of 121m a.s.l, nearChojnice, in the Tuchola Forest, northern Poland (Fig. 1). It is one ofthe largest lakes in Poland (1336 ha). Its maximum depth is 30.5 m,and its average depth is 9.8 m. The water volume is an estimated134,533.2 m3. The water pH is 8.4, and the conductivity is 300 mS/

te Glacialeearly Holocene environmental changes in Charzykowskieera data, Quaternary International (2013), http://dx.doi.org/10.1016/

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Fig. 1. Location of Charzykowskie Lake. ChL e drilling point of sediment core; SLS e Seven Lakes Stream; Mq e Ma1e qowne peat bog (Milecka and Tobolski, 2008); SK e SkrzynkaLake (Apolinarska et al., 2012).

J. Mirosław-Grabowska, E. Zawisza / Quaternary International xxx (2013) 1e112

cm (Ja�nczak, 1997). Lake Charzykowskie lies in a large glacialchannel with a NeS orientation andmeasures 10 km long by 2.4 kmwide (Ja�nczak, 1997).

Charzykowskie Lake is located in the northwestern part of theBrda outwash plain, which accumulated during the Pomeranianstage of the Weichselian glaciation approximately 16,200 BP(Kozarski, 1995). The Brda outwash plain consists of sands andgravels underlain by glacial tills and/or Neogene deposits (Pozna�nsilts) that crop out in morainal plateaus and along river valleys(Galon, 1953).

The lake sediments were sampled in March 2006 using a Liv-ingstone type corer (Tobolski, 2010). The sampling point waslocated a significant distance from the mouth of the Brda River andthe Seven Lakes Stream (Fig. 1), in the lower part of the northernlake basin. The water depth at the sampling point was 8.4 m. Thetotal length of the sediment core was 14.2 m. The drilling site wasselected for its location outside the main flow, where the thickestsequence of sediments could be obtained. Generally, the depositswere composed primarily of calcareous gyttja and lake marl. Thelowest deposits, below a depth of 1420 cm b.l.f. (below lake floor),consisted of olive-beige, sandy silt (CaCO3 content above 40%;Fig. 2). At a depth of 1420e1340 cm, this sandy silt was replaced bylight grey, sandy, calcareous detritus gyttja, with a very low CaCO3content, below 5%. Above this, at a depth of 1340e460 cm, lightgrey, calcareous, fine detritus gyttja with increasing carbonatecontent was present. The gyttja is interbedded with white, ho-mogenous lakemarl (CaCO3 content above 80%). Overlying this, at adepth of 460e210 cm, light olive-beige, calcareous detritus gyttjawith a CaCO3 content of 40e65% occurred. The uppermost lake bedsediment (depth: 0e210 cm) consisted of olive, calcareous detritusgyttja, characterized by a CaCO3 content that decreases to 20%. Thesediment core was sliced every 5e10 cm and was subject tonumerous paleoecological analyses, including pollen, isotope andsubfossil Cladocera analysis. In this paper, the results of

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investigation of a 6-m section of the bottom sediments, whichaccumulated during the Late Glacial and early Holocene, arepresented.

3. Methods

3.1. Stable isotopes

Stable isotope ratios of authigenic carbonates are often appliedto palaeolimnological studies. The oxygen isotope compositions oflacustrine carbonates are controlled by the isotopic composition ofhost water and the water temperature at which carbonate precip-itation takes place. The oxygen isotope composition of lake water isdetermined in part by the atmospheric component of the globalhydrological cycle (air mass source), and reflects the volume-weighted mean oxygen isotopic composition of catchment pre-cipitation, and the precipitation/evaporation ratio (Craig, 1953;Hoefs, 1996; Schwalb, 2003; Leng et al., 2006).

The carbon isotope composition of authigenic carbonates isdetermined by the isotopic composition of bicarbonate (HCO3

�). The13C content in sediments is influencedmainly by exchange betweenCO2 in water and the atmosphere, by the volume of incominggroundwater and the influx of dissolved carbonates, by photosyn-thesis/respiration of aquatic plants and plankton within the lake,and by CO2 production during the decay of organic matter (Craig,1953; Ró _za�nski et al., 1998; Leng and Marshall, 2004).

Many of these factors are connected with climatic conditionse.g. the volume of lake water e with ratio of precipitation toevaporation. The results of investigations of oxygen and carbonisotopes enable the interpretation of past climate (Stuiver, 1970)and past environment (e.g. Miros1aw-Grabowska, 2009).

Analyses for oxygen and stable carbon isotopes were performedon 105 samples of carbonate sediments from depths of 810e1420 cm b.l.f., using the classical phosphoric acid method (McCrea,


Fig. 2. Schematic lithology, content of CaCO3 and oxygen and carbon isotope curves of the sediments from Charzykowskie Lake. A e Lithology: 1 e calcareous detritus gyttja, 2 e

sandy calcareous detritus gyttja, 3 e sandy silts; B e Content of CaCO3; C e Results of oxygen and carbon isotope analysis of the carbonate sediments; b.l.f. e below lake floor.

J. Mirosław-Grabowska, E. Zawisza / Quaternary International xxx (2013) 1e11 3

1950). The isotopic compositions were measured using a FinniganMAT Deltaþ gas spectrometer at the Institute of Geological Sciencesin Warsaw, Poland. The oxygen and carbon isotope ratios are pre-sented in standard delta notation (d18O, d13C), versus the V-PDBstandard and are presented in the form of curves of variation ofd18O and d13C (Fig. 2). The analytical error was �0.1& for d18O and�0.05& for d13C.

3.2. Cladocera analysis

Cladocera remains are often found in lacustrine sediments.Cladocera are primitive crustaceans that are dominant amongzooplankton and inhabit lakes with various chemical conditions.The presence and frequency of certain species reflect the climaticand hydrologic conditions of the lake. Fluctuations in pH correlatewith faunal diversity. The fauna also responds to changes in waternutrient concentrations. Cladocera analysis has been applied manytimes to paleoenvironmental reconstruction of the Late Glacial andHolocene periods (e.g., Szeroczy�nska, 1998, 2002; Hofmann, 2003;Milecka and Szeroczy�nska, 2005; Sarmaja-Korjonen et al., 2006;Sienkiewicz et al., 2006). There have been several attempts toreconstruct water level changes based on changes in the cladoceranrecord (Hofmann, 1999; Sarmaja-Korjonen et al., 2003) because theratio of planktonic to littoral forms reflects changes in the propor-tional areas of littoral and open water habitats (Alhonen, 1970;Korhola et al., 2000; Szeroczy�nska and Gasiorowski, 2002). Clado-cera analysis has also been used to estimate the trophic status ofand human impact on lakes (Alhonen, 1985; Goslar et al., 1999;Szeroczy�nska, 2002).

The Cladocera analysis was conducted on 113 samples spanningthe core depth interval 810e1420 cm b.l.f. The sediments wereprocessed in a laboratory in accordance with the standard method(Frey, 1986; Korhola and Rautio, 2001). Each sample (1 cm3 of freshsediment each) was boiled in a 10% KOH solution for 20 min andwas stirred using a magnetic stirrer to remove humic matter and


later treated with HCl to eliminate carbonates. The residue waswashed and sieved using a 40-mm sieve and diluted in 10 cm3 ofdistilled water. A tenth of a milliliter of solution was used for everymicroscope slide, and two to four slides were counted from eachsample. The extracted remains were identified using an OLYMPUSmicroscope, based on the papers by Flössner (1972, 2000) andSzeroczy�nska and Sarmaja-Korjonen (2007). In each sample, all ofthe skeletal elements (head shield, shell, postabdomen, claw,ephippium) were counted and used to calculate the number ofindividuals per cm3 of fresh sediment. The results of the qualitativeand quantitative analyses are expressed as the relative abundanceand Cladocera habitat preferences diagrams (Fig. 3). Cladocerazones (CAZ) were distinguished based on significant changes in therelative abundance of Cladocera and composition of species. Theclassifications of Cladocera habitat preferences were based on thefindings of Hann (1990) and Fryer (1985, 1993).

3.3. Carbonate content

The carbonate content of the lake sediments was estimatedbased on 20 samples taken from depths of 820e1420 cm and usingthe loss on ignition (LOI) method (Heiri et al., 2001). The sedimentsamples were dried at 105 �C and homogenized in an agate mill.Next, the samples were subjected to a two-step procedure. In a firstreaction, organic matter is oxidized at 500e550 �C to carbon di-oxide and ash. In a second reaction, carbon dioxide is evolved fromcarbonate at 900e1000 �C, leaving an oxide. The carbonate contentwas calculated using the relationship Carb ¼ 1.36 LOI925, whereLOI925 ¼ CO2 evolved from the sample, as defined by Heiri et al.(2001).

3.4. AMS radiocarbon dating

AMS radiocarbon dating was performed on seven samples ofterrestrial plant macrofossils at the Pozna�n Radiocarbon Laboratory


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Please cite this article in press as: Miros1aw-Grabowska, J., Zawisza, E., Late Glacialeearly Holocene environmental changes in CharzykowskieLake (northern Poland) based on oxygen and carbon isotopes and Cladocera data, Quaternary International (2013), http://dx.doi.org/10.1016/j.quaint.2013.07.019

Fig. 4. Depth-age model based on calibrated AMS 14C data. Error bars for individualAMS 14C dates indicate calibrated 2s ranges. The grey shading represents the 2sprobability range. The age model was constructed by linear interpolation betweenmidpoints of the individual 2s ranges.


in Pozna�n, Poland (Table 1). All of the conventional radiocarbonagesmentioned in the text were calibrated using “The CalPal OnlineRadiocarbon Calibration” (http://www.calpal-online.de).

Table 1AMS 14C dates of macrofossils from the Charzykowskie Lake sediments.

Depth b.l.f. (cm) Laboratory number Dated material AMS 14C yr BP cal. yr BP (68% range) 2s cal. yr BP used in age model

848 Poz-36565 Pine bark 5740 � 60 6472e6621 6547998 Poz-36566 Pine bark 7350 � 50 8078e8263 81711139 Poz-32393 Pine bark 8940 � 70 9941e10174 10,0581145 Poz-32392 Pine bark 8960 � 80 9942e10190 10,0661223 Poz-36567 Pine needle 9800 � 120 10971e11409 11,1901298 Poz-36568 Birch bark 9900 � 130 11241e11650 11,4461337 Poz-36569 Fruit of birch and peat 17180 � 100 20320e20929 20,625

b.l.f. e below lake floor.

3.5. Statistical methods

Detrended correspondence analyses (DCA) (Hill and Gauch,1980) were performed using the average abundance of Cladoceraspecies and rescaling by equalizing the average dispersion withinsamples, based on a standard deviation of one. The samples wereordinated in bidimensional coordinated spaces (biplots) defined bythe biological data, i.e., the Cladocera species composition (Fig. 5A).In addition, the species scores on axes 1 and 2 (Fig. 5B) were plottedstratigraphically, by age (depth), which provides a simplified pic-ture of environmental changes through time. DCA analysis waspreferred over others because it avoids arch effect derived fromlinearity assumptions. The gradient lengths of the DCA areexpressed as standard deviation (SD) units and may be interpretedas a measure of ecological turnover (Gauch, 1982). All data pro-cessing was performed using the R statistics program (RDevelopment Core Team, 2009).

4. Results

4.1. Stable isotopes

The oxygen isotope ratio varies between �9.3 and �5.0&,whereas the carbon isotope ratio oscillates between �5.7and þ0.4& (Fig. 3). The lowest deposits (below a depth of 1380 cmb.l.f.), consisting of olive-beige, sandy, calcareous silt, display low


values of both d18O, approximately �8.1&, and d13C, approximately�4.6&. In the overlying sediments (depths of 1380e1330 cm b.l.f.),consisting of sandy, calcareous detritus gyttja, the values of d18Oand d13C abruptly increase, to �5& and þ0.4&, respectively. Bothd18O and d13C reach their maxima in this depth interval. The later(higher) sediments are more calcareous. Specifically, calcareousdetritus gyttja and then white lake marl were deposited at depthsof 1330e1130 cm b.l.f. In this depth interval, the lowest values ofd18O and d 13C were measured. Both d18O and d13C reach theirminima in this depth interval. The d18O values drop to �9.3&. Onlyat depths of 1215e1235 cm b.l.f. do they increase, to approximately1&. The d13C values systematically decrease to�5.7&, and only at adepth of 1220 cm do they slightly increase, to approximately 0.7&.Higher, at depths of 1130e1045 cm b.l.f., the value of d18O starts torise and reaches approximately �7.7&. The d13C slightly increasesto approximately �5.1&. In the overlying sediments (depths of1045e810 cm), d18O values remain constant, at approximately�8&, and values of d13C oscillate around e5&, a trend whichslightly increases upward.

4.2. Cladocera analysis

In the studied material, 24 species of Cladocera belonging to fivefamilies, Bosminidae, Daphniidae, Leptodoridae, Chydoridae and

Sididae, were found. The predominant species throughout thesediments were Bosmina longirostris (O.F. Müller), Bosmina (E.)longispina (O.F. Müller), Bosmina (E.) coregoni (Baird), Alona affinis(Leydig) and Chydorus sphaericus (O.F. Müller).

The changes in Cladocera populations and species compositionin the studied part of the Charzykowskie sediment core (LateGlacial to Middle Holocene) allow us to distinguish six Cladoceraassemblages zones (CAZ), which reflect lake-development stages.The results of the analyses are shown in the absolute frequencydiagrams in Fig. 3.

4.2.1. Phase I (>1420 cm depth)The lowest sediments, deposited at depths below 1420 cm,

belong to phase CAZ I. The Cladocera species assemblage in thislayer was so distinct that it was possible to distinguish the oldestsediments as a separate Cladocera phase, CAZ I (Figs. 3, 5A and 5B).In this phase, the remains of 18 Cladocera species, both pelagic andlittoral, were found. Among the littoral Cladocera, plant-associatedspecies were dominant, particularly Alona affinis, Alona quad-rangularis, and Alona rectangula. Planktonic species were repre-sented by 3 species of the Bosminidae family.

4.2.2. Phase II (1380e1335 cm depth)In phase CAZ II, which spans the depth interval of 1380e

1335 cm, only six Cladocera species were found. The predominantspecies were pelagic: Bosmina (E.) longispina and Bosmina


http://www.calpal-online.de

Fig. 5. 5A. DCA analysis performed for the sediments of Lake Charzykowskie based on Cladocera species. Aaffi e Alona affinis, Acost e Alona costata, Arect e Alona rectangula, Aquare Alona quadrangularis, Alexc e Alonella excisa, Alnana e Alonella nana, Aharp e Acroperus harpae, Aelong e Alonopsis elongta, Bcore e Bosmina (E.) coregoni, Blongr e Bosminalongirostris, Blongs e Bosmina (E.) longispina, Csphae e Chydorus sphaericus, Cpiger e Chydorus piger, Crecti e Camptocercus rectirostris, Drost e Disparalona rostrata, Dlongi eDaphnia longispina group, Elamel e Eurycercus lamellatus, Gtestu e Graptoleberis testudianria, Ptrigo e Pleuroxus trigonellus, Puncin e Pleuroxus uncinatus, Rfalca e Rynhotalonafalcata, Scryst e Sida crystallina. 5B. DCA analysis Axis 1 and 2 scores versus depth, respectively.


longirostris. Cladocera from the littoral zone constitute only 5% oftotal Cladocera. Littoral species were represented by four taxa, i.e.,Alona affinis, Alona quadrangularis, Alonella nana, and Chydorussphaericus, but their frequency was very low and did not exceed1200 individuals per cm3 of sediment.

4.2.3. Phase III (1335e1280 cm depth)Phase CAZ III spans the sediment depth interval from 1335 to

1280 cm. In this phase, the number of Cladocera species increased


to 18 from only 6 species in the previous phase, and the total Cla-docera density significantly increased (Fig. 3). Cladocera living inthe open-water zone, namely Bosmina longirostris, Bosmina (E.)longispina, and the Daphnia longispina group, were dominant andconstituted more than 60% of total Cladocera. Littoral species livingin association with plants were also numerous, dominated mostlyby Alona quadrangularis, Alona rectangula, Alonella nana, Chydorussphaericus and Sida crystallina. The rare species Alonopsis elongataappeared in the lower part of this phase.



4.2.4. Phase IV (1280e1165 cm depth)Phase CAZ IV, which spans depths of 1280e1165 cm, is charac-

terized by an increase in Cladocera density to 4500 individuals percm3. The open-water species were dominant. Among these species,the frequency of the D. longispina group significantly increased. Inaddition, a new species, Bosmina (E.) coregoni, appeared in thisgroup. Species living in association with plants were also verynumerous, represented mostly by the Aloninae subfamily. Clado-cera living in sediments were represented byMonospilus dispar andPleuroxus trigonellus.

4.2.5. Phase V (1165e1060 cm depth)In phase CAZ V (depth interval 1165e1060 cm), the frequency

and amount of pelagic species decreased (from 4 in the previousphase to 2), except Bosmina longirostris, which reached theirhighest frequency in the studied sequence. Bosmina (E.) longispinacompletely disappeared by the end of the phase. Littoral Cladocerawere represented by subfamily Aloninae, i.e., Alona affinis, Alonarectangula. The frequency of Chydorus sphaericus significantlydecreased, and its frequency position was taken by Chydorus piger.

4.2.6. Phase VI (1060e810 cm depth)Phase CAZ VI spans the sediment depth interval of 1060e

810 cm. The lower part of this phase (depths of 1060e990 cm) wascharacterized by the reappearance and continuous presence of thepelagic Bosmina (E.) longispina and Bosmina (E.) coregoni. In addi-tion, the population of Bosmina longirostris decreased in this part ofcore and reached its lowest frequency in any part of the core, i.e.,200 individuals per cm3). The pelagic species comprised approxi-mately 60% of the total Cladocera. Littoral Cladocera were repre-sented primarily by plant-associated species, namely Alona affinis,Alona quadrangularis and Alona rectangula. In the sediments fromdepths of 1016e1006 cm, the rare species Alonopsis elongataappeared again.

Bosmina (E.) longispina disappeared from the sediments above adepth of approximately 990 cm. In this part of the core, the fre-quency of Bosmina longirostris increased again; this species hadbeen continuously present in high abundance. In the second part ofphase VI, pelagic species dominated and constituted more than 65%of the total Cladocera. Species living in association with plants andsediments were represented by Alona affinis, Alona rectangula,Alona quadrangularis, Chydorus piger, Monospilus dispar, and Pleu-roxus uncinatus.

4.2.7. Results of DCADCAwas intended to delineate the ecological space taken up by

the various Cladocera species. The species arrangement as depictedin the reduced biplot shows an environmentally associated distri-bution along axes 1 and 2. Based on the distribution of Cladoceraspecies, the distribution along axis 1 likely reflects the watertemperature (Fig. 5A, B). Cold-tolerant species (e.g., D. longispinagroup, Alonella nana, Chydorus sphaericus) are plotted on thenegative part of the axis, whereas species with higher thermal re-quirements are plotted on the positive part of axis 1 (e.g., Dis-paralona rostrata, Alonella excisa, Alona costata, Bosmina coregoni).The DCA results indicate that the environmental gradient associ-ated with axis 2 (Fig. 5A, B) is also a very important elementdifferentiating the samples through time, although it is difficult tointerpret the axis 2 plot. It is possible that axis 2 may be associatedwith changes in water level.

4.3. Radiocarbon dating

Seven AMS 14C dates for terrestrial plant macrofossils (samplesof Pinus and Betula) were obtained from the Charzykowskie Lake


sediments (Table 1). The calibrated ages of all these samples are instratigraphical order. An age-depth model based on calibrated AMS14C dates was constructed by linear interpolation between mid-points of the individual 2s ranges (Fig. 4).

The 14C age of sample Poz-36569 does not match with thepollen-based biostratigraphy, as the midpoint age of 20,625 calBP appears too old to correspond to the late/mid-Younger Dryas.Based on the pollen data, “.the analyzed sediment core fromLake Charzykowskie includes all Holocene climatic periods, andthe bottom section of the sediments was accumulated duringthe Younger Dryas” (Tobolski, 2010). In addition, the outwashsediments building up the bottom of Charzykowskie Lake weredeposited during the Pomeranian stage of the Weichselianglaciation, only approximately 16,200 BP (Kozarski, 1995).

5. Interpretation

We used the isotopic and Cladocera data to reconstruct thepaleoenvironmental conditions of the Charzykowskie Lake, inparticular the changes in water level, trophic state, and watertemperature, and connected these parameters with vegetationdevelopment.

5.1. Before ca. 11,600 cal BP e Late Glacial (LG)

The oldest analyzed sediments in Charzykowskie Lake accu-mulated before approximately 11,700 cal BP (during the LateGlacial). The carbonate sediments, primarily sandy calcareous silts,displayed d18O values of approximately �8.1& and d13C values ofapproximately �4.6&. These isotopic values might be characte-ristic of the early autochthonous lake carbonates.

The high number of Cladocera species (18) living in both thelittoral and pelagic zones during phase CAZ I strongly suggest thatfavorable ecological conditions for Cladocera zooplankton devel-opment prevailed at this time. The abundances of species associ-ated with macrophytes and macrophytic sediments indicate thatplants were common in the littoral zone of the lake (Fig. 3). Thecomposition of Cladocera species suggests that sediments of phaseCAZ I were deposited in a quite warm and humid period and thatthe edaphic status of the lake might be reconstruct as a-mesotro-phic. The DCA results (axis 1 score, Fig. 5B) also suggest that theoldest sediments accumulated in a warm climate.

Next, approximately 11,700e11,600 cal BP, higher values of d18Oand d13C, i.e., approximately 2e3& and 4e5&, respectively,occurred. These isotope values were close to the d18O and d13Cvalues of marine limestones, respectively �5.25& and þ0.56&(Keith andWeber, 1964), suggesting an input of older carbonates tothe lake due to erosion of the lake’s catchment. At that time, siltswere replaced by sandy low-calcareous detritus gyttjawith a CaCO3content of 3e15%. The increase in erosion in the surrounding areamay have been caused by a scarcity of plant cover associated withcold climate conditions.

The reduction in the density and numbers of Cladocera species(from 18 to 4) in phase CAZ II indicates important ecologicalchanges in the lake and its catchment. At that time, the predomi-nant species were pelagic, constituting 95% of the population(Fig. 3). The presence of the very cold-tolerant species Bosmina (E.)longispina and Bosmina longirostris suggests unfavorable climateconditions, in particular, cold water and low productivity. The lowfrequencies of two littoral species, Alona affinis and Chydorussphaericus, are noted. These species are known for their tolerance ofcold climate conditions (Szeroczy�nska, 1985; De Eyto and Irvine,2001; Sarmaja-Korjonen, 2001; Bennike et al., 2004; Belyaevaand Deneke, 2007). Sometimes these species are even called “arcticspecies” (Whiteside, 1970). It is quite possible that the area of the



littoral zone occupied by plants was significantly reduced. Thepresence of only these four species in the sediments and significantdomination by planktonic species suggests that the lake’s water atthat time was cold and oligotrophic. The unfavorable climate con-ditions during CAZ II are also suggested by low DCA values alongaxis 1, which indicate cold lake water (Fig. 5B). The results of theDCA plotted on axis 2 suggest that the lake likely was quite shallowat that time (Fig. 5B).

5.2. Ca. 11,600e11,400 cal BP (Late GlacialeHolocene)

The transition from Late Glacial to Holocene conditions (YoungerDryas/Preboreal) took place from 11,600 to 11,400 cal BP and is veryclearly reflected in the isotopic curves, expressed as a decrease in thed18O (of approximately 4&) and d13C values (of approximately 5&).These lower d18O and d13C values were caused by a change in theisotopic composition of the lake water. These large drops in thevalues of d18O and d13C were due to an inflow of water rich in lightisotopes, which may suggest an inflow of water from melting deadice, a rising water level and/or increased precipitation.

The Late Glacial/Holocene transition period is also very wellmarked in the cladoceran community (CAZ III) by an increasedpopulation density and species count (from4 to 18). The dominationby pelagic species suggests a rising water level in the lake. A pres-ence of littoral species, particularly those living in association withplants, suggests the development of a littoral zone and an increasingamount of biogenic material in the water (Fig. 3). However, thepresence of species typically associated with cold water, the D.longispina group and particularly Alonopsis elongata (Hofmann,2000; Bennike et al., 2004; Szeroczy�nska and Zawisza, 2010), in-dicates that the lake water was still quite cold at that time. Thelowest DCA values along axis 1 also indicate the lowest water tem-perature (Fig. 5B). Simultaneously, the DCA values along axis 2suggest rising lake water levels. The rising water level and decreasein the lake’s water temperature can be explained, as they can frominterpretation of the isotopic data, by the melting of dead ice.

5.3. Ca. 11,400e10,200 cal BP (Preboreal)

During the period 11,400e10,200 cal BP, the lithology of thesediments changed and calcareous-rich deposits (calcareous gyttjaand lake marl) accumulated. Initially, the d18O and d13C values inthese sediments were low, below �8.5& for d18O and below �5&for d13C. Only at approximately 11,200e11,150 cal BP was there asmall shift of 0.8e1& in the isotopic values, likely due to a fluc-tuation in the evaporation/precipitation ratio and oscillations in thewater level. The low oxygen isotopic values suggest a humidclimate and a higher water level during the Preboreal period. Thelow carbon isotopic values indicate good levels of oxidation in thewater. Approximately 11,000 cal BP, the isotopic values drop andreach their minima of�9.3& (for d18O) and�5.7& (for d13C). Theselowest isotopic values suggest a high water level; the lake waslikely its deepest at this time.

The start of the Preboreal period is reflected in the cladocerancommunity (CAZ IV) by an increasing number of species andincreasing populations, suggesting that ecological conditions in thelake were favorable for Cladocera (Fig. 3). The DCAvalues along axis1 suggest a rise in water temperature at the beginning of the Ho-locene (Fig. 5B), which had significance for the development of theCladocera community. The continuous presence of four planktonicspecies, which became dominant, and the presence of numerouslittoral species demonstrate that Charzykowskie Lakewas deep andhad well-developed pelagic and littoral zones at the beginning ofthe Holocene. The highest DCA values along axis 2 confirm thatCharzykowskie Lake was deepest during Preboreal time. In


addition, Cladocera assemblages, particularly domination bypelagic species, indicate that the lake was quite rich in biogenicmaterial and was in a a-mesotrophic state at that time.

5.4. Ca. 10,200e9000 cal BP (Boreal)

Approximately 10,200 cal BP, a systematic increase of ca.1.5& ind18O values and ca. 0.5& in d13C values occurred. Such trends maybe associated with climatic warming (oxygen isotope) and an in-crease in the photosynthetic activity of phytoplankton and mac-rophytes (carbon isotope). At that time, a drop in the water levellikely occurred. An increasing CaCO3 content in the gyttja wasassociated with climatic warming, which led to higher bio-production (Nitychoruk, 2000).

A slightly lower water level during the Boreal period (CAZ V)was also noted in the Cladocera assemblages, mostly as a decreasein the density and number of species of pelagic Cladocera. Awarming of the climate likely caused an abundance of phyto-plankton and macrophytes, which covered a significant portion ofthe lake, reduced the open-water area and consequently decreasedthe planktonic species. The highest frequency of Bosmina long-irostris at this point in the core may suggest that the trophic statuswas quite high andmay be classified as b-mesotrophy. At the end ofthis phase (approximately 9270 cal BP), Bosmina (E.) longispinacompletely disappeared from the sediments. This species is knownto be strictly planktonic and favor cold, low-nutrient waters(Vijverberg, 1980; Herzig, 1984; Nevalainen et al., 2013). Thedisappearance of this species documents a shrinking of open waterarea and an increasing trophic level. The results of the DCA,particularly the low values along axis 2, suggest a drop in the waterlevel during the Boreal period (Fig. 5B). The DCA results along axis 1show a continuously increasing trend, indicating a gradual rise inwater temperature.

5.5. Ca. 9000 cal BP (Atlantic)

Since approximately 9000 cal BP, the constant values of bothd18O (approximately �8&) and d13C (approximately �5.0&) sug-gest stable climatic and hydrologic conditions and/or a fast sedi-mentation rate. At that time, the isotopic composition of the lakewater was constant and carbonates are marked by similar isotopicratios. Only approximately 8200 cal BP did a slight decrease of 0.2e0.3& in d 18O occur (cooler and/or higher water level?). In addition,we observed a slight positive trend in d13C values, whichwere likelyassociated with a higher trophic state.

The beginning of the Atlantic period (Middle Holocene) ischaracterized by the reappearance of two pelagic species from thesubgenus Eubosmina and a decrease in Bosmina longirostris (CAZ VI,Fig. 3). Such changes in the makeup of the Cladocera communityindicate that important ecological changes occurred in Charzy-kowskie Lake in the beginning of the Atlantic period. It is possiblethat the reappearance of Eubosmina is connected with anexpanding open-water area and a decrease in water fertility andtemperature (Hofmann, 1984). Decreasing nutrient levels are alsodocumented by a decrease in the Bosmina longirostris population,which reaches its minimum at this point in the core. This speciesusually inhabits fertile lakes (Szeroczy�nska, 1998; Sarmaja-Korjonen, 2003). Between 8400 and 8200 cal BP, the appearanceof Alonopsis elongata, known to be a cold-water species (Korhola,1999; Nevalainen et al., 2013), indicates decreasing water temper-ature. The changes in the Cladocera community, primarily theappearance of Alonopsis elongata and changes in Bosminidae, mightbe a result of temperature and trophic status changes and/or arising lake water level related to the period of climate coolingknown as episode 8.2 ka (Weninger et al., 2006; Magny et al., 2007;



Sarmaja-Korjonen and Seppä, 2007; Szeroczy�nska and Zawisza,2010). The high water level is also suggested by the DCA valuesalong axis 2 (Fig. 5B), which increased at the beginning of theAtlantic period.

Bosmina (E.) longispina, which prefers cold and low-nutrientwaters (Hofmann, 1984), disappeared for approximately 200years beginning approximately 8000 cal BP. At that time, the pop-ulations of pelagic species preferring more fertile waters, i.e., Bos-mina longirostris and Bosmina (E.) coregoni, increased. In addition,the populations of species associated with littoral macrophytes andmacrophytic sediments also rose (Fig. 3). It seems that changes inCladocera assemblages were triggered by the warming Atlanticclimate. Climate warming may have provoked development ofaquatic vegetation and increasing water fertility. The Holoceneclimatic optimum is also well documented by the highest DCAvalues along axis 1 (Fig. 5B). These results suggest that of all of thetimes represented by the cores, the lake temperatures were thehighest during Atlantic time.

6. Discussion

The comparison of the isotope and Cladocera data from thesediment cores made it possible to reconstruct the changes in thewater level, trophic state, and water temperature of CharzykowskieLake during the transition from the Late Glacial to the early Holo-cene periods.

The environmental changes observed in Northern Hemispherelakes after the end of the last glaciation (Weichselian) were causedby global factors, primarily climatic amelioration expressed as a risein temperature and then a decline in herb vegetation and thespread of Betula (Bos et al., 2007; Kupryjanowicz, 2007; Stan�cikaitéet al., 2008), modified by many local factors. The local factors arelinked with, e.g., geographic location, particularly the distance fromthe North Atlantic Ocean and adjacent ice sheets. CharzykowskieLake was situated only approximately 600 km from the Scandina-vian ice sheet during the Younger Dryas (Lundqvist and Wohlfarth,2001).

At the end of the last glaciation (approximately 11,600 cal BP),northern Poland was under the influence of anticyclonic windsover the Scandinavian ice sheet and strengthening easterly windssouth of the ice (Yu and Harrison, 1995). Thus, cold and dry con-ditions dominated in this region.

In many locations in Europe, the transition between the LateGlacial and Holocene is expressed by an increase in d18O values,which is primarily interpreted as the effect of temperature in-creases (e.g., Ró _za�nski, 1987; Mayer and Schwark, 1999; Starkelet al., 1999; Schwander et al., 2000; Ralska-Jasiewiczowa et al.,2003; Magny et al., 2006; Bos et al., 2007; Apolinarska and Ham-marlund, 2009).

In Charzykowskie Lake, the isotopic record from the beginningof the Holocene shows the opposite trend, i.e., a decline in d18Ovalues. This decrease in isotopic values reflects a substantial changein the isotopic composition of the lake, which likely was caused byan inflow of water enriched in light oxygen isotopes. This may havebeen due to an increase in precipitation and/or inflowof water frommelting dead ice. Climate warming, particularly increasing airtemperature, initiated this melting of dead ice and the inflow ofcold water enriched in light oxygen isotopes. This process is alsosuggested by the DCA results, in particular the values along axis 1,which reach their minimum at this point. The values along axis 2indicate a rising water level associated with the melting of dead iceat this time. The presence of cold lakewater is also suggested by theCladocera assemblage, in particular the presence of Alonopsiselongata, a cold-tolerant species typical of Northern areas (Hessenand Walseng, 2008; Szeroczy�nska and Zawisza, 2010; Nevalainen


et al., 2013). The possibility that dead ice persisted for a relativelylong time, until the Allerød, is suggested by findings from thenearby Lake Skrzynka basin (Apolinarska et al., 2012).

In northern Poland, the influence of warm, moist air massesfrom the North Atlantic, causing the rise of summer temperatures,was possible after the retreat and then decline of the ScandinavianIce Sheet and unblocking of the Westerlies. This temperature in-crease had to cause melting and then a disappearance of stagnantice, which delivered the light oxygen isotopes to the lakes (a rapiddecrease in the d18O values was noted in the studied profile). Inaddition, an influx of cold melt water from disintegrating stagnantice and rapid sedimentation may have kept lake’s organic produc-tivity low and delayed the response of the limnic environment tothe temperature increase (Wohlfarth et al., 2007).

The isotopic record of the Preboreal period is characterized by afurther decline and finally minima in the d18O and d13C values.Approximately 11,200 cal BP, a small increase in d18O and d13Cvalues occurred. A similar spike at that time is observed in the re-cord from other lakes in northern Poland: Skrzynka and Ha�ncza(Lauterbach et al., 2011; Apolinarska et al., 2012).

The low d18O values may be explained by an increase in pre-cipitation and/or a rise in the water level. The larger amount ofmeteoric water may have caused a rise in the water level andrelative depletion in heavy isotopes. The lake was likely at itshighest level (and deepest) at that time. A rising water level is alsosuggested by the highest DCA values along axis 2, which reflectschanges in the water depth. A similar substantial decrease in d18Ovalues and then a minimum value, occurring at the onset of theHolocene and during the early Holocene, were observed in PergusaLake (Sicily). These trends were interpreted as indicators of awettest period (Zanchetta et al., 2007). The makeup of the Clado-cera community, particularly an increase in planktonic species,indicates a large area of openwater at the lake and also suggests anincreasing lake depth.

During the Boreal period, a shift to higher d18O values is noted.Such a trend may have been caused by progressive climaticwarming and a rise in the precipitation/evaporation ratio, likelydue to an influence of the Westerlies expressed by a rise in tem-perature. This trend of rising temperature and falling water levelare well supported by the results of the DCA. Similar ecologicalchanges are also noted in the Cladocera assemblages, i.e., adecreasing planktonic Cladocera population indicates a fallingwater level in Charzykowskie Lake at this time. This is the oppositeof the trends observed in the adjacent peatbog named Ma1e qowne(Milecka and Tobolski, 2008). In the nearby Skrzynka Lake, “.thedeep water conditions noted in the Preboreal period might havecontinued in the early Boreal.” (Apolinarska et al., 2012).

In the early Atlantic period, a decrease in fertility and watertemperature and a slight decrease in the oxygen isotope data arenoted, suggesting a brief cooling. These changes were well corre-lated with cold episode 8.2 ka, which occurred at the beginning ofthe Atlantic period (Sarmaja-Korjonen and Seppä, 2007; Hessenand Walseng, 2008; Szeroczy�nska and Zawisza, 2010). Since 8 ka,stable hydrologic conditions and other conditions favorable forzooplankton development were present in Charzykowskie Lake.The increases in planktonic and littoral Cladocera species werelikely a consequence of increasing water fertility, which, in turn,was a consequence of increasing water temperature.

7. Conclusions

� The sedimentation in Charzykowskie Lake started before11,700 cal BP. The isotopic values reflect the early autochtho-nous lake carbonates. The relative abundance of certain



Cladocera species suggests that sediments were deposited in awarm, humid period.

� Approximately 11,700e11,600 cal BP, a period of intenseerosion of the lake’s catchment occurs. The correspondencebetween the lake’s isotope values and those of marine lime-stones suggest an input of older carbonates into the lake. Thepresence of only “arctic” species and domination by planktonicspecies indicate that the lake was cold and oligotrophic at thattime.

� In Charzykowskie Lake, the transition between the Late Glacialand Holocene is expressed by a decline in d18O values, likelycaused by an inflow of water frommelting dead ice, and by thepresence of species typical of cold water, D. longispina groupand, in particular, Alonopsis elongata. Domination by pelagicspecies and low d18O values suggest a high lake level duringthis transition.

� In the Preboreal period, conditions favorable for Cladoceradevelopment are revealed by an increasing number of speciesand populations. At that time, Charzykowskie Lake had well-developed pelagic and littoral zones, and likely its water leveland depth were greatest.

� During the Boreal period, a progressive warming of the climateis indicated by a shift to higher d18O values, which also suggestsa falling water level. A declining planktonic Cladocera popu-lation confirms a falling water level in Charzykowskie Lake atthat time.

� During the early Atlantic period, a decrease in lake fertility andtemperature and a slight lowering in d18O values are noted,suggesting a brief period of cooling (8.2 ka event?). Since 8 ka,stable hydrologic conditions and other conditions favorable forzooplankton development were present again, An increase inplanktonic and littoral Cladocera species was likely an effect ofincreasing water fertility, which was an effect of rising tem-perature. During the Atlantic period, lake temperatures werethe highest of any times represented by the sediment cores.

� The cladoceran assemblies indicate an initial decrease of thelake’s trophic state from a-mesotrophic to oligotrophic andthen an increase up to the b-mesotrophic state.

Acknowledgments

Wewould like to express our thanks to Prof. Kazimierz Tobolskiand Dr. Mariusz Ga1ka for providing the sediment samples and veryinteresting cooperation and Dr. Micha1Woszczyk for the estimationof CaCO3 content. The studies of Charzykowskie Lake sedimentswere financed by the Institute of Geological Sciences Polish Acad-emy of Sciences.

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Late Glacial–early Holocene environmental changes in Charzykowskie Lake (northern Poland) based on...

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