Long (MIS 5e – 3) environmental history of a paleolake in central Poland recorded in the succession from Kubłowo

Long (MIS 5e – 3) environmental history of a paleolake in central Poland recorded in the succession from Kubłowo

Quaternary International xxx (2016) 1e17 Contents lists available at ScienceDirect Quaternary International journal homepage: www.elsevier.com/locat...

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Quaternary International xxx (2016) 1e17

Contents lists available at ScienceDirect

Quaternary International journal homepage: www.elsevier.com/locate/quaint

Long (MIS 5e e 3) environmental history of a paleolake in central Poland recorded in the succession from Kubłowo Joanna Mirosław-Grabowska a, *, Monika Niska b, Małgorzata Roman c a

Institute of Geological Sciences, Polish Academy of Sciences, Research Centre in Warsaw, INGPAN, Twarda St. 51/55, PL-00-818 Warsaw, Poland w St. 27, PL-76-200 Słupsk, Poland Institute of Geography and Regional Studies, Pomeranian University in Słupsk, Partyzanto c dz, Narutowicza St. 88, PL-90-139 Ło dz, Poland Department of Geomorphology and Palaeogeography, Faculty of Geographical Sciences, University of Ło b

a r t i c l e i n f o

a b s t r a c t

Article history: Available online xxx

One of the longest (above 7 m thick) uninterrupted Eemian-Vistulian (Weichselian) pollen successions in Poland is known from Kubłowo (central Poland). The huge importance of this succession is manifested in the possibility of study the evolution of the lake environment under the influence of changing, from interglacial to glacial climatic conditions. The reconstruction of the environmental changes at Kubłowo and its surroundings show the short climate fluctuations and a detailed picture of the natural conditions prevailing in central Poland (central Europe) during the period from approximately 126,000 to 55,000 years BP. Based on the results of multiproxy analysis, four lacustrine stages linked with a higher water level and three boggy stages (low water level) of evolution of the Kubłowo paleolake were observed. The pollen diagram from Kubłowo reveals a succession of eighteen local pollen assemblage zones (K1K18 L PAZ). The K1 to K7 LPAZ zones record the interglacial succession correlated with the Eemian. The next pollen zones, K8 to K18, are linked with the Vistulian. The pollen sequence shows, between the Eemian Interglacial (MIS 5e) and the last glaciation (second stadial of the Plenivistulian (MIS 4-3), four cold intervals that are interpreted as stadial phases: Herning, Rederstall, Schalkholtz and Ebersdorf € rup, Odderade and Oerel. separated by three temperate intervals (interstadials): Bro In the studied sediments, 25 species of subfossil Cladocera were identified, and ten zones of fauna development were distinguished. The unique feature of this profile is the good preservation of above 100,000-year-old Cladocera remains and a high diversity of species, which are primarily littoral. The Middle and Late Eemian were periods convenient to Cladocera development due to high temperatures, high water levels, high trophy and occurrence macrophytes. A deterioration of fauna living conditions started in the Late Eemian and next took place in the Vistulian cold events. The results of isotopic and geochemical analyses of organic matter from Kubłowo have enabled the reconstruction of varying environmental conditions and the origin of organic matter. The organic carbon content (TOC) changes from 1% to 40% and is closely related to climatic conditions. During the Eemian, the amount of TOC systematically increased. The lowest values are recorded in the coldest periods during the Herning, Rederstall and Schalkholz Stadials. The C/N atomic ratio fluctuates from 13 to 34 and the € rup Interstadial. The values of d13C change from 27.9 to 12.9‰, and highest values occur in the Bro d15N values oscillate between 1.6 and þ5.3‰. © 2016 Published by Elsevier Ltd.

Keywords: Cladocera Carbon and nitrogen stable isotopes Pollen stratigraphy Palaeoenvironment Eemian Vistulian (Weichselian) Glaciation

1. Introduction

* Corresponding author. E-mail addresses: [email protected] (J. Mirosław-Grabowska), [email protected] (M. Niska), [email protected] (M. Roman).

The Kubłowo paleolake contains a long Eemian-Vistulian (MIS 5e-MIS 3), lake-bog sedimentary succession. These sediments, which represent the last interglacial and the following Vistulian (Weichselian) cold period, are of great significance for investigations of dynamic environmental changes during the Late Pleistocene (Zagwijn, 1989; Kupryjanowicz, 2008; Boettger et al.,

http://dx.doi.org/10.1016/j.quaint.2016.06.027 1040-6182/© 2016 Published by Elsevier Ltd.

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 nas and 2009; Malkiewicz, 2010; Kołaczek et al., 2012; Satku _ 2012; Baltru  nas et al., 2013; Mirosław-Grabowska et al., Grigiene, 2015) and to better understand the nature of the last interglacialglacial transition. The Kubłowo paleolake is located in the Kujawy region, within an area of special importance for regional Late Pleistocene stratigraphy and particularly for the paleogeography of the last Scandinavian ice-sheet expansion onto the European lowlands (Mojski, 2005; Wysota et al., 2009; Roman, 2010; Marks, 2012). From the numerous Eemian-Vistulian sites recognized in the central Polish Plain (Mirosław-Grabowska and Niska, 2007; Mirosław-Grabowska, 2009; Kołaczek et al., 2012; Rychel et al., 2014; Kupryjanowicz and Drzymulska, 2015), which lies in the eastern part of the North European Plain, only a few are of singular importance for their long undisturbed records (MirosławGrabowska et al., 2015). A particularly long record of the last interglacial-glacial environmental changes is represented in the Kubłowo paleolake, which has been previously examined for pollen (Roman and Balwierz, 2010), and now provides rich Cladocera and stable isotopes data. Organic sedimentation at Kubłowo started with the beginning of the Eemian (MIS 5e) and lasted continuously throughout that interglacial, the Early Vistulian (MIS 5d-a) and a significant part of the Plenivistulian (MIS 4-3). In the Early Vistu€ rup and Odderade) and two cool (Herning lian, two temperate (Bro and Rederstall) intervals have been distinguished. The next three climatic oscillations correspond to the Shalkoltz and Ebersdorf Stadials and to the Oerel Interstadial. This paper presents new data from the Kubłowo paleolake sediments. Thanks to comprehensive studies of the whole lake-bog 7.6 m-thick sedimentary succession, a reconstruction of the environmental evolution of the Kubłowo paleolake has been produced. The aim of our study was to describe the development history and limnological conditions (water level, trophic state and water temperature) of the Kubłowo paleolake using multiproxy analyses (pollen, subfossil cladoceran remains, carbon and nitrogen isotopes). An attempt has been made to compare these findings, which reflect the lake evolution, with pollen data that define the vegetation history of the surrounding area. We paid special attention to the period from the Early to Middle Plenivistulian because the lakebog deposits of that age are especially rare (Kołaczek et al., 2012; Marks et al., 2015; Mirosław-Grabowska et al., 2015). We use the stratigraphic nomenclature of Poland, where the Vistulian, Eemian and Wartanian periods correspond to marine isotope stages (MIS) 2e5d, 5e and 6, respectively (Ber et al., 2007; Marks, 2011). The Vistulian is subdivided into Early (MIS 5d-a), Plenivistulian (Lower e MIS 4, Middle e MIS 3 and Upper Plenivistulian e lower part of MIS 2) and Late Vistulian (upper part of MIS 2 and lowermost part of MIS 1) (cf Roman et al., 2014; Marks et al., 2015). We correlated the Vistulian with the Western Europe Weichselian glacial period and the Wartanian with the late Saalian glacial period. 2. Regional setting The Kubłowo (52 2103000 N, 19 040 5100 E) site is located in the northernmost part of the Kłodawa Plateau, central Polish Plain, and standing barely 1 km to the south from the limit of the last Scandinavian ice-sheet (Fig. 1). To the north of that limit lies the Kujawy Lakeland, the area occupied by the Vistula ice-lobe, which delineated the Last Glacial Maximum (LGM) in central Poland. Ice advanced into the area during the Late Vistulian (MIS 2), and according to luminescence (OSL) dating, the advance occurred between 22.9 and 18.7 ka BP (Roman, 2010). A dominant landform in the Kujawy Lakeland is an undulating morainic plain, at 120e130 m a.s.l., which is mostly composed of till and is covered in local patches by glaciofluvial sands with gravels that are 1.0e2.0 m thick.

The morainic plain is cut by numerous tunnel valleys (glacial troughs), with lakes or peat plains at places (115e120 m a.s.l.). The distal part of the valleys together with the beginnings of glaciofluvial fans mark the maximum extent of the last ice-sheet in that area (Fig. 1). It can be observed that glaciofluvial fans at 125e134 m a.s.l. are the highest element in the relief of the area under consideration. Those forms are built of variable grained sands and gravels, sporadically with intercalations of flow till (Fig. 2). The fan surface mildly inclines southwards and passes into a widespread outwash plain within which the Kubłowo (Ku) borehole and two exposures (Ku-1 and Ku-2) have been placed (Figs. 1 and 2). The borehole is located in a slight close depression, at a depth 3.5 m, which is only just the visible sign of a deep steep-slope kettle-hole that developed during the decay of the Wartanian (late Saalian, MIS 6) ice-sheet and lasted as a lake or bog at least up to the second stadial of the Plenivistulian. The borehole sediment core from Kubłowo (Ku), which was taken from a depth of 3.8e11.1 m, has previously been examined for lithology and pollen content and also for radiocarbon dating at its top (Roman and Balwierz, 2010). At present, those studies are being complemented by Cladocera, selected geochemical and stable isotope research. 3. Methods 3.1. Cladocera analysis The Cladocera analysis was performed using 115 samples taken ca. every 5 cm from the core depths of 405e1050 cm. Samples of 1 cm3 were prepared according to a slightly modified standard procedure (Frey, 1986). Each sample was boiled in 10% KOH for 20 min. After washing with distilled water, the residue was sieved through a 40-mm mesh sieve. The fine material was transferred into a polycarbon test tube. Prior to counting, the remains were colored with safranine. The samples were analyzed under a light microscope with a 100e400x magnification. A minimum of 200 remains of Cladocera e 3 to 5 slides e was examined from each sample. Firstly, all of the remains from each slide were enumerated (headshields, shells, postabdomina, postabdominal claws, exopods and antennules) and the most abundant remains of each taxa were chosen to represent the number of individuals. The identification and ecological interpretation of the Cladocera remains were conducted based on the studies by Duigan (1992), Frey (1958, 1962), Goulden (1964), Hofmann (1986, 2000), € ssner (2000), Szeroczyn  ska (1985) and Korhola (1990), Flo  ska and Sarmaja-Korjonen (2007). Szeroczyn The Cladocera species were classified into three littoral habitat preference groups, bottom-dweller species, species associated with €ssner, 1964; vegetation and species restricted to vegetation (Flo Whiteside, 1970; Hofmann, 1987; Whiteside and Swindoll, 1988; Korhola, 1990) and were also categorized as planktonic (offshore) and littoral (meiobenthic) groups following Mueller (1964). The species Chydorus sphaericus and Bosmina longirostris were classified into littoral and open water zones (Whiteside, 1970). The lake water acidity was estimated on the basis of the classification of species into five pH groups developed by Krause-Dellin and Steinberg (1986). However, the distribution of Cladocera is also affected by factors such as trophy, acidification, water temperature and fish predation (Krause-Dellin and Steinberg, 1986; Korhola and Rautio, 2001). The diagram of the absolute number of Cladocera individuals and the total number of Cladocera specimens, species and species diversity were performed using the PolPal computer program (Nalepka and Walanus, 2003). Cladoceran zones were determined by Constrained Single Linkage method (ConSLink) by PolPal (Nalepka and Walanus, 2003).

Please cite this article in press as: Mirosław-Grabowska, J., et al., Long (MIS 5e e 3) environmental history of a paleolake in central Poland recorded in the succession from Kubłowo, Quaternary International (2016), http://dx.doi.org/10.1016/j.quaint.2016.06.027

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Fig. 1. Location of the Kubłowo palaeolake and geomorphology of the study area.

Preliminary results of a Cladocera analysis from only the Eemian section (1050e820 cm) of the Kubłowo profile have been published (Niska and Roman, 2014). 3.2. Isotopic and geochemical analyses The isotopic and geochemical analyses were performed using 106 samples collected every 5e10 cm from depths of 405e1050 cm. The sediments were dried at 60  C and ground. The

carbonate fraction was removed with hydrochloric acid. The carbon and nitrogen isotope compositions were analyzed using a Flash Elemental Analyser 1112 and a Thermo MAT 253 mass spectrometer, which were calibrated using an internal nicotinamide standard and reported as per mill (‰) deviations versus atmospheric N2 (d15N) and Vienna Pee Bee Belemnite (d13C). The analytical errors (l SD) for the d13C and d15N measurements were 0.17‰ and 0.24‰, respectively. The organic carbon and nitrogen concentrations were analyzed using an Elemental Analyser Vario

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Fig. 2. Lithological logs and stratigraphy of the Kubłowo sedimentary succession.

Micro Cube. The analyses were performed at the Laboratory for Isotope Dating and Environmental Studies at the Institute of Geological Sciences of the Polish Academy of Sciences in Warsaw, Poland. The d15N and d13C ratios of bulk organic matter (OM) in lake sediments can be used to infer changes in organic matter (aquatic or terrestrial source). Changes in d13C and d15N can also provide information on nutrient cycling, lake mixing regimes and water

column stability (e.g., Meyers and Ishiwatari, 1993; Meyers and s, 1999). The carbon and nitrogen stable isotopes Lallier-Verge were used to define the changes in the quantity and source of organic matter transported into the paleolake. Lake morphology, catchment topography and the relative abundances of lake and catchment plants significantly influence the relative contributions from terrestrial and aquatic sources to lake sediments (Meyers and Ishiwatari, 1993).

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The carbon to nitrogen (C/N) ratio is an indicator for nitrogen limitation in plants and other organisms. The C/N ratio indicates the primary source of organic matter in lakes, e.g., C/N ratios in the range 4e10 are usually from marine sources, whereas higher ratios are likely to come from terrestrial sources. C/N ratios less than 10e12 are typical for a mainly algal sources, and those greater than 20 are suggestive of terrestrial plant origins. Therefore, the C/N ratios of sediments have frequently been used to infer the relative proportions of organic detritus derived from algae and terrestrial plants (Smeltzer and Swain, 1985; Tyson, 1995; Meyers and Lalliers, 1999). Verge 4. Results of the investigation of the Kubłowo paleolake sediments From numerous boreholes performed in the Kubłowo vicinity, only three reached organic deposits older than the Holocene, and the Kubłowo (Ku), which is presented here, proved to be the most complete (Fig. 1). The bottom of the fossil lake basin, in which accumulations of lacustrine and bog deposits occur, lies at 112.9e114.7 m a.s.l. on Wartanian (late Saalian) glaciofluvial sands with gravels underlined with till. In the Kubłowo borehole (Ku), the lake-bog sediment series occur at depths of 3.5e11.1 m (120.5e112.9 m a.s.l.), and their thickness is 760 cm (Fig. 2). At the

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bottom (depth of 1050e1110 cm), bright grey silty sands occur and gradually pass upwards into very fine laminated sands with organic admixtures (1035e1050 cm). They are overlain by highly compressed dark grey gyttja (940e1035 cm), and a laminated and massive clayey silt (881e940 cm) lies above. Next, they are covered by dark brown loamy peat boggy deposits (820e881 cm) and then brownish-black organic shales (745e800 cm) and strongly decomposed peat with silt laminae in increasing amounts at the top (710e745 cm). Higher, at depths of 632e710 cm, laminated grey silt occurs and gradually passes into brownish-gray silty gyttja (600e632 cm) with a compressed structure. Brownish-black strongly decomposed peat appears again at depths of 570e600 cm. Subsequently, dark gray silty gyttja arises (532e570 cm), and a bright gray sandy silt with organic detritus (depth of 380e532 cm) completes the lake-bog sedimentary succession. However, the upper boundary of the succession is sharp, and the contact with overlaying glaciofluvial deposits is erosional. The Kubłowo lake-bog deposits characteristically lack CaCO3.

4.1. Previous pollen data and stratigraphy The Kubłowo pollen diagram reveals a succession of eighteen local pollen assemblage zones (K1-K18 L PAZ, Fig. 3). In Table 1, each

Table 1 Characterization of local pollen assemblage zones from Kubłowo (by Z. Balwierz, cf. Roman and Balwierz, 2010). Local pollen assemblage zones L PAZ

Depth [cm]

Description of local pollen assemblage zones L PAZ

K18 Poaceae-Artemisia-Cyperaceae

380e395

K17 Pinus-Poaceae

405e435

K16 Poaceae-Artemisia-Pinus

445e490

K15 Pinus-Poaceae-Artemisia

500e530

K14 Pinus-Betula

540e590

K13 Betula-Pinus

600

K12 Pinus-Betula

620

K11 Artemisia-Poaceae-Juniperus

625e730

The zone is marked by the dominance of herbs and dwarf shrubs. The values of AP are the lowest (15.1%) in the whole profile. The dominants are Poaceae (49,3%), Cyperaceae and Artemisia. Herbs attain a great variety, including Armeria, Helianthemum, Pleurospermum, Sanguisorba officinalis, Plantago major, P. media, Rumex acetosa, R. acetosella, Ranunculus acris t., Caryophyllaceae, Rubiaceae and Cichorioideae. AP increases to 65.3%, mainly in Pinus (46.2%), and Poaceae falls. Artemisia, Cyperaceae and Sphagnum pollen are more abundant. Herbs are less diversified than in the preceding zone. Pollen of Salix and Juniperus as well as Botrychium spores are present. A rapid decrease in the AP value indicates the outstart of the next zone. The AP content in this zone varies from 44.2 to 58.2%. The values of Betula pollen are lower than Pinus and more stable. The percentage of spores in the zone is relatively high. The spores belong mainly to Sphagnum and Polypodiaceae. Botrychium spores are also present. A rise in AP, mainly Pinus, marks the upper limit of the zone. The curve of Pinus falls. The share of Betula is slightly below 20%. The variety of herbaceous plants taxa grows. The upper limit of the zone is indicated by a decline in AP and an abrupt increase of Poaceae pollen. The AP value reaches 94.9%, and Pinus dominates again (72.5%). The upper boundary of the zone is denoted by a drop in AP and an increase in Artemisia and Poaceae. The AP share slightly rises (83.5%), but Betula pollen dominates (57.3%). The rise in Pinus marks the boundary of the next zone. The AP content climbs to 75.6% in this zone. Pollen of Pinus prevails (46.7%). Artemisia and Poaceae descend, and the diversity of herbs falls. The limit of the next zone is marked by an increase in Betula. In this zone, herbs and dwarf shrubs (NAP) are the prevalent components in the spectra. The AP value is higher than 50% in only two samples, and in the (continued on next page)

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Table 1 (continued ) Local pollen assemblage zones L PAZ

Depth [cm]

K10 Pinus

740e790

K9 Betula

800e870

K8 Poaceae-Artemisia-Juniperus-Salix

880e915

K7 Pinus-Picea

925e960

K6 Picea-Abies-Carpinus

965

K5 Carpinus-Alnus-Corylus-Tilia

975e995

K4 Corylus-Tilia-Alnus

1010e1020

K3 Quercus

1030

Description of local pollen assemblage zones L PAZ remaining samples, it remains lower than 40%. The share of the Betula pollen, after a distinct fall in the preceding zone, increases to approximately 20%. The Pinus value is less than 10%. Pollen of Juniperus and Salix occur. There is a growth in Artemisia (max. 33.6%), Poaceae (20.8e34.6%) and Cyperaceae. The is a large variety of herbaceous taxa (Helianthemum, Rumex acetosella, R. acetosa, Pleurospermum, Polygonum viviparum, Sanguisorba officinalis, Plantago maritima s. str., Linum catharticum t., Ranunculus acris t., Caryophyllaceae and Chenopodiaceae). The upper limit of the zone is marked by a growth in AP. The AP value is persistently very high (max. 99.5%), but with Pinus dominating. Only some pollen grains of Larix and Juniperus are present. IN the upper part of the zone, there is a rapid fall in the AP content accompanied by a growth in the Artemisia value. A sharp decrease in AP indicates the boundary with the next zone. There are high values of AP (97%), and Betula dominates. A small but distinct decrease in AP is visible in the upper part of the zone, accompanied by a growth in the share of Artemisia and Juniperus pollen. The top of the zone is marked by a drop in the Betula pollen and an increase in Pinus. In the pollen composition of this zone, the AP content declines to 53.5%, particularly in Pinus (23.5%). Betula pollen values persist on the same level. The share of Poaceae rises to 24.4%. Pollen of Salix, Juniperus and Artemisia (8.4%) is present. Herbs attain greater variety, i.al. Cyperaceae, Calluna, Chenopodiaceae, Rubiaceae, Caryophyllaceae, Thalictrum, Rumex acetosella, Helianthemum, and Sanguisorba officinali. The value of Sphagnum reaches 12.5%. There is a sharp rise in Pinus (77.1%) and a slight decrease in Betula (18.6%). The conifer pollens replace those of the deciduous taxa in the preceding zones and are accompanied by increasing frequencies of Betula. The curves of Quercus, Corylus, Alnus, Carpinus and Abies disappear, and Picea drops noticeably. Herbs remain poorly represented, although only in the lower part of the zone. The continuous curves of Poaceae and Artemisia appear for a second time, which together with a significant decrease in the AP value, indicates the upper limit of the zone. The spectra are dominated by coniferous trees. The shares of Picea and Abies reach 43.9% and 4.9%, respectively, and Pinus ascends to 14.8%. Alnus continues to be frequent, but decreases in other deciduous taxa are conspicuous. The boundary with the next zone is marked by a decrease in Picea pollen and a progressive increase in Pinus. This zone is marked by the dominance of Carpinus pollen, the frequencies of which reaches 47.4%. Corylus and Alnus pollen decrease, and Tilia disappears. The upper limit of the zone is denoted by a decrease in Carpinus accompanied by an arrival and progressive increase in Picea pollen and also the emergence of Abies. The zone is distinguished by a sharp increase in Corylus together with other broad-leave tree pollen. The level of AP exceeds 99.1%. A reduction in the frequencies of Quercus gives way to the peak of Corylus, up to its maximum value of 68.9%, and Alnus pollen. Tilia makes its first appearance and reaches its maximum value (14.9%). A decline in Corylus pollen together with an increase in Carpinus characterize the upper zone boundary. This zone is represented by a single spectrum dominated by Quercus. The AP content reaches 93.4%. The maximum value of Quercus is 59.4%, whereas Pinus and Betula pollen occur in low

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Table 1 (continued ) Local pollen assemblage zones L PAZ

Depth [cm]

K2 Pinus-Betula-Quercus

1040e1090

K1 Pinus-Artemisia-Juniperus

1090e1100

Description of local pollen assemblage zones L PAZ amounts. The top of the zone is marked by a decline in Quercus and an increase in Corylus. The AP value is persistently high. The Pinus and Betula pollen content are at the same level. Artemisia and Juniperus disappear, and Quercus pollen emerges. A continuous curve of Ulmus appears and attains a maximum value of 2.7%. The upper limit of the zone is indicated by a rise in Quercus and a drop in Pinus and Betula. In this zone, the pollen spectra are dominated by trees and shrubs. The AP content is over 90%, and Pinus pollen dominates (66.9%). Betula reaches 22.4%. Additionally, Poaceae, Cyperaceae, Artemisia and Juniperus pollen are present. The low percentages of herb pollens and the low diversity of its taxa indicate a closed forest cover. The upper boundary of the zone is marked by a decrease in Pinus and a rise in Betula.

Fig. 3. Pollen percentage diagram of the Kubłowo succession according to Balwierz (Roman and Balwierz, 2010). AP e trees and shrubs, NAP e herbaceous plants, L PAZ e local pollen assemblage zones.

PAZ is named after the dominant taxa and/or taxa showing percentage maxima within the zone. The K1 to K7 LPAZ zones record the interglacial succession concurrent with the regional pollen assemblage zones of the Eemian defined for Poland by Mamakowa (1989). Thus, the next pollen zones, K8 to K18, may be correlated with the Vistulian (Table 2). The Kubłowo pollen sequence shows, between the Eemian Interglacial and the last glaciation, four cold intervals that are interpreted as stadial phases: Herning, Rederstall, Schalkholtz and Ebersdorf (Behre and Lade, 1986; Behre, 1989). These are €rup, separated by three temperate intervals (interstadials): Bro Odderade and Oerel. The Kubłowo sequence, when correlated with the Western European chronology, agrees well with those of

Northern Germany and accordingly spans the period between 126,000 and 55,000 years BP (cf. Roman et al., 2014; Marks et al., 2015). The organic sediments from pollen zone K14, which is related to the upper part of the Odderade Interstadial, have been radiocarbon dated (Roman and Balwierz, 2010, Figs. 2 and 3). Their age has been determined to be 42,700 ± 3,000 14C BP and older than 45,000 BP. However, at the Oerel site, the top of the Odderade deposit has been dated to 60,800 þ 2,300/1.800 BP 14C BP (Behre and Plicht, 1992). The first dating from Kubłowo is thus much younger than expected, however, difficult to interpret, especially when contamination by a recent plant material has been excluded, but other factors, such as long-term sample storage or sample preparation have to be

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Table 2 Correlation of local pollen assemblage zones (L PAZ) from Kubłowo succession (cf. Roman and Balwierz, 2010) with regional pollen assemblage zonation (R PAZ) schemes for Western Europe (Menke and Tynni, 1984; Behre and Lade, 1986; Behre, 1989) and Poland (Mamakowa, 1989). Kubłowo L PAZ (Roman and Balwierz, 2010) K18 PoaceaeArtemisiaCyperaceae K17 PinusPoaceae K16 PoaceaeArtemisiaPinus K15 PinusPoaceaeArtemisia K14 Pinus-Betula K13 Betula-Pinus K12 Pinus-Betula K11 ArtemisiaPoaceaeJuniperus K10 Pinus K9 Betula K8 PoaceaeArtemisiaJuniperus-Salix K7 Pinus-Picea K6 Picea-AbiesCarpinus K5 CarpinusAlnus-CorylusTilia K4 Corylus-TiliaAlnus K3 Quercus K2 Pinus-BetulaQuercus K1 PinusArtemisiaJuniperus

Poland R PAZ (Mamakowa, 1989)

EV4 Pinus-Betula

Western Europe chronostratigraphy (Menke and Tynni, 1984; Behre and Lade, 1986; Behre, 1989)

MIS

WP III

Ebersdorf Stadial

3

WP II

Oerel Interstadial

WP I

Schalkholz Stadial

WE IV

Odderade Interstadial

Plenivistulian

Vistulian Glaciation (Weichselian)

4

Early Vistulian (Weichselian)

5a

EV3 Gramineae-Artemisia-Betula nana WE III

Rederstall Stadial

5b

EV2 Betula-Pinus

€rup Interstadial Bro

5c

EV1 Gramineae-Artemisia-Betula nana WE I

Herning Stadial

5d

E7 E6

Pinus Picea-Abies-Alnus

E III

Late

E5

Carpinus-Corylus-Alnus

E II

Middle

E4

Corylus-Quercus-Tilia

E3 E2

Quercus-Fraxinus-Ulmus Pinus-Betula-Ulmus

El

Pinus-Betula

WE II

Eemian interglacial

5e

EI

Early

considered as likely contamination sources (cf Wohlfarth et al., 1998). The second dating is reliable, although scarcely significant. Pollen stratigraphy was essential to establish the timing of environmental changes obtained from other methods. 4.2. Cladocera data and environmental indications The subfossil Cladoceran fauna of deposits from the Kubłowo paleolake are represented by 25 species that belong to four families: Bosminidae, Chydoridae, Sididae and Daphnidae (Fig. 4). Most of the remains belonged to the families of Chydoridae and Bosminidae, and two species of Chydoridae, Alona affinis and Camptocercus rectirostris, which represent a group that inhabited a littoral

zone in clear and calm water, depending on presence of aquatic € ssner, 1972), were dominant. In the deposits, the plants (Flo ephippial eggs of the Daphnia pulex group also have been identified. During the coolest period of the Kubłowo paleolake, the dominant species became Chydorus sphaericus, which replaced Alona affinis and Camptocercus rectirostris. Considering the age of the remains above 128,000 years BP (Shackleton and Opdyke, 1976), the Cladocera remains were well preserved and abundant. Ten faunal zones (Cladocera Assemblage Zone e CAZ) were identified (Fig. 4) based on quantitative and qualitative analyses supported by numerical analyses. Characterization of Cladocera Assemblage Zones (CAZ) distinguished in the Kubłowo profile are presented in Table 3.

Table 3 Characterisation of Cladocera Assemblage Zones (CAZ) distinguished in the Kubłowo profile (by M. Niska) Symbol of zones

Depth [cm]

Main features of CAZ

CAZ e I

1050e1017

CAZ e II

1017e980

Identification of 8 species; total number of individuals of Cladocera is around 400 ind./cm3 of sediments; predominant species are: Acroperus harpae, Fig. 5), Chydorus sphaericus, Eurycercus lamellatus, Sida cristalina; there additional occur: Monospilus dispar and Camptocercus rectirostris. At the beginning of this zone, the biodiversity is low (0.8) and then increases. Increase of number of species to 14 and number of individuals to 1400 ind./cm3 of sediments (Fig. 5); a participation of occurring species is aligned; there

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9

Table 3 (continued ) Symbol of zones

Depth [cm]

CAZ - III

980e922

CAZ e IV

922e885

CAZ e V

885e805

CAZ e VI

805e752

CAZ e VII

752e690

CAZ e VIII

690e620

CAZ e IX

620e522

Main features of CAZ occur: Acroperus harpae, Alona affinis, small Alona, Camptocercus rectirostris, Pleuroxus uncinatus, and Pleuroxus trigonellus (Fig. 4); Biodiversity index is high e 1.0 (max 1.2). The best conditions for the zooplankton development; progressive increase of number species to 18 and frequency of Cladocera remains up to 8000 ind./cm3 of sediments (Fig. 5); predominant: Alona affinis (3200 ind./cm3 of sediments), Camptocercus rectirostris (3900 ind./ cm3), numerous: Pleuroxus uncinatus and Bosmina coregoni; a high frequency of species preferring a high content of nutrient in water: Pleuroxus uncinatus, Leydigia acanthocercoides, Graptoleberis testudinaria; in the middle part of this zone, more open water species: Bosmina coregoni, Bosmina longirostris and Daphnia pulex-group occur. In this zone, two short periods of lower Cladocera frequency (at the depth 965 and 940 cm) are noted; the biodiversity is a little lower e 0.8. Deterioration of living conditions due to progressive cooling of the climate. In this zone only 8 species of Cladocera are indentified (Fig. 4); an attendance of the Cladocera individuals significantly decreases (max 400 ind./cm3 of sediments, Fig. 5); only cold water tolerant species occur: Alona Affinis, Acroperus harpae, Eurycercus lamellatus and eurybiotic Chydorus sphaericus. Improving the conditions of zooplankton development; high frequency of zooplankton (above 1660 specimens/cm3 of sediments); identification of 17 species of Cladocera; a presence of cladoceran fauna connected with sediments e.g. Pleuroxus laevis, Alona quadrangularis and Monospilus dispar and macrophyte e Alona affinis, Camptocercus rectirostris, Acroperus harpae; an occurrence of Chydorus sphaericus, Leydigia leydigi linked with a meso-eutrophic status; an occurrence of Camptocercus rectirostris, Graptoleberis testudinaria preferring higher temperature of water. At the end of this zone, a frequency of individuals decreases to 150 ind./cm3 of sediments and number of species drops to 5. Further improvement of living conditions; the highest number of species (20), and the highest rate of biodiversity (1.2). The number of individuals reaches over 6000 ind./cm3 of sediments; predominat: A. affinis, C. sphaericus, C. rectirostris, Sida crystallina, Alonella nana, small Alona and A. quadrangularis; significant attendance of species preferring a higher temperature of water e P. uncinatus, P. trigonallus, G. testudinaria; a presence of species Pleuroxus sp. and L. acancocercoides, C. sphearicus, Bosmina longirostris, M. dispar, Alonella exigua e connected with a higher trophy level; a significant share of the pelagic species e B. longirostris, E. coregoni. The deterioration of living conditions; a strong decline in attendance of species to 9; drop in the participation of individuals up to 300 ind./cm3 of sediments; lack of deep-water species and species with a higher thermal requirements; higher attendance only species with broad environmental tolerance e A. affinis, A. quadrangularis and C. sphaericus. Improvement of living conditions; an increase of attendance of species to 17; a number of individuals increases to 1500 ind./cm3 of sediments; a return of species with a higher thermal and nutrients requirements: C. rectirostris, G. testudinaria, M. dispar, Pleuroxus and open water specie e B. coregoni, B. longirostris, Daphnia pulex e group; high biodiversity e 1.0 (max. 1.2). (continued on next page)

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Table 3 (continued ) Symbol of zones

CAZ e X

Depth [cm]

Main features of CAZ Further improvement of living conditions; a significant increase in attendance of individuals to about 2000 ind./cm3 of sediment; a higher variety of species composition e 20; predominant: C. sphaericus (Fig. 4), C. rectirostris, A. harpae, small Alona, A. affinis; presence of species from all habitats group; high level of biodiversity index (1.0 max. 1.2). Visible deterioration of living conditions; a decline in attendance of Cladocera remains between 0 and 770 ind./cm3 of sediments; a reduce of amount of species, initially to 9 and then to 4 species; a significant attendance of only broad tolerant species e.g.: C. sphaericus; above a depth of 490 cm there are almost no Cladocera remains; above depth of 425 cm, an improvement in the living conditions and presence of: Bosmina longirostris and B. coregoni, C. sphaericus; biodiversity index declines to a maximum of 0.5.

522e405

4.3. Isotopic and selected geochemical data 4.3.1. TOC, TN, and C/Natomic The lowest sediments (below 1025 cm) are characterized by a variable concentration of organic carbon in the range of 2e10% and a nitrogen content below 0.5% (Fig. 5). The C/Natomic ratio oscillates from 18 to 24. Then the contents of both TOC and TN rapidly increase to 20% and 1.5%, respectively, but the C/N atomic ratio decreases to 16 (at a depth of 1025 cm). This increase is followed by a drop to 9% and 0.6%, respectively. The C/N atomic ratio rises to 20. Next, in the deposits at depths ranging from 1025 to 925 cm, the Corg content varies from 20 to 28%, and the N content oscillates from 1.3 to 1.8%. The C/N atomic ratio oscillates around approximately 18. In the overlying deposits (to a depth of 880 cm), the TOC and TN rapidly decrease to 2% and 0.1%, respectively. The C/N atomic ratio oscillates from 16 to 23. At depths of 820e875 cm, the contents of both TOC and TN rapidly increase to 37.6% and 2%, respectively, and the C/N atomic ratio first rises to 34 and then decreases to 20. These increases are followed by drops to 11% and 0.6%, respectively. The C/N atomic ratio oscillates approximately 19 (depths of 805e815 cm). In the overlying deposits, there are again increases in the organic carbon and nitrogen contents (to a depth of 760 cm). The TOC and TN attain the highest values of 38.5% and 2.8% (a maximum of TN), respectively. The C/N atomic ratio systematically drops to 16. From depths of 755 to 600 cm, the TOC and TN achieve the lowest values of 1.6% and 0.1%, respectively (minimum at a depth of 730 cm). The C/N atomic ratio initially declines to a range approximately 13 (depth of 695 cm) and then systematically increases to 19. The high peaks of both TOC and TN occur (23% and 1.5%, respectively) only at a depth of 625 cm. Above 600 cm, the contents of TOC and TN rapidly increase to 40.3% (maximum of TOC) and 2.5%, respectively, and their values then systematically drop. The C/N atomic ratio first rises to 23 and then decreases to 20. From a depth of 545 cm, the contents of TOC and TN rapidly decline within the ranges of 5e8% and 0.3e0.6%, respectively. At depths above 495 cm, these values remain at their lowest levels, 1.1e2.2% and 0.1e0.15%, respectively. The C/N atomic ratio decreases to 13. 4.3.2. d13 and d15N In the lowest sediments (below 1330 cm), the d13C values slightly oscillate from 26.7 to 27.8‰, and the d15N values change from 1.4 to 2.3‰ (Fig. 5). At depths of 1030e1025 cm, the values of d13C increase to 24.8‰, and the d15N values are constant (1.1‰). Then, the values of d13C rise from 27.4 to 21.8‰, but d15N

irregularly drops from 2.9 to 0.2‰ (depth: 1020e970 cm). In the overlying deposits (depth of 965e920 cm), d13C continues to increase and reaches high values of -19-18‰, and d15N fluctuates from 0.8e1.1‰. Next, the d13C values systematically decline to approximately 28‰ (the lowest values). d15N shows an opposite trend and increases to 4‰ (depth: 915e870 cm). From 870 to 800 cm, a systematic increase in the d13C values to 22.5‰ and a fluctuation of d15N from 1.5 to 3.7‰ are observed. Then, (depth: 795e750 cm), the carbon isotopic values increase and attain a maximum of 12.9‰. In this interval, d15N first rapidly drops to 1.6‰ (minimum of values) and then rises to ca. 1‰. At depths of 730e680 cm, d13C firstly increases to ca. 20‰ and then slightly decreases to 22.6‰. d15N again rises to ca. 4‰. From 675 to 635 cm, d13C first rapidly drops to 25.8‰ and then increases to 16.6‰. The d15N values drop to ca. 2‰. Next (depth: 630e495 cm), d13C fluctuates between 24.8 and 21.7‰, and a systematic increase in the d15N values from 0.7 to 3.2‰ is noted. Above 485 cm, the values of d13C decrease and oscillate around ca. 27‰. The d15N values are constant at ca. 5‰ (the highest values). 5. Environment of the Kubłowo paleolake during the Eemian-Vistulian The ecological preferences of Cladocera were used to reconstruct the changes in water level, trophic status and temperature of the lake water in the study paleolake. The isotopic and geochemical data were used to help describe the characteristics of the organic sediments and define the source of the organic matter. 5.1. Pre-early Eemian In the final phase of the Wartanian Stadial (Late Saalian), deglaciation of the area took place (Baraniecka, 1979; Roman, 2010). Many lake basins e.g., in Kubłowo was formed, and mineral deposition started. 5.2. Early Eemian e K1 (Pinus-Artemisia-Juniperus) and K2 (PinuseBetula-Quercus) L PAZs During the early Eemian, an open pine forest with a distinct birch share (K1 L PAZ) occurred in the neighborhood, and the scarce open terrains were grassy. The climatic amelioration led to a

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Fig. 4. Diagram of the absolute number of Cladocera individuals and total number of Cladocera specimens, species and diversity in the sediments of Kubłowo profile (M. Niska). CAZ e Cladocera assemblage zones, L PAZ e local pollen assemblage zones (acc. Roman and Balwierz, 2010). Explanations of Lithology e see Fig. 2.

distinct reduction in heliothytes, which were replaced by a dense birch-pine forest (K2 L PAZ) (Roman and Balwierz, 2010). At the beginning of the early Eemian Interglacial, only one species of Cladocera e Eurycercus lamellatus was identified (CAZ I, Fig. 4). This species is characterized by its cold-water tolerance and inhabits the littoral zone in clear water (Poulsen, 1944). At the end of the early Eemian, above a depth of 1045 cm, species that prefer a low content of nutrients and are tolerant to cold water, such as Acroperus harpae, Eurycercus lamellatus, Sida crystalina Alona affinis (Whiteside, 1970) and ubiquitous Chydorus sphaericus (Alhonen, 1970), occurred. Additionally, E. lamellatus, S. crystallina and € ssner, 1972). The A. affinis, point to the existence of macrophytes (Flo species composition was characteristic for a littoral zone, and the

absence of open water species suggest a low water level in this lake. The presence of Monospilus dispar in the sediment confirms a sandy bottom. The trophy was at the oligo/mesotrophic level. At the time, the conditions in this reservoir were inconvenient for the development of zooplankton. In the early Eemian Interglacial, organic sediments began to be deposited in the Kubłowo paleolake. The concentrations of organic carbon and nitrogen were low, ca.10% and 0.5%, respectively (Fig. 5). The amount of organic matter reflected a very low primary productivity and the development of macrophytes in the lake. The C/ Natomic ratio, which is above 20, indicates a higher plant component during the period. The deposits were characterized by d13C org values around 27‰ and d15N values of ca. 1.5‰ (Fig. 5). The low

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Fig. 5. Results of selected geochemical and carbon and nitrogen isotopic analyses of the Kubłowo palaeolake sediments (J. Mirosław-Grabowska). R PAZ e regional pollen assemblage zones (acc. Mamakowa, 1989), L PAZ e local pollen assemblage zones (acc. Roman and Balwierz, 2010), TOC eTotal Organic Carbon, TN e Total Nitrogen. Explanations of Lithology e see Fig. 2.

d13Corg values may reflect the dominance of C3 plants in the catchment. These isotopic and geochemical data suggest that the organic matter present in sediments at that time were predominantly of terrestrial origin. 5.3. Middle Eemian e K3 (Quercus), K4 (Corylus-Tilia-Alnus), and K5 (Carpinus-Alnus-Corylus-Tilia) L PAZs The interglacial climatic optimum started with an oak forest with a small admixture of elm and an ash e oak phase (K3 L PAZ). The progressing warming resulted in changes in the main tree stands. The dominant species e oak, elm and ash trees e were replaced by a multi-species deciduous forest with hazel and linden trees, as well as alders (K4 L PAZ) that prevailed in wet habitats. Next, hornbeam began to spread and at the end of K5 L PAZ became dominant (Roman and Balwierz, 2010). At the beginning of the middle Eemian (K3 L PAZ), species that preferred a low content of nutrients and were tolerant to cold water dominated the Kubłowo paleolake. Next, with the changes in climate conditions, the zooplankton composition began to alter. Cladocera species, which preferred warmer water, appeared for the first time (Camptocercus rectirostris, Pleuroxus uncinatus and Pleuroxus trigonellus), suggesting slightly warmer water conditions (CAZ II) (Poulsen, 1944). From the hazel phase (K4 L PAZ), there was also an increased frequency of species that preferred water richer in nutrients e Leydigia acanthocercoides, Chydorus sphaericus and, as

mention before, P. uncinatus (Frey, 1958). The dominant species were characteristic of a littoral zone, but at the middle of CAZ II (above 1005 cm), the share of planktonic taxa (Bosmina (E) coregoni) started to increase, which suggests an increase in water level. The increasing temperature and water level and an increase in nutrients and food (phytoplankton) in the lake resulted in greater biodiversity, which reflected an improvement in the living conditions in the lake. In the organic deposits in the Kubłowo paleolake, the concentrations of TOC and TN fluctuated and significantly increased (to ca. 25% and 1.3e1.8%, respectively e Fig. 5). Firstly, C/Natomic dropped and then remained at a level of approximately 18 (Fig. 5). The lower C/Natomic ratios indicate an increased share of algae in the organic matter. After the initial decrease (below 27‰), d13Corg systematically increased to ca. 22‰ (Fig. 5). Initially, d15N irregularly rose to ca. 3‰ and then rapidly dropped to 0.2‰ (Fig. 5). The systematic increase in the d13Corg values likely reflects an increase in photosynthesis, but the high d15N values may suggest a high trophic status (Brenner et al., 1999). 5.4. Late Eemian e K6 e Picea-Abies-Carpinus and K7 e Pinus e Picea L PAZs A decline in the climatic optimum was indicated by the preeminence of a hornbeam forest. Gradual cooling led to withdrawal of the thermophilous trees and the appearance of spruce, fir

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and yew e the spruce phase (K6 L PAZ). Next, the spread of pine with some spruce and initially also fir (K7 L PAZ) marked a decline of the Eemian Interglacial (Roman and Balwierz, 2010). The zooplankton composition responded to climatic and environmental changes. The climatic conditions during the spruce phase (K6 L PAZ) were optimal for the development of zooplankton (CAZ III), which resulted in increased number of the Cladocera species and respective individuals, including the thermophilous species (C. rectirostris, P. trigonellus, P. uncinatus and G. testudinaria). A higher attendance of species Leydigia acanthocercoides, Alona rectangula and Pleuroxus uncinatus suggests a higher content of nutrients in the lake water (Korhola, 1990). In the middle part of this period (approximately 950 cm), there were a little more species that preferred open water: Bosmina (E) coregoni, Bosmina longirostris. This faunal set suggests a rise in the lake water level. An increase in attendance such species as: C. rectirostris, small Alona, Alona affinis, and Alonella excisa probably indicates a growth of macrophyte zone. At the end of the pine phase (K7 L PAZ), there was a reduction in both number of the Cladocera species and the number of specimens of individual species (the end of CAZ III). During the late Eemian Interglacial, there were two short periods with a lower frequency of Cladocera (depths of 965 cm and 940 cm e CAZ III), which could have been due to fluctuations in the thermal conditions in the lake and the occurrence of a short cooling and/or shallowing. The Cladocera composition reflects the occurrence of the most unfavorable conditions for zooplankton development in this lake, probably due to a progressing cooling. In this period, the concentrations of TOC and TN were still high: 28.6% and 1.8%, respectively (Fig. 5). The C/Natomic slightly increased (to 20 e Fig. 5). The d13Corg values systematically rose to the Eemian maximal values (17.8‰). The opposite trend was observed for the d15N values; they decreased and varied between 0.4 and 1‰ (Fig. 5). The more positive d13Corg, as well as high TOC values, generally correspond to the marshy sediments, implying the shrinkage of the water body and intrusion of the terrestrial and near shore aquatic plants (Wei et al., 2010). The reduce of water body is also suggested by a high abundance of cladocerans, especially preferring macrophytes (Pleuroxus sp., Alona affinis, Camptocercus rectirostris) and a higher C/Natomic ratio refers to dense macrophyte in littoral (Tyson, 1995). The end of this period was marked by a drop in the organic carbon and nitrogen contents. This phenomenon corroborates the reduction of primary production as well as vegetation that was likely associated with cooling. 5.5. Early Vistulian 5.5.1. Herning Stadial e K8 e Poaceae-Artemisia-Juniperus-Salix L PAZ At the onset of the Vistulian, the forest, particularly the pine forest, began to retreat, and open vegetation communities spread (K8 LPAZ). Shrub communities, mainly with juniper and willow, grew, and meadow-type grassland communities dominated. In open settlements and leached soils, heathers developed (Roman and Balwierz, 2010). Progressive cooling in the Herning Stadial resulted in the habitation of the Kubłowo paleolake by only cold water tolerant species, Acroperus harpae, Eurycercus lamellatus and eurybiontic Chydorus sphaericus, and a disappearance of species preferring warm water (CAZ IV, Fig. 4). These species, which tolerate cold water and a lower status of trophy (Harmsworth, 1968), confirm the unfavorable environmental conditions for zooplankton development. In this period, the composition of the organic matter significantly changed. The concentrations of TOC and TN rapidly decreased to minimal values (2% and 0.2%, respectively e Fig. 5). The C/Natomic ratio fluctuated between 16 and 23. The values of

13

d13Corg dropped to ca. e26‰. The opposite trend was observed for d15N. These values increased to 4‰ (Fig. 5). The geochemical and isotopic data suggest a reduction in primary production and the sporadic input of terrestrial organic matter into the lake (Meyers s, 1999). The increase in d15N suggests lower niand Lallier-Verge trogen availability in the surface waters (Talbot and Laerdal, 2000). € rup Interstadial e K9 e Betula and K10 e Pinus L PAZs 5.5.2. Bro At first, birch was dominant and resulted in a significant reduction in the herbaceous communities (K9 L PAZ). The end of the birch phase was a short-duration, cool climatic event that was expressed by the growth of herbaceous communities and shrubs with Juniperus. Then, dense pine forests, with larch and spruce, spread (K10 L PAZ). At the end of the pine phase, a subsequent cooling occurred. Similarly to the former one, it was characterized by the increased growth of the herbaceous communities, including heliophytic sage-brush, followed by a short-term return of the pine forests. The pine phase rapidly terminated (Roman and Balwierz, 2010). €rup Interstadial caused improving Climate warming in the Bro conditions for zooplankton development e CAZ V (Fig. 4). The cladoceran succession is divided into two parts. The first part is described by a high frequency of zooplankton (above 1660 specimens/cm3 of deposits, CAZ V) and an occurrence of 17 species of Cladocera. The characteristic feature of this period is the presence of cladoceran fauna connected with sediments, e.g., Pleuroxus laevis, Alona quadrangularis and Monospilus dispar, and a macrophyte zone (Alona affinis, Camptocercus rectirostris and Acroperus harpae), and there are no Cladocera from the open water zone. This Cladocera composition suggests a lowering of the water level. The occurrence of Chydorus sphaericus and Leydigia leydigi indicates a high trophic status meso/eutrophy in the lake. The moderate thermal conditions were conducive to the development of species that preferred warmer water (Camptocercus rectirostris, Graptoleberis testudinaria) as well as species that tolerated lower water € rup Interstadial finished with temperatures. The first part of the Bro a significant drop in the abundance of species and specimens of Cladocera, which suggests a deterioration in the living conditions (approximately 810 cm). In the second part of this interstadial (CAZ VI), the frequency of both species and specimens of Cladocera noticeably increased. In this stage, the highest number of species (20) and also the highest rate of biodiversity (1.2) occurred (Fig. 4), which can be interpreted as a restoration of the lake. This period was one of two in the whole profile with the highest attendance of remains in the sediments. During this period, A. affinis, C. spaericus, C. rectirostris, Sida crystallina, Alonella nana, Alona rectangula and A. quadrangularis were dominant. A significant turnout also reached the species that preferred higher water temperatures (P. uncinatus, P. trigonallus and G. testudinaria), which might indicate favorable thermal conditions for the more demanding species. The presence of species of the genus Pleuroxus sp. and L. acancercoides, C. sphearicus, Bosmina longirostris, M. dispar and Alonella exigua suggests a higher trophy e at the level of a meso/ eutrophic state (Adamska and Mikulski, 1968). The occurrence of G. testudinaria is also connected with a higher content of decaying plants in water e a high content of carbon (Diugan, 1992; Niska, 2008). This phenomenon usually happens after periods of low water levels and the development of peat-forming plants. In this stage, a significant participation of a pelagic species group e B. longirostris and E. coregoni was also reported, which might indicate an improvement in the living conditions that was also associated with a higher water level. At the period, the geochemical and isotopic data diversified into two phases, highly correlated with the Cladocera zones: CAZ V and CAZ VI and local pollen zones (K9 and K10 L PAZs). Since the

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beginning of the period, the content of TOC rapidly increased to above 38%, and TN systematically rose to 2.8%. At depths of 810e815 cm, a short time, rapid decreases to 11% and 0.6%, respectively, occurred (Fig. 5). The C/N atomic ratio first reached a double maxima (of 34 and 31) and then steadily decreased (to 17e18). The lower deposits (depth: 800e875 cm) were characterized first by a rapid drop in the d13C org values to minimum values (ca. 28‰, at a depth of 870 cm) and then an increase of 4e6‰. d15N oscillated between 1.5 and 3.7‰. In the upper deposits (depth: 750e800 cm), d13C org rose quickly to 14‰ and then started to decrease. d15N exhibited an opposite trend. An initial drop to minimum values (1.6‰, at depth: 790 cm) was followed by systematic increase (to ca. 1‰). A significant, rapid increase in TOC and the highest values of C/N atomic suggest an initial significant supply of organic matter of terrestrial origin into the lake. Next, with an improvement in the thermal conditions, an enhancement in the primary productivity probably occurred (the lower values of d13C org are typical for freshwater algae e Leng et al., 2005). The systematic increase in the d13Corg values likely reflects an increase in photosynthesis, because at periods of enhanced productivity, the aquatic plants preferentially take up of 12C during photosynthesis (Leng and Marshall, 2004). The high d15N values suggests a high trophic status (Brenner et al., 1999) and the Cladocera compostion also the high trophic state confirms (presence of Ch. sphaericus, M. dispar and Alonella exigua). The results of proxies are similar to the Eemian period, thus environmental conditions would be similar to interglacial ones.

5.5.3. Rederstall Stadial e K11 e Artemisia-Poaceae-Juniperus L PAZ During the Rederstall Stadial, climatic cooling occurred. Probably only at the beginning of this stadial, there were patches of pine and birch forest. Then, a very low share of pine pollen suggests that pines were absent around the Kubłowo paleolake. The heliophytic communities and mainly grasses and sage-brushes were dominant at that time and were much more numerous than in the Herning Stadial. Juniper and willow were also common. At the end of the stadial, birch significantly spread. In the Rederstall Stadial, two stages of Cladocera development were also observed. In the first part (CAZ VII), there was a strong decline in the attendance of individuals and species of Cladocera (7 species, Fig. 4). A lowering of the water level in the reservoir is suggested by the absence of deep-water zone species. The lack of species with higher thermal requirements indicates climatic cooling. The dominant species was ubiquitous C. sphaericus. In the second part (above 680 cm, CAZ VIII), a slight improvement in the living conditions was observed. The turnout of species increased (17 species, Fig. 4), but the attendance of individuals was low. In the Kubłowo paleolake, the species of the deep-water group and thermophilics reappeared. At the time, the low organic carbon and nitrogen contents occurred (to 3e5% and ca. 0.3%, respectively). Only at depths of 625 cm, a rapid increase to 23% and 1.5%, respectively, was noted (Fig. 5). Initially, the C/N atomic ratio decreased to 13 and then rose to 18. Large fluctuations in the values of d13C org were observed (from 26‰ to 16.6‰) and finally dropped to ca. 24‰. Initially, the d15N values remained at the same level of ca. 4‰ and then decreased to ca. 2‰. The low values of TOC and C/N atomic suggest a low primary production and aquatic vegetation in the lake during this period. The appearance of planktonic Cladocera especially Daphnias, and decreased number of plant associated species corroborates a low primary production and suggests an oligotrophic condition.

5.5.4. Odderade Interstadial e K12 e Pinus-Betula, K13 e BetulaPinus, K14 e Pinus-Betula and K15 e Pinus-Poaceae-Artemisia L PAZs During the Odderade Interstadial, a short spread of pine (K12 L PAZ) followed by a short dominance of birch (K13 L PAZ) is observed. The pine returned later to the investigated area (K14eK15 L PAZs). The constant appearance of heliophytic plants indicates that the forests were not very dense. There were also open site communities, which were especially most numerous at the onset and towards the end of the interstadial. Birch was dominant at the beginning of the interstadial, and later its share decreased (Fig. 3). In this interstadial, a substantial improvement in the living conditions took place at the Kubłowo paleolake. The attendance of specimens (approximately 2500 ind./cm3 of sediments) and species (max. 21 species, Fig. 4, CAZ IX) was higher than expressed by the earlier pollen. In the paleolake, species from all groups, including those that required or preferred deep water, higher temperatures and higher concentrations of nutrients, were represented. This interstadial was the last period of the occurrence of convenient conditions for the development and reproduction of zooplankton and the presence of thermophilic species. In this period, the contents of organic carbon and nitrogen were at first very low. Then, especially during the pollen zone (K14 L PAZ), they abruptly increased to 40% and 2.5%, respectively (Fig. 5). The C/N atomic ratio first oscillated from 16 to 23 and then oscillated approximately 20. The d13C org values initially changed from 24.8 to 21.7‰, and next they remained at a level of 23e24‰. The d15N values fluctuated approximately 2‰ (Fig. 5). The C/N atomic ratios indicate a higher plant component during the period. The isotopic and C/N atomic values suggest a short-term supply of terrestrial plants into the reservoir. 5.6. Plenivistulian 5.6.1. Schalkholz Stadial e K16 e Poaceae-Artemisia-Pinus L PAZ This period was a mainly forestless period around the Kubłowo paleolake (Fig. 3). However, there could have been small tree stands or isolated trees. Herbs, mainly grasses, were predominant. The share of Artemisia was much lower than in the Rederstall Stadial, which was likely due to the influence of the continental climate or higher groundwater levels, and the more widespread moist and wet locations possibly indicating grassland communities (Roman and Balwierz, 2010). The beginning of Middle Vistulian (Lower Plenivistulian, Schalkholz Stadial) was associated with a gradual disappearance of the remains of Cladocera in the sediments (CAZ X). Initially, the remains of five species were identified and between them C. gibbus very tolerating towards electrolyte rich water. Then only C. sphaericus was found in the reservoir to a depth of 460 cm. There was a pause in the succession of Cladocera caused by adverse climatic conditions. At that period, the contents of organic carbon and nitrogen were low (2% and ca. 0.15%, respectively e Fig. 5). Initially, the C/N atomic ratio oscillated between 17 and 19. d13Corg systematically decreased to 27‰. The d15N values increased and reached their maximum of 5.3‰ and then remained at the same level of ca. 5‰ (Fig. 5). The very low TOC and TN reflect a low primary production and water vegetation in the lake during this period. The increase in d15N suggests lower nitrogen availability in the surface waters (Talbot and Laerdal, 2000). The geochemical data corroborated the unfavorable stadial conditions. 5.6.2. Oerel Interstadial e K17 e Pinus-Poaceae L PAZ During this interstadial, the share of trees, mainly pines, grew (Fig. 3). The amelioration of the climate probably removed the

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northern limit of the forest, however the landscape around the Kubłowo paleolake could have remained forestless. Herbaceous communities still dominated this area (Roman and Balwierz, 2010). During the Oreal Interstadial (CAZ X), Bosmina (E) coregoni, B. longirostris and C. sphaericus returned. These species are found in lakes with unfavorable thermal and edaphic conditions and in reservoirs in the absence of macrophytes towards the end of a lake's existence (Bjerring et al., 2009). This reappearance of Cladocera suggests a slightly higher water level and probably the development of zooplankton. At that time, the contents of organic carbon and nitrogen were extremely low (ca. 1% and ca. 0.1%, respectively e Fig. 5). The C/N 13 atomic ratio dropped to 13. d C org systematically decreased 15 to 27.6‰. The d N values remained at the same level of ca. 5‰ (Fig. 5). The geochemical data reflect a continuation of unfavorable conditions. 5.6.3. Ebersdorf Stadial e K18 e Poaceae-Artemisia-Cyperaceae L PAZ During the Ebersdorf Stadial, only the topmost sediments accumulated. At that time, herbaceous plants reached a maximum proportion in the entire profile. Thus, around the Kubłowo paleolake, no trees were present. A high percentage of Poaceae indicates a dominance of fresh/moist grassland settlements, but the simultaneous presence of species characteristic of dry settings suggests that at that time tundra and steppe communities could have cohabited (Roman and Balwierz, 2010). The organic deposits accumulated in the Ebersdorf stadial were only palynologically analyzed. The lack of sediments made it impossible to investigate Cladocera. The very small amount of organic matter did not allow for geochemical and isotopic analyses. 6. Conclusions The unique character of the paleolake at Kubłowo is related to the very long, uninterrupted sedimentation that occurred during the entire Eemian Interglacial (MIS 5e), Early Vistulian (MIS 5a-d) and for part of the Middle Vistulian (MIS 4-3). The reconstruction of the environmental changes at Kubłowo and its surroundings show the short climate fluctuations and a detailed picture of the natural conditions prevailing in central Poland during the period from approximately 126,000 to 55,000 years BP. The paleoenvironmental data have been into the context of the last interglacial and early glacial conditions in the central Europe. The pollen analysis has provided a schema of the biostratigraphy, vegetation development and climatic changes during the Eemian Interglacial and the Early Vistulian Glaciation in the Kujawy region (central Poland). The Cladocera and geochemical data illustrate the environmental conditions that occurred in the lake and confirm the climatic changes postulated using the pollen data. The comparison of the Cladocera and the carbon and nitrogen stable isotopes data enabled the reconstruction of the development of the paleolake in relation to the pollen record of the lake-bog sediments. Based on the results of multi-proxy analysis, four lacustrine stages linked with a higher water level and three boggy stages (low water level) of evolution of the Kubłowo paleolake have observed (Fig. 6). The characteristics of the environmental conditions were summarized in Fig. 6. Vegetation development around the Kubłowo paleolake was connected with climatic conditions, akin to that of other Eemian sites from the European lowlands (Zagwijn, 1996; Aalbersberg and  nas and Grigiene, _ 2012) as well in Litt, 1998; Kühl et al., 2007; Satku  wek and Łanie˛ ta site (Janczyk-Kopikowa, 1997; the nearby Ruszko Balwierz and Roman, 2002). At the beginning of the Eemian Interglacial, pine and birch dominated, and a boreal forest spread. A

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further warming of climate was demonstrated by the appearance of deciduous trees such as oak, elm and lime. During the Eemian optimum, thermophilous deciduous forests, dominated first by hazel and then by hornbeam, occupied the area. The post-optimum cooling was expressed by a deterioration of the deciduous forest and the expansion of boreal coniferous trees. The further regression of the forest communities and the domination by herbaceous plants was caused by the approaching continental ice sheet e the Vistulian Glaciation. During the Early Vistulian, two cold and two warmer periods alternately occurred. In the stadials, the herbaceous plants with Artemisia and Poaceae dominated. During the interstadials, the shares of pine and birch increased, and forest communities prevailed. In the Plenivistulian, two stadials separated by one interstadial have been distinguished. A low proportion of trees reflects a woodless period in the study region during the Middle Vistulian (Roman and Balwierz, 2010). The different climatic conditions recorded in the pollen diagram are also clearly visible in the lithology e the changes in the amount of organic matter observed at the boundaries between the Eemian Interglacial and the Vistulian and between the interstadials and stadials within the Vistulian e as well in the Cladocera composition, observed as changes in the number of specimens, number of species and biodiversity. Climatic warming is evident not only from the development of the vegetation but also by an increase in the primary production and an increase in the abundance and diversity of Cladocera (e.g., the Eemian optimum). Climatic cooling not only caused the disappearance of certain species of plants (e.g., deciduous trees) but also produced unfavorable conditions for the development of zooplankton and the production of small amounts of organic matter (Fig. 6). This phenomena occurred e.g. during the € rup Interstadial (Fig. 6). Bro The succession of Cladocera recorded in the sediments of the Kubłowo paleolake is unique on a European scale, especially noting the continuous function of the reservoir for more than 70,000 years. There are currently no papers that compare the record of a succession of zooplankton over such a long period of time, but there are similar but shorter profiles: Solniki (northern Podlasie, Kupryjanowicz, 2008; Mirosław-Grabowska et al., 2015) and  wek (Janczyk-Kopikowa, 1997; Mirosław-Grabowska et al., Ruszko 2009), which include the Early Vistulian (without the Plenivistulian). The variability in species and in individual attendance reflects regional climate changes, and changes in the local living conditions in the lake are also documented by the palynological, geochemical and isotopic data. The maximum of the reservoir development and the most convenient conditions for fauna development occurred in the late Eemian Interglacial (Fig. 6). The end of the Eemian Interglacial and the beginning of the Vistulian were associated with the almost complete absence of Cladocera fauna. The progressive cooling associated with the Stadial Rederstall and Schalkholz Stadials also contributed to the decline in the attendance of species and individuals of Cladocera. The remains of Cladocera identified in sediments, despite their age, were very well preserved, and identified species also occur now in Central Europe. Several paleolakes with sequences of Eemian-Vistulian sediments have been found near the paleolake at Kubłowo (eg. Balwierz and Roman, 2002; Mirosław-Grabowska and Niska, 2007; Mirosław-Grabowska, 2009; Mirosław-Grabowska et al., 2009; Roman and Balwierz, 2010). The Kubłowo paleolake differs from the other sites in the region because of its organic sedimentation. That lithology has made it possible to obtain carbon and nitrogen stable isotope records for the entire Eemian and Vistulian succession. The results of the analyses of organic matter correlated with Cladocera data have enabled a reconstruction of the varying environmental conditions and geneses of organic matter. The organic

Please cite this article in press as: Mirosław-Grabowska, J., et al., Long (MIS 5e e 3) environmental history of a paleolake in central Poland recorded in the succession from Kubłowo, Quaternary International (2016), http://dx.doi.org/10.1016/j.quaint.2016.06.027

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Fig. 6. Reconstruction of environmental conditions in the Kubłowo palaeolake. R PAZ e regional pollen assemblage zones (acc. Mamakowa, 1989), L PAZ e local pollen assemblage zones (acc. Roman and Balwierz, 2010), AP e trees and shrubs, NAP e herbaceous plants (acc. Roman and Balwierz, 2010), CAZ e Cladocera assemblage zones, TOC eTotal Organic Carbon. Explanations of Lithology e see Fig. 2.

carbon content is closely related to climatic conditions (Fig. 6). The lowest values are recorded in the records of the coldest periods during the Herning and the Rederstall Stadials of the Early Vistulian and during the Plenivistulian. The highest values occur in the re€rup cords of the Eemian post-optimum (E6-E7) and in the Bro Interstadial (Early Vistulian). Acknowledgements We would like to express our thanks to Dr. Zofia Balwierz for very interesting discussion. The study was partially financed by the Institute of Geological Sciences Polish Academy of Sciences (EEM). The research project was supported by the Foundation for Polish Science, Bridge Program 2012 e project: “Reconstruction of the development of lake environmental in the Eemian interglacial based on subfossil Cladocera (Crustacea) analysis”. Fieldworks and pollen analysis were supported by the Ministry of Science and Higher Education, project 2PO4E 02329. References Aalbersberg, G., Litt, T., 1998. Multiproxy climate reconstructions for the Eemian and early Weichselian. Journal of Quaternary Science 13, 367e390. Adamska, A., Mikulski, J.S., 1968. Cladocera remains in the superficial sediments of lakes as a typologic indicator. Z. Naukowe UMK 25. Prace Stacji Limnolog w Iławie 5, 41e48. Alhonen, P., 1970. The paleolimnology of four lakes in southwestern Finland. Annales Academiae Scientiarum Fennicae Series A III. Geologica-Geographica 105, 1e39.

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Please cite this article in press as: Mirosław-Grabowska, J., et al., Long (MIS 5e e 3) environmental history of a paleolake in central Poland recorded in the succession from Kubłowo, Quaternary International (2016), http://dx.doi.org/10.1016/j.quaint.2016.06.027