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Quaternary International 268 (2012) 9e20

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Quaternary International

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Vegetation patterns under climate changes in the Eemian and Early Weichselianin Central Europe inferred from a palynological sequence from Ustków (centralPoland)

Piotr Ko1aczek a,*, Monika Karpi�nska-Ko1aczek b, Joanna Petera-Zganiacz c

aDepartment of Biogeography and Palaeoecology, Faculty of Geographical and Geological Sciences, Adam Mickiewicz University, ul. Dziegielowa 27, 61-680 Pozna�n, PolandbDepartment of Palaeobotany and Palaeoherbarium, Faculty of Biology and Earth Sciences, Jagiellonian University, Lubicz 46, 31-512 Kraków, PolandcDepartment of Quaternary Research, Faculty of Geographical Sciences, University of Łód�z, ul. Kopci�nskiego 31, 90-142 Łód�z, Poland

a r t i c l e i n f o

Article history:Available online 7 May 20127 May 2012

* Corresponding author.E-mail address: pkolacz@amu.edu.pl (P. Ko1aczek)

1040-6182/$ e see front matter � 2012 Elsevier Ltd adoi:10.1016/j.quaint.2012.05.004

a b s t r a c t

This paper presents a fresh examination of a pollen profile from Ustków (central Poland), which afterpreliminary results was thought to span a period between the Early Eemian (MIS 5e) and the upperPleni-Weichselian (MIS 3). The newly obtained results confirmed the age assessment of the bottomseries, but revealed a much older age for the uppermost part of the profile as far back as the Rederstallstadial (MIS 5b). Palynological research showed slightly different patterns of vegetation in comparison toother sites located in Central Europe. Among them were an early Ulmus maximum coinciding with theBetula optimum in the Early Eemian, a relatively late optimum of Taxus baccata during the decline of theMiddle Eemian, and a distinct division of the older part of the Late Eemian into phases of Abies-Picea andPicea-Pinus forest domination. The Herning stadial (MIS 5d) falls into a typical bipartition reflected byheathland domination during its older part and the prevalence of Juniperus thickets and Artemisia-Poaceae steppe during its younger part, which makes this succession similar to those from northernGermany. The pollen spectra reflecting the Brørup interstadial (MIS 5c) distinctly revealed an intra-Brørup cold oscillation rarely detected in profiles from Central Europe and a period of Larix dominatedforests during the latest part of the interstadial (extraordinarily high percentages exceeding 15% of thetotal pollen sum). Local pollen taxa, together with the lithological composition of deposits, revealed thatthere was a water body in the area during the periods of the Late Saalian/Early Eemian transitioneMiddleEemian (the decline of the Tilia phase), the Herning stadial (MIS 5d), the early Brørup interstadial (MIS5c), and the Rederstall stadial (MIS 5b). Between these time-intervals a poor and/or rich fen and/or bogfunctioned.

� 2012 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction

Research carried out on profiles with series which span periodsof the InterglacialeGlacial cycle is crucial to understand patterns inthe palaeoclimate, palaeohydrology and palaeoecology of thatperiod, and they are important for building predictive climatemodels (Reille et al., 1998; Müller and Sánchez-Go�ni, 2007). Thereis nothing new in the statement that such sites are unique to thearea of Central andWestern Europe, as a result of several damagingmechanical processes these sites were subjected to during theWeichselian and subsequent Holocene. Of about 300 pollen profilesspanning the Eemian interglacial (MIS 5e) examined from the area

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of Poland (Kupryjanowicz et al., 2011), only the profile from Hor-oszki Du _ze in eastern Poland (Granoszewski, 2003) reached thestadial after the Denekamp interstadial (MIS 3). This fact promptedreexamination of the palynological analysis of the profile fromUstków (central Poland) which, after preliminary pollen analysisand radiocarbon dating, was thought to span the period betweenthe Saalian/Eemian transition and the decline of the Pleni-Weichselian (Klatkowa and Za1oba, 1991) i.e. the lengthiestorganic sequence from the EemianeWeichselian period. Due to therelative uniformity of the Eemian and EarlyWeichselian patterns ofvegetation in Central Europe (comp. Turner, 2002; Velichko et al.,2007), a detailed pollen analysis of the Ustków profile would bethe best tool for verifying its age. Moreover, fewer than a dozensites from the area of Poland spanning the EemianeEarly Weich-selian period have been incorporated into synthetic reconstructionsof more regional and/or global climateevegetation relationships

P. Kołaczek et al. / Quaternary International 268 (2012) 9e2010

(comp. Aalbersberg and Litt, 1998; Caspers and Freund, 2001).Hence, the results of such an analysis might be a valuable contri-bution to such reconstructions for that time interval in the future.

2. Regional setting

The Ustków site is located in central Poland within the TurekUpland e a part of the South Wielkopolska Lowland (Fig. 1). In theeast, the Turek Upland is adjacent to the Warta River valley, one ofthe main rivers in Poland. The Turek Upland developed asa moraine plain during the Warta stade, a recessional stade of theSaalian Glaciation (in Polish stratigraphy after Lindner, 2005 e theOdranian Glaciation). From the north and south, this plain is cut bydry denudation valleys.

The characteristic feature of the southern part of the TurekUpland is the occurrence of numerous small closed depressions,only just visible in the present-day landscape. Preliminarygeological studies show that these depressions developed as deepsteep-slope kettle-holes and organic deposits mainly fill them(Klatkowa and Za1oba, 1991).

3. Materials and methods

3.1. Fieldwork and description of deposits

The site presented in this paper is one of these closed depres-sions, located in a moraine plain composed of till from the time of

Fig. 1. Location maps of the Ustków site and cross-section of the kettle-hole filling. A. Locatisites cited in the text; 1 e Ustków (Klatkowa and Za1oba, 1991; this paper), 2 e Zgierz-Rudun(Granoszewski, 2003), 5 e Nidzica (Bi�nka et al., 2010), 6 e Czaple (Bi�nka and Nitychoruk, 201e Kub1owo (Roman and Balwierz, 2010), 10 e Mikorzyn (Stankowski and Nita, 2004), 11 e Le(Malkiewicz, 2010), 14 e Klinge (Novenko et al., 2008), 15 e Grabschütz (Litt, 1994; Litt et al.,2003), 18 e Bispingen (Field et al., 1994), 19 e Rederstall (Menke and Tynni, 1984), 20 e Füra(Beets et al., 2006), 23 e Sokli (Helmens et al., 2011), 24 eMondsee (Drescher-Schneider andrelation to central Poland (the numbers of other sites marked on the subfigure are compacollected; 1 e glacial till, 2 e mineral loam, 3 e organic loam, 4 e bituminous shale, 5 e p

the Warta stade. An 11 m long profile from the site was collected in1990 from the central part of the buried kettle-hole mire(133 m.a.s.l.; 51� 4503600N, 18�3600800E). Drilling was performedunder the management of B. Wicik who used a piston corer(Wieckowski type).

The lithology of the core was determined by H. Klatkowa and M.Za1oba and is presented in Table 1. Deposits above 262 cm wereverified by M. Ga1ka.

3.2. Laboratory treatment and pollen analysis

Twenty samples from the core were selected for primary pollenanalysis, which was carried out by M. Jastrzebska-Mame1ka (Fig. 2),but there is no information in the available literature about thetechnique used to prepare the samples (Klatkowa and Za1oba,1991). Afterwards, 82 samples from a profile section of890e90 cm were assigned for higher resolution analysis, but thisdid not happen until 2009, when the present working grouprenewed the research. For the final pollen analysis, all samples(1 cm3 volume) were prepared by a standard preparation procedureand then acetolysis was applied (Berglund and Ralska-Jasiewiczowa, 1986). To every sample, a weighed Lycopodiumtablet was added for further calculations of pollen concentration(Stockmarr, 1971). More than 500 arboreal pollen grains per samplewere counted (except for one sample where only slightly over 100AP grains were the basis for calculation, because of a very lowpollen concentration).

on map of Ustków in relation to Central, Western and Northern Europe and location ofki II (Jastrzebska-Mame1ka, 1985), 3 e Bieganin (Malkiewicz, 2003), 4 e Horoszki Du _ze1), 7 e Dziewule (Bi�nka and Nitychoruk, 2001), 8 e Imbramowice (Mamakowa, 1989), 9szczyno (Krupi�nski et al., 2006), 12 e qanieta (Balwierz and Roman, 2002), 13 e Gutów1996), 16 e Neumark (Litt, 1994; Litt et al., 1996), 17 e Gröbern (Litt, 1994; Kühl and Litt,moos (Müller et al., 2003), 21 e Jammertal (Müller et al., 2005), 22 e Amsterdam basinPapesch, 1998), 25 e Starunia (Stachowicz-Rybka et al., 2009); B. Location of Ustków intible with subfigure A); C. Cross-section of the kettle-hole from which the profile waseat, 6 e various-grained sand with admixture of gravel.

Table 1Simplified lithostratigraphic description of the Ustków sequence.

Depth [cm] Lithology

0e30 Humus with sand30e85 Various-grained sand with

gravel and stone admixture85e262 Organic loam, black262e303 Decomposed peat black, with

visible macroscopic plant remains303e455 Moss peat with macroscopic

remains, bright reddish-brownrapidly changing in colour toviolet black under oxidation

455e535 Loam with peat admixture,brown/black

535e600 Moss peat with plant macroscopicremains; reddish-brown, changingcolour to brown/black under oxidation

600e640 Moss peat reddish-brown; twobigger macroremains (probably roots)

640e718 Moss peat light reddish-brown718e725 Moss peat, compacted changing

into shale725e820 Bituminous shale, dark grey820e885 Organic loam, brownish-black885e895 Mineral loam, bluish-gray, with strata

of brown/black organic loam895e945 Mineral loam bluish-gray

(washed away till?)945e1100 Plastic till with stones (diameter 10 cm),

grayish-blue

P. Kołaczek et al. / Quaternary International 268 (2012) 9e20 11

The pollen taxa were determined with the assistance of themodern pollen slide collection of the W1adys1aw Szafer Institute ofBotany, Polish Academy of Science, and special keys and atlases

Fig. 2. Preliminary pollen diagram from the Ustków site carried out by Jastrzebska-Mame1organic loam, 3 e bituminous shale, 4 e peat, 5 e various-grained sand with admixture ofLiguliflorae, Comp. Tub. e Compositae Tubuliflorae, Chenop. e Chenopodiaceae, Juniper. etristachyum, Ranunc. t. acer e Ranunculus type acer, Ranunculus trych. e Ranunculus trycho

(Moore et al., 1991; Reille, 1992; Beug, 2004). The pollen percentagevalue of each taxon in each sample was calculated as a ratio of thetaxon to the sum of trees and shrubs (AP) and herbs (NAP, excludingaquatic and wetland plants, but including Cyperaceae and spores)in the sample. The percentages of excluded taxa were calculatedindividually as a ratio of the taxon to the sum of AP þ NAP þ taxon.Palynological diagrams were plotted using POLPAL software(Nalepka andWalanus, 2003). Palynological diagrams were dividedinto local pollen assemblage zones (LPAZs) and units (where zoneswere distinctly divided), local pollen assemblage subzones (LPASZs)defined and named following Birks and Birks (1980) and Janczyk-Kopikowa (1987). Afterwards LPAZs were referred to regionalpollen assemblage zones (RPAZs) (Mamakowa, 1988, 1989) andfurther to the chronostratigraphic units for north-western Europeproposed by Menke and Tynni (1984), Behre and Lade (1986) andBehre (1989), and also correlated with Marine Isotope Substages(Helmens et al., 2000).

3.3. Chronology

Bulk samples from selected depths were chosen for radiocarbondating and all were preparedwithout any screening of the material.The content of 14C in 6 bulk samples was measured by radiometricmethods in the Radiocarbon Laboratory of the ArchaeologicalMuseum in qód�z (lab. code e Lod) in 1990 and 1992, but only 3dates from the top were published (Klatkowa and Za1oba, 1991).Additionally, 3 thermoluminescence (TL) dates were done in the TLLaboratory of Gda�nsk University (Tables 2 and 3; Klatkowa andZa1oba, 1991).

Unfortunately, the long storage time, and an insufficient amountof material made any revision of these early results by AMS

ka (Klatkowa and Za1oba, 1991), redrawn and slightly changed; 1 e mineral loam, 2 e

gravel, 6 e humus with sand, Caryophyl. e Caryophyllaceae, Comp. Lig. e CompositaeJuniperus, Lycop. cf. alpinus e Lycopodium cf. alpinus, Lycop. tristach. e Lycopodium

phyllus, Typha an./Sparg. e Typha angustifolia/Sparganium, Vibur. e Viburnum.

Table 3Thermoluminescence (TL) dates from the Ustków profile.

Depth [cm] Lab. code Material dated TL date [yr before 1990]

60e80 UG-805 Coarse-grained sandwith gravel admixture

16,200 � 2400

880e900 UG-806 Mineral silt with organicsilt strata

68,100 � 10,200

10200 UG-807 Till with stones >50,100

P. Kołaczek et al. / Quaternary International 268 (2012) 9e2012

radiocarbon dating impossible, because of the reliable probabilityof age rejuvenation (Wohlfarth et al., 1998).

4. Results and discussion

The pollen spectra revealed 199 taxa of pollen and spores, whichcould be allocated to 12 local pollen assemblage zones (LPAZs)(Table 4; Figs. 3 and 4). Moreover, the patterns of vegetation shownby the analysis (unit 4.1) failed to support the chronology based on14C and TL dates. Even though the age of the bottom series of theprofile is in accordancewith the previous assessment (Saalian/EarlyWeichselian transition), the upper part consists of materialdeposited during the Rederstall stadial (MIS 5b) and there aredefinitely no traces of Pleni-Weichselian patterns of vegetation(comp. Klatkowa and Za1oba, 1991).

4.1. Patterns of vegetation reflected in the palynological profile fromUstków

4.1.1. Early Eemian; Marine Isotope Substage 5e (MIS 5e)4.1.1.1. Late Saalian/E1 RPAZ, Us-1 Pinus LPAZ. According to severalauthors, the transition between the late glacial of the SaalianGlaciation and the Early Eemian caused a rise in the meantemperature of the warmest month (MTWA) from 10 �C in the lateglacial to 16 �C in the early Eemian in Poland. Similarly, the meantemperature of the coldest month (MTCO) rose from �2 to 0 �C atthe beginning of the interglacial (Miros1aw-Grabowska, 2009 onthe basis of Mamakowa, 1989; Litt et al., 1996; Zagwijn, 1996;Aalbersberg and Litt, 1998; Cheddadi et al., 1998; Granoszewski,2003; Klotz et al., 2003; Kupryjanowicz, 2008). Although the highpercentages of Pinus sylvestris type pollen suggest the occurrence offorest at that time, the low pollen concentration seems to contra-dict this. The record of pinemight have been caused by the selectivecorrosion of grains, making this taxon more numerous relative toother taxa. Pollen of the P. sylvestris type is easy to identify amongother corroded grains (personal observation). Although not everypollen profile from Central and Western Europe shows a domina-tion of pine at the decline of the Late Saalian and the beginning ofthe Eemian interglacial (e.g. Granoszewski, 2003), most do reflectthis pattern (e.g. Mamakowa, 1989; Litt, 1994; Bi�nka andNitychoruk, 2001; Malkiewicz, 2003; Kühl and Litt, 2003;Kupryjanowicz, 2008; Novenko et al., 2008). Patches of openvegetationwith Poaceae, Selaginella selaginoides and Helianthemumwould then be likely (Figs. 3 and 4).

4.1.1.2. E1 RPAZ, Us-2a Betula (Ulmus) LPASZ. Rapidly expandingBetulawas a dominant taxon in pioneer woodland communities, inwhich it occurred together with pine (P. sylvestris type) and scat-tered specimens of Larix. Pine constantly replaced birch during thisperiod. Rapid climate amelioration at the beginning of the Eemianinterglacial resulted in a great rise in pollen production. The moreabundant occurrence of deformed or anomalously formed Betulagrains (with 4 or 5 pores) might have been a trace of Betulapubescens and Betula nana hybridisation (comp. Karlsdóttir et al.,

Table 214C dates from the Ustków profile.

Depth [cm] Lab. code Material dated 14C date [yr BP]

100 Lod-388 Bulk sample 16 100 � 200200 Lod-389 Bulk sample 19 400 � 250290 Lod-390 Bulk sample 25 300 � 310450 Lod-523 Bulk sample 25 800 � 500480e490 Lod-521 Bulk sample 28 200 � 500550 Lod-522 Bulk sample 29 700 � 800

2008). Even though dwarf birch was not detected in pollen, thereare known cases where this species, being a glacial relic in CentralEurope, failed to show a continuous record in pollen (comp.Ko1aczek et al., 2010). Elm (Ulmus), poplar and/or aspen (Populus),oak (Quercus) and ash (Fraxinus excelsior) occupied wet and dampniches forming forests similar to modern riparian ones. The firsttaxon reached the earliest Eemian optimum in Central Europe (seee.g. Menke and Tynni,1984;Mamakowa,1989; Field et al., 1994; Littet al., 1996; Drescher-Schneider and Papesch, 1998; Bi�nka andNitychoruk, 2001, 2011; Granoszewski, 2003; Beets et al., 2006;Kupryjanowicz, 2008; Novenko et al., 2008; Bi�nka et al., 2010;Roman and Balwierz, 2010). The edges of these woodlands wereovergrown by Humulus lupulus which reached the Eemianoptimum during that time, but earlier than in other sites in Poland(comp. Krupi�nski et al., 2006; Bi�nka et al., 2010; Bi�nka andNitychoruk, 2011) and this optimum lasted a shorter time (comp.Mamakowa, 1989; Bi�nka and Nitychoruk, 2001; Granoszewski,2003; Krupi�nski et al., 2006). The lakeshore was the habitat ofSalix thickets. Patches of heliophilous open vegetation with Juni-perus as well as steppe communities with Poaceae and Artemisiawere sparsely represented in the landscape. The belt of rushessurrounding the lake consisted of members of the Cyperaceaefamily, Phragmites australis, Equisetum and Typha latifolia.Submersed macrophytes included Myriophyllum spicatum andCeratophyllum sp.

4.1.1.3. E2 RPAZ, Us-2b Betula (PinuseQuercus) LPASZ. The amelio-ration of edaphic conditions led to the spread of P. sylvestris, whichgradually replaced pioneer birch-dominated woodlands. Fertilesoils in wet depressions were also constantly colonized by Quercuswhichmight have replaced birch as well as elm. Even though borealcommunities were in regression, Pinus cembra, Larix, B. nana, andSalix polaris still had enough habitats. Subsequent traces of ivy(Hedera helix) flowering in the declining part of the zone signifieda rise in the minimum mean January temperature up to �4.5 �C oreven �2 �C (comp. Iversen, 1944; Bi�nka and Nitychoruk, 2011).Open vegetation covered an area of similar size as in the previouszone. A new component of the belt of rushes surrounding the lakewas Cladium mariscus. Among water macrophytes, Potamogetonsubgen. Eupotamogeton was recorded.

4.1.2. Middle Eemian; Marine Isotope Substage 5e (MIS 5e)4.1.2.1. E3 RPAZ, Us-3 QuercuseFraxinus LPAZ. Oak dominatedterrestrial habitats and pushed pioneer birch-pine forests intomarginal niches. Beech (Fagus) might have occasionally occurred inthese forests, which is compatible with other sites in Poland(Granoszewski, 2003; Kupryjanowicz, 2008), but some authorsinterpreted the presence of its pollen as the effect of rebedding(�Srodo�n, 1990). In the contemporary equivalent of riparian wood-lands, the second important tree was ash (F. excelsior) reachingoptimal conditions for development in the whole Eemian. Elm andalder (Alnus) could have also grown in those forests as an admix-ture. A rise in soil fertility caused by the domination of deciduousforest enabled the rapid spread of hazel (Corylus avellana) whichgradually blocked oak regeneration and caused its subsequent

Table 4Summary of the local pollen assemblage zones (LPAZs) in the Ustków profile (see Figs. 3 and 4).

LPAZ, LPASZ Depth [cm] Description

Us-12 Pinus-Artemisia-Alnus 195e90 Rise in Pinus sylvestris type (44e59%), Alnus undiff. (4e9%), Picea abies (2e6%), Carpinus (0.5e3%) andCorylus avellana (1e2%). Decline in Pinus cembra type. Stable values of Artemisia (3e6%).

Us-11 Betula-Poaceae-Artemisia 285e195 Decline in AP. Fall and subsequent rise in Pinus sylvestris type, increase in Betula undiff., followed bydecline in the upper part of the zone. Rise in Artemisia (up to 10%).

Us-10c Pinus (Larix) 339e285 Rise in AP, fluctuation of Pinus sylvestris type and Betula undiff. The highest values of Larix (18%) andPinus cembra type (13%) in the profile. Rapid fall in Artemisia and gradual decline in Poaceae undiff.Fluctuation of Filicales monolete (0.7e19%) and Pteridium aquilinum (0.1e3%, max.).

Us-10b Pinus (Artemisia) 346.5e339 Single spectrum; rapid fall in AP together with Artemisia maximum in the profile (18%), rise in Poaceaeundiff. Minimum pollen concentration in the profile (w14 � 103/cm3).

Us-10a Pinus (Pinus) 392.5e346.5 Sharp increase in Pinus sylvestris type (78%, max. in the zone) and subsequent decline; correlated withdecrease in Betula undiff.

Us-10 Pinus 392.5e285 Fluctuation of Pinus sylvestris type (34e62%), rise in Pinus cembra type.Us-9 Betula 455e392.5 Maximum of Betula undiff. (89%) and Betula nana type in the profile (8%). Decline in Artemisia

and Sphagnum.Us-8b Pinus-Poaceae (Juniperus) 475e455 Pinus sylvestris type falls constantly. The highest values of Juniperus (9%) and Poaceae undiff. (14%)

in the profile.Us-8a Pinus-Poaceae (Calluna) 535e475 Gradual decrease in Pinus sylvestris type, rise in Alnus undiff., Corylus avellana and Picea abies type. Rise in

Calluna vulgaris (max. 4%), Poaceae undiff. and Artemisia. Maximum of Sphagnum (28%).Us-8. Pinus-Poaceae 535e455 Gradual fall in Pinus sylvestris type still dominating arboreal taxon, increase in Pinus cembra type. Rise

in Poaceae undiff.Us-7 Pinus 599e535 The highest values of Pinus sylvestris type (max. 90%) in the profile, increase in Pinus cembra type. Fall in

Alnus undiff., Picea abies type, Carpinus, Corylus avellana and Abies alba.Us-6b Picea (Alnus) 645e599 Fluctuation of Pinus sylvestris type (12e42%),rise in Alnus undiff. (24% max. in the profile) and Picea abies

type (second maximum 30%). Stable occurrence of Sphagnum and Pteridium aquilinum.Us-6a Picea (Abies) 680e645 The highest percentages of Abies alba in the profile (48%) and the first maximum of Picea abies type (32%).

Us-6 Picea 680e599 Maximum of Picea abies type, high values of Alnus undiff. Sample between depths 620e640 cm homogenizedinto one sample.

Us-5b Carpinus (Taxus) 717.5e680 Taxus baccata reached the highest values in the profile (3.7%), stable values of Alnus undiff.Us-5a Carpinus (Tilia) 757.5e717.5 Rapid rise in Carpinus (from 8 to 58.5%, max. in the profile), after maximum of Tilia cordata type (12%), fall in its

values. Increase in Alnus undiff. Maximum of Filicales monolete (25%) and Salvinia microsporocarps (2.8%).Us-5 Carpinus 757.5e680 Domination of Carpinus.Us-4 Corylus 795e757.5 The highest percentages of Corylus avellana in the profile (max. 83%). Rise in Tilia cordata type in the upper part

(up to 6%). Maximum pollen concentration (w12.6 � 106/cm3).Us-3 Quercus-Fraxinus 813.5e795 After Quercus maximum (64%) visible decline in the curve, maximum of Fraxinus excelsior (5%). Strong increase

in Corylus avellana (from 6 to 29%).Us-2b Betula (Pinus-Quercus) 852.5e813.5 Decrease in Betula undiff., rise in Pinus sylvestris type nad P. cembra type. Visible increase in Quercus

percentages (from 7 to 25%).Us-2a Betula (Ulmus) 885e852.5 Rapid increase in Betula undiff. (up to 67%, the first maximum in the profile) simultaneous with sharp

fall in Pinus sylvestris type. Maximum of Ulmus curve (5%).Us-2 Betula 885e813.5 The first maximum of Betula undiff. in the profile, the highest values of Ulmus, rise in Quercus. Stable

percentages of Poaceae undiff.Us-1 Pinus 890e885 The zone is represented by a single spectrum. Domination of Pinus sylvestris type (75%), high values

of P. cembra type (6%).

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retreat. Syringa and/or Ligustrum (pollen of Ligustrum type, sensuBeug, 2004), which were also more common in other parts ofCentral Europe in the Middle Eemian (Mamakowa, 1989; Litt, 1994;Krupi�nski et al., 2006; Bi�nka and Nitychoruk, 2011), might haveoccupied the understorey under looser canopies. The shadedgroundcover was overgrown by ivy but in places with better lightconditions, bracken (Pteridium aquilinum) prevailed. C. mariscusprobably retreated from the belt of rushes, although it occurredirregularly until the end of the Middle Eemian. Nympheaceae(mainly Nymphaea alba) spread in the water body, which led to thelimitation of the population of light-demanding Myriophyllum andprobably allowed the spread of Ceratophyllum sp. e a taxon knownfor its higher tolerance of shade (comp. Matuszkiewicz, 2005).

4.1.2.2. E4 RPAZ, Us-4 Corylus L PAZ and Us-5a Carpinus (Tilia) LPASZto 752.5 cm. The climate of the E4 RPAZ as well as the subsequentE5 RPAZ had suboceanic features and this period is treated as theinterglacial climatic optimum, when the minimum MTWA wasestimated at 19e20 �C and the minimumMTCO at 2e4 �C (Zagwijn,1996; Aalbersberg and Litt, 1998). On the other hand, somereconstructions pointed to an MTCO below 0 �C during theseperiods in central Germany (Kühl and Litt, 2003). At that time, thedynamic expansion of hazel (C. avellana), which probably built

dense homogenous thickets and/or short-stemmed forests, led toa change in deciduous forest structures and limited oak reproduc-tion. Patches of woodland with oak and ash overgrew the placeswhere ground water disallowed or restricted hazel existence.Despite difficulties in tree regeneration caused by dense thicketsand/or canopies of hazel, maple (Acer) had optimal conditions fordevelopment in the whole Eemian, similarly to other areas inCentral Europe (e.g. Mamakowa, 1989; Litt, 1994; Granoszewski,2003). In the declining phase of the E4 RPAZ small-leaved lime(Tilia cordata) and hornbeam (Carpinus) expanded starting to formforests similar to present woodlands from the Tilio-Carpinetumclass. Simultaneously yew (Taxus baccata) could have been anadditional element of those forests. Although its pollen grains weredetected irregularly, this taxon is known for poor dispersal andeven a single grain may indicate its in situ occurrence(Nory�skiewicz, 2001; Krupi�nski et al., 2004).

4.1.2.3. E5 RPAZ, Us-5 Carpinus LPAZ above 752.5 cm. Rapidlyexpanding hornbeam, which might have established pure stands inmost areas, replaced homogenous communities with hazel,although this was probably an important component of theunderstorey of those forests. At the beginning of Carpinus domi-nation T. cordata remained an important component of forests, but

Fig. 3. Pollen diagram from the Ustków site e selected curves included in the total pollen sum; 1 e mineral loam, 2 e organic loam, 3 e bituminous shale, 4 e peat, 5 e various-grained sand with admixture of gravel, 6 e humus withsand, Aln. e Alnus, Art. e Artemisia, Call. e Calluna, Fr.e Fraxinus, Jun. e Juniperus, Pin. e Pinus, Pleur. austr. e Pleurospermum austriacum, Q e Quercus. Gray area of each percentage column expresses 10� exaggeration of percentagevalues.

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Fig. 4. Pollen diagram from the Ustków site e selected curves of herbs excluded from the total pollen sum (except Filipendula and Caltha type), Cryptogams and other NPPs; 1 e

mineral loam, 2 e organic loam, 3 e bituminous shale, 4 e peat, 5 e various-grained sand with admixture of gravel, 6 e humus with sand, Aln. e Alnus, Art. e Artemisia, Call. eCalluna, Carpin. e Carpinus, Fr.e Fraxinus, Jun. e Juniperus, Koen. isl. e Koenigia islandica. Pin. e Pinus, Q e Quercus. Gray area of each percentage column expresses 10� exaggerationof percentage values.

P. Kołaczek et al. / Quaternary International 268 (2012) 9e20 15

afterwards the occurrence of this taxon was limited by expandinghornbeam in the younger part of the E5 RPAZ. This phenomenonhad a similar course in the Central Europe (Mamakowa, 1989;Table 5, see Litt, 1994; Granoszewski, 2003; Kupryjanowicz, 2008;

Novenko et al., 2008). Alder (Alnus) gradually colonized damp andmoist areas, where ferns from the Osmunda genus (Osmunda cin-namomea type) might have overgrown the undercover (see Lordand Travis, 2011). In the younger phase of the E5 RPAZ (Us-5b

Table 5Local pollen assemblage zones of the Ustków profile (LPAZs), in comparisonwith northern Germany biozones (Menke and Tynni, 1984 and; Caspers and Freund, 2001), central Germany biozones (Litt, 1994), regional pollen zones(RPAZs) for Poland (Mamakowa, 1988, 1989), RPAZs for the Konin region e a westerly located area to Ustków (Tobolski, 1991), LPAZs of the Horoszki Du _ze profile in eastern Poland (Granoszewski, 2003) referred to chro-nostratigraphic units for north-western Europe (Menke and Tynni, 1984; Behre and Lade, 1986; Behre, 1989). WFIIbKr e intra-Brörup kryomer (Caspers and Freund, 2001). A correlation of the Eemian biozones between northernand central Germany is in accordance with Aalbersberg and Litt (1998) and Kühl and Litt (2003).

Northern Germany Central Germany Konin Region e western Poland Ustków Horoszki Wielkie e eastern Poland Poland Chronostratigraphy

WFIII WFIII NAPI Us-12 Pinus-Artemisia-Alnus HD-22 NAP-B. nana-Salix polaristype-P. cembra type

EV3. Gramineae-Artemisia-Betula nana

Rederstall

Us-11 Betula-Poaceae-Artemisia HD-21 NAP-Pinus-P. cembra typeWFIIb WFII Pinus Us-10c Pinus (Larix) HD-20 Pinus-P. cembra type-NAP EV2. Betula-Pinus Brørup

HD-19 Pinus-Larix-PiceaHD-18 Pinus-P. cembra type-PiceaHD-17 Pinus-Betula-LarixHD-16 Betula-NAP

WFIIbKr Us-10b Pinus (Artemisia) HD-15 NAP-BetulaWFIIb Us-10a Pinus (Pinus) HD-14 BetulaWFIIa Betula-NAP Us-9 Betula

Betula-LarixNAP-Betula

WF Ia WF I Artemisia-NAP Us-8b Pinus-Poaceae(Juniperus)

HD-13 Poaceae-Cyperaceae-Juniperus(Salix polaris type)

EV1. Gramineae-Artemisia-Betula nana

Herning

WF Ib Us-8a Pinus-Poaceae(Calluna)

E7 Pinus E7 Pinus Pinus Us-7 Pinus HD-12 Pinus-Larix E7. Pinus Late EemianHD-11 Pinus-Betula-PiceaHD-10 Pinus

E6 Pinus, Picea, Abies E6b Pinus, Picea, Abies Picea-Abies Us-6b Picea (Pinus-Alnus) HD-9 Picea-Abies-Carpinus (Pinus) E6. Picea-Abies-Alnus6a Carpinus, Abies Us-6a Picea (Abies)

E5 Carpinus, Picea E5 Carpinus Carpinus Us-5b Carpinus (Taxus) HD-8 Carpinus-Alnus-Picea E5. Carpinus-Corylus-Alnus Middle EemianUs-5a Carpinus (Tilia) HD-7 Carpinus-Corylus-Tilia (Alnus)

E4b Corylus, Taxus, Tilia E4b Corylus,Taxus, Tilia

Corylus Us-4 Corylus HD-6 Corylus-Tilia-Alnus E4. Corylus-Quercus-TiliaHD-5 Corylus-Taxus-Ulmus (Quercus)

E4a Quercetummixtum, Corylus

E4a Quercetummixtum, Corylus

Quercus Us-3 Quercus-Fraxinus HD-4 Quercus-Fraxinus-Ulmus E3. Quercus-Fraxinus-Ulmus

E3 Pinus, Quercetummixtum

E3 Pinus, Quercetummixtum

Pinus-Betula Us-2b Betula (Pinus-Quercus) HD-3 Pinus-Betula-Quercus-Ulmus E2. Pinus-Betula-Ulmus Early Eemian

E2 Pinus, Betula E2 Pinus, BetulaE1 Betula E1 Betula Us-2a Betula (Ulmus) HD-2 Betula-Pinus E1. Pinus-Betula

E1/S Us-1 Pinus HD-1 NAP-Juniperus-Pinus Late Saalian/Early Eemian

P.Kołaczek

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P. Kołaczek et al. / Quaternary International 268 (2012) 9e20 17

LPASZ) T. baccata reached its optimum in Eemian forests. Thisphenomenon took place later in comparison with other sites fromCentral Europe (e.g. Mamakowa, 1989; Litt, 1994; Drescher-Schneider and Papesch, 1998; Kupryjanowicz, 2008) but occurredwithin at least a 50 km radius of Ustków (Jastrzebska-Mame1ka,1985; Malkiewicz, 2003). Although forest density was the highestin the whole Eemian succession, the presence of Calluna vulgarispoints to places with relatively good light conditions under cano-pies. In the hornbeam-dominated forests ivy found favourableconditions for growth and flowering. In the undergrowth, and/or inthe belt of rushes surrounding the lake, Polypodiaceae (Filicalesmonolete) were more common. In the last phase of the waterbody’s existence Salvinia natans reached its optimum. The occur-rence of this fern indicates calm shallow eutrophic waters anda thick layer of organic substrate in the bottom sediments. In allprobability, an association similar to present-day Lemno minoris-Salvinietum natantis (Slavni�c, 1956) Korneck 1959 occurred there,where other members of Lemna type and Ceratophyllum were alsopresent (Michalska-Hejduk and Kope�c, 2002). The shallowing ofthe water body and a rise in Salvinia frequency restricted thedevelopment of Nymphaeaceae.

4.1.3. Late Eemian; Marine Isotope Substage 5e (MIS 5e)4.1.3.1. E6 RPAZ, Us-6 Picea LPAZ. A dynamic expansion of borealforest composed mainly of spruce (pollen of Picea abies type), fir(Abies alba) and birch replaced the forests with Carpinus ina significant manner. Nonetheless, patches of hornbeamwoodlandswere still present in the landscape. These processes were inducedby climate cooling expressed in a drop in the MTWA to 15e18 �Cand the MTCO to �2e0 �C at sites from central and southernPoland and some sites from central, eastern and northern Germany(Aalbersberg and Litt, 1998; Kühl and Litt, 2003). The younger partof the zone (Us-6a LPASZ) was the period when fir was the domi-nant forest species, as in other sites from Central and WesternEurope (Litt, 1994; Litt et al., 1996; Kühl and Litt, 2003; Müller et al.,2003, 2005). In the subsequent phase (Us-6b LPASZ) P. sylvestris andP. cembra substituted A. alba in forests. In contemporary woodlandcommunities, H. helix still existed: Ustków was probably one of thelatest stands where it flowered in western/central Poland (comp.Tobolski, 1991; Stankowski and Nita, 2004). Nevertheless, pollen ofthis species was identified in the Late Eemian deposits fromsouthwestern and central Poland (Mamakowa, 1989) as well asfrom central and eastern Germany (Litt,1994; Novenko et al., 2008).Less frequent in comparisonwith the E5 RPAZ was T. baccata. Dampareas were dominated by alder, which strongly replaced other taxawith high moisture requirements like elm and oak. In the Us-6bLPASZ, pollen spectra suggest a decline in woodland cover, whichenabled a slight spread of light-demanding patches of steppe withPoaceae, Artemisia, Gypsophila and Asteraceae, together with Juni-perus thickets and sandy grasslands/heathlands with C. vulgaris andpatches of P. aquilinum. Unfortunately, this slight vegetation shiftwas detected only in one amalgamated sample from depths of640e620 cm, so that it is impossible to speculate about climateoscillation. During the period reflected by the Us-6a LPASZ, themirewas still dominated by Cyperaceae, whereas in the subsequentphase (Us-6b LPASZ) sedge communities retreated and Sphagnummosses were more common. The constant occurrence of Utriculariaduring the E6 RPAZ points to the existence of small meso-/dystro-phic ponds (Matuszkiewicz, 2005).

4.1.3.2. E7 RPAZ, Us-7 Pinus LPAZ. During this period the MTWAdrop to 15 �C whereas the decrease in the MTCO reached valuesbetween �5 and �10 �C (Kühl and Litt, 2003). The consequentcooling engendered P. sylvestris domination in most habitats. Thisspecies together with the accompanying P. cembra, P. abies, Betula

sp. and Larix probably built woodlands/forests then. The increase inlandscape openness triggered the spread of tundra communitieswith B. nana and S. selaginoides similarly to other sites from CentralEurope (e.g. Granoszewski, 2003). The pollen spectra failed toreflect a drastic cool shift demonstrated by a rise in Betula whichwas detected in eastern Poland (Granoszewski, 2003;Kupryjanowicz, 2008). As in the E6 zone, small pools inhabited byUtricularia were present on the mire.

4.1.4. Herning stadial, Marine Isotope Substage 5d (MIS 5d)4.1.4.1. EV1 RPAZ, Us-8 Pinus-Poaceae LPAZ. The beginning of theHerning stadial brought a drop in the minimum MTWA from 8 to10 �C in central Poland (Aalbersberg and Litt, 1998) to an MTWAabout 10 �C in northern Germany (Caspers et al., 2001) anda minimumMTWA of about 13 �C in eastern Germany (Aalbersbergand Litt, 1998). The minimum MTCO dropped to �8 to �12 �C ineastern Germany (Aalbersberg and Litt, 1998) or even below�15 �Cin central Germany (Walkling and Coope, 1996). This climaticdeterioration caused a drastic retreat of pine-dominated wood-lands, which was recorded in the whole area of Central Europe (e.g.Menke and Tynni, 1984; Mamakowa, 1989; Jastrzebska-Mame1ka,1985; Litt, 1994; Balwierz and Roman, 2002; Granoszewski, 2003;Kupryjanowicz, 2008). Patches of them were still present in thelandscape, but birches were a more common component. Taxo-nomically rich steppe communities overgrew dryer areas, wherePoaceae, Artemisia, Chenopodiaceae, Brassicaceae, Rubiaceae, Tha-lictrum and Rumex acetosa/acetosella dominated. The occurrence ofAmbrosia (Ambrosia artemisiaefolia type, sensu Punt and Hoen,2009), which is nowadays represented in Europe by one nativespecies e Ambrosia maritima (Makra et al., 2005), may suggest thatthis species had a broader range (nowadays it is restricted to coastalareas of the Mediterranean Sea), or that there was a broaderdistribution of American species. In other Eemian and/or Weich-selian profiles from the area of Central Europe this taxon wasrecognized in the Pleni-Weichselian (originally Plenivistulian) layerin the Horoszki Du _ze site in eastern Poland (Granoszewski, 2003)and in the Starunia site in southwestern Ukraine (Stachowicz-Rybka et al., 2009). Heats with C. vulgaris, Bruckenthalia (Erica)spiculifolia with the probable addition of dwarf willows (S. polaristype) and Diapensia lapponica as well as shrub willows played animportant role in the landscape. Communities of dwarf shrubswereprobably replaced by thickets with Juniperus and grasslandcommunities with Poaceae and Artemisia in the younger part of theHerning stadial. At that time, the cover of arboreal plant fell toa minimum. The bipartition of the stadial is well reflected innorthern Germany and the Netherlands and it is explained bya shift from more oceanic (in the older part) to a more subconti-nental/continental climate (in the younger part). In these cases,climatic continentalisation was manifested mostly by the expan-sion of Poaceae and Artemisia (Caspers and Freund, 2001). Polishsites represent different patterns of the vegetation of the Herningstadial i.e. with a distinct division into two phases (Jastrzebska-Mame1ka, 1985), sites without that clear division (Malkiewicz,2010; Roman and Balwierz, 2010) and sites where Poaceae andArtemisia dominated through the whole stadial (eastern Poland;Granoszewski, 2003). From less representative communities,patches of dwarf-shrub tundra with B. nana, as well as herb tundracommunities with Polygonum bistorta/vivipara and members ofGentianaceae and Saxifragaceae may have occupied wetter habi-tats. Cooling led to the revival of a eutrophic water body witha similar composition of submersed macrophytes to that in theinterglacial. However, in the first stage Ranunculus trichophyllus andCeratophylum sp. communities were more common and afterwardsthe frequency of Myriophyllum and Potamogeton subgen. Eupota-mogeton increased. At the end of the stadial Ceratophyllum sp.

P. Kołaczek et al. / Quaternary International 268 (2012) 9e2018

appeared again, at the same time as Koenigia islandica, whichprobably occupied the lakeshore (Hedberg, 1997).

4.1.5. Brørup interstadial; Marine Isotope Substage 5c (MIS 5c)4.1.5.1. EV2 RPAZ, Us-9 Betula and Us-10 Pinus LPAZs. The Brørupinterstadial (MIS 5c) brought climate warming reflected in the risein the minimumMTWA up to approx. 15 �C, whereas the minimumMTCO temperature in that area was approx. �13 �C (Aalbersbergand Litt, 1998). At that time, birch rapidly spread and builtpioneer woodland, the light herb layer of which was suitable for thesurvival of steppe vegetation. Simultaneously, patches of tundrawith dwarf birch reached the most optimal conditions for theiroccurrence in the Early Weichselian. The second stage of forestdevelopment (Us-9 LPAZ) was the expansion of P. sylvestriswith theaddition of P. cembra and Larix. In that period, a rapid shift invegetation took place (Us-9b LPASZ), in which an Artemisiamaximum suggests the expansion of steppe. However, the vege-tation pattern after this relatively short-lived episode shows littleevidence of regeneration with a typical succession sequence fromBetula to Pinus as at the beginning of the interstadial. The mostcredible explanation of this phenomenonmight be a rapid decreasein the production of arboreal pollen in relation to herbs (especiallyArtemisia), which only creates an impression of rapid steppeexpansion. However, the pollen spectra reflect a retreat of forestafter the Artemisia maximum. Evidence of an intra-Brørup coldperiod was found in some diagrams from Western, Central andNorthern Europe (Caspers and Freund, 2001; Granoszewski, 2003).The last stage of interstadial forest development was the spread oflarch (Larix), which, taking into consideration its pollen frequency,probably dominated in communities surrounding the site (comp.Huntley and Birks, 1983). A similar pattern has been observed ineastern Poland, where during that time larch reached the Weich-selian optimum (Granoszewski, 2003), and northern Germany(Caspers and Freund, 2001). The constant presence of Alnus andCorylus may signify their sparseness around the mire. The formertaxon especially seems to be more widespread. Alder pollen wasidentified at sites located in northern Germany, the Netherlandsand Denmark (Caspers and Freund, 2001). Moreover, macroscopicremains of alder indicate that it reached areas north of the ArcticCircle, which is reflected at the Sokli site in northern Fennoscandia(Helmens et al., 2011). Another component of the forest might bePicea, whose values point to its in situ occurrence during the Larixphase (comp. Harmata, 1987; Björkman, 1996), but the values ofspruce were more comparable to eastern Poland, whereas theywere lower than those detected in northern Germany (Caspers andFreund, 2001).

At the beginning of the Brørup interstadial the shallowing waterbody was colonized by Nyphaeaceae and Ceratophyllum sp., andafterwards underwent terrestrialization. The domination ofCyperaceae points to nutrient enrichment of the developed fen/bog(comp. Lata1owa et al., 2004). Finally, a rise in Polypodiaceae (Fili-cales monolete) and P. aquilinum may suggest that this taxa spreadon themire surface, whichmight have been caused by a decrease inthe water level.

4.1.6. Rederstall stadial; Marine Isotope Substage 5b (MIS 5b)4.1.6.1. EV3 RPAZ, Us-11 Betula-Poaceae-Artemisia and Us-12PinuseArtemisiaeAlnus LPAZs. Climate cooling manifested bya drop in the minimum MTWA to about 10 �C (reconstruction foreastern Germany; Aalbersberg and Litt, 1998), led to the re-expansion of steppe communities, heliophytous juniper thicketsand dry heathlands similar to those in the Herning interstadial.However, in this case there was no significant differentiation intoa Calluna and subsequent Juniperus phase (Caspers andFreund, 2001; Granoszewski, 2003). The high percentages of

thermophilous arboreal taxa such as Alnus, C. avellana, Carpinusand T. cordata and spores of O. cinnamomea type were probablycaused by pollen redeposition, which also manifested itself in thehigh number of corroded pollen grains. Pollen spectra whichpurportedly point to the occurrence of pine-birch woodlands withan addition of larch may be invalidated by the possible resedi-mentation of the pollen of those taxa. The rise in the frequency ofPicea pollen was probably induced by these processes as well,because most sites from Central and Western Europe failed toreflect the expansion of spruce in that period (e.g. Caspers andFreund, 2001; Granoszewski, 2003; Behre et al., 2005). Duringthis stadial the ground water level rose and a small eutrophic lake(pools) similar to that from the Herning stadial (MIS 5d) devel-oped. In the earlier phase, reflected by the Us-11 LPAZ, a belt ofrushes was composed of Cyperaceae, Typhaceae/Sparganiaceaeand Equisetum accompanied by T. latifolia (?), Hippuris vulgaris andButomus umbellatus. In the younger phase (Us-12 LPAZ) P. australiswas more common. A Sphagnum peat bog probably existed at theedge of the basin, where Drosera rotundifolia and Menyanthes tri-foliata may have occurred. A succession of submersed macro-phytes during the stadial started with Nymphaeaceae,Myriophyllum and Potamogeton subgen. Eupotamogeton, and aftertheir decrease R. trichophyllus became more frequent, contrary tothe succession of macrophytes in the Herning stadial. The pres-ence of bulrushes (T. latifolia, Typha angustifolia) and Ceratophyllumsp. indicates a minimum mean June temperature of >13 �C (seevan Geel et al., 1981; Litt, 1994; Isarin and Bohncke, 1999) which isincompatible with the results of Aalbersberg and Litt (1998).Unfortunately, the relatively high number of rebedded taxa withhigher temperature demands casts doubt on the abovementionedestimates, because the pollen from both bulrushes may have alsobeen on a secondary bed.

5. Conclusions

Palynological analysis of the profile collected from the palaeo-kettle mire in Ustków pointed to a shorter time-span ascompared with the preliminary results and chronology establishedon 14C and TL dating. This fact should encourage other authors totake a critical view of the results of dating obtained from bulksamples, especially in situations where profiles are relatively shortand where dating reveals an age earlier than the Late Glacial of theWeichselian.

Despite the relatively low resolution of the pollen diagram, theprofile showed several patterns which differentiate it from otherslocated in Central Europe. The early elm maximum in the EarlyEemian, simultaneous with the phase of birch woodland domina-tion, the late maximum of yew in the declining part of the MiddleEemian, and the distinct division of the older part of the LateEemian into phases of fir-spruce and pine-spruce forest dominationare noteworthy in the Eemian part of the profile. The part of theprofile which reflects the Early Weichselian also indicated manyinteresting patterns, including a division of the Herning stadial intotwo phases of vegetation development (older with Calluna andyounger with Juniperus and Artemisia domination), and a phase oflarch domination during the late Brørup, which sets absolute recordvalues among pollen profiles from Central Europe (more than 15%of the total pollen sum).Moreover, the intra-Brørup cold oscillation,which is rarely detected in pollen profiles from this part of Europe,was distinctly marked in the Ustków profile. Local pollen taxarevealed 3 phases of lake existence in the site i.e. (i) Late Saalian-middle E5 zone (a decline of the lime-phase), (ii)Herning-earlyBrørup, (iii)Rederstall. Between those phases, poor and/or rich fenand/or bog functioned.

P. Kołaczek et al. / Quaternary International 268 (2012) 9e20 19

Acknowledgements

We would like to dedicate our paper to Prof. H. Klatkowa, MScM. Za1oba and Dr M. Jastrzebska-Mame1ka (University of qód�z,Poland), because it was thanks to their scientific enthusiasm thatthe site was discovered and preliminarily described. Unfortunately,none of them lived to see the submission of our manuscript. Wewish to express our gratitude to Prof. K. Szczepanek (JagiellonianUniversity, Poland) for the transfer of material to us, and P. Trzeciak,Msc (the Radiocarbon Laboratory of the Archaeological Museum inqód�z) for providing 3 unpublished radiocarbon dates from theUstków profile. We would also like to thank Ms. Barbara Now-aczy�nska (Jagiellonian University, Poland) for the laboratory prep-aration of samples and Dr M. Ga1ka (A. Mickiewicz University,Poland) for the revision of lithology of the upper part of the profile.We would also like to express our special appreciation to Prof. K.Milecka (A. Mickiewicz University, Poland) and the anonymousreviewers who helped to improve our manuscript.

Appendix A. Supplementary material

Supplementary material associated with this article can befound, in the online version, at doi:10.1016/j.quaint.2012.05.004.

References

Aalbersberg, G., Litt, T., 1998. Multiproxy climate reconstructions for the Eemian andEarly Weichselian. Journal of Quaternary Science 13, 367e390.

Balwierz, Z., Roman, M., 2002. A new Eemian interglacial to Early Vistulian site atqanieta, Central Poland. Geological Quarterly 46, 207e217.

Beets, D.J., Beets, C.J., Cleveringa, P., 2006. Age and climate of the late Saalian andearly Eemian in the type-area, Amsterdam basin, The Netherlands. QuaternaryScience Reviews 25, 876e885.

Behre, K.-E., Lade, U., 1986. Eine Folge von Eem und 4 Weichsel-Interstadialen inOerel/Niedersachsen und ihr Vegetations e ablauf. Eiszeitalter und Gegenwart36, 11e36.

Behre, K.-E., Hölzer, A., Lemdahl, G., 2005. Botanical macro-remains and insectsfrom the Eemian and Weichselian site of Oerel (northwest Germany) and theirevidence for the history of climate. Vegetation History and Archaeobotany 14,31e53.

Behre, K.-E., 1989. Biostratigraphy of the last glacial period in Europe. QuaternaryScience Reviews 8, 25e44.

Berglund, B.E., Ralska-Jasiewiczowa, M., 1986. Pollen analysis and pollen diagrams.In: Berglund, B.E. (Ed.), Handbook of Holocene Palaeoecology and Palae-ohydrology. John Wiley & Sons, Chichester, pp. 455e484.

Beug, H.-J., 2004. Leitfaden der Pollenbestimmung für Mitteleuropa und angren-zende Gebiete. Verlag Dr. Friedrich Pfeil, München.

Bi�nka, K., Nitychoruk, J., 2001. Late Saalian climate changes in Europe in the light ofpollen analysis and the problem of two-step deglaciation at the oxygen isotopestage 6/5e transition. Boreas 30, 307e316.

Bi�nka, K., Nitychoruk, J., 2011. Cyclicity in the Eemian climate? A case study of theEemian site at Czaple, Eastern Poland. Review of Palaeobotany and Palynology164, 39e44.

Bi�nka, K., Nitychoruk, J., Dzier _zek, J., 2010. Climate stability during the Eemian enew pollen evidence from the Nidzica site, northern Poland. Boreas 40,342e350.

Birks, H.J.B., Birks, H.H., 1980. Quaternary Palaeoecology. Edward Arnold Limited,London.

Björkman, L., 1996. The Late Holocene history of beech Fagus sylvatica and Norwayspruce Picea abies at stand-scale in Southern Sweden. Lundqua Thesis 39, LundUniversity, Lund.

Caspers, G., Freund, H., 2001. Vegetation and climate in the Early- and Pleni-Weichselian in northern Central Europe. Journal of Quaternary Science 16,31e48.

Caspers, G., Merkt, J., Müller, H., Freund, H., 2001. The Eemian Interglaciation inNorthwestern Germany. Quaternary Research 58, 49e52.

Cheddadi, R., Mamakowa, K., Guiot, J., de Beaulieu, J.-L., Reille, M., Andrieu, V.,Granoszewski, W., Peyron, O., 1998. Was the climate of the Eemian stable? Aquantitative climate reconstruction from seven European pollen records.Palaeogeography Palaeoclimatology Paleoecology 143, 73e85.

Drescher-Schneider, R., Papesch, W., 1998. A contribution towards the reconstruc-tion of Eemian vegetation and climate in Central Europe: first results of pollenand oxygen-isotope investigations from Mondsee, Austria. Vegetation Historyand Archaeobotany 7, 235e240.

Field, M.H., Huntley, B., Müller, H., 1994. Eemian climate fluctuations observed ina European pollen record. Nature 371, 779e783.

Granoszewski, W., 2003. Late Pleistocene vegetation history and climatic changes atHoroszki Du _ze, eastern Poland: a palaeobotanical study. Acta Palaeobotanica(Suppl. 4), 3e95.

Harmata, K., 1987. Late-glacial and Holocene history of vegetation at Roztoki andTarnowiec near Jas1o (Jas1o-Sanok depression). Acta Palaeobotanica 27 (1),43e65.

Hedberg, O., 1997. The genus Koenigia L. emend. Hedberg (Polygonaceae). BotanicalJournal of the Linnean Society 124, 295e330.

Helmens, K.F., Räsänen, M.E., Johansson, P.W., Jungner, H., Korjonen, K., 2000. TheLast Interglacial-Glacial cycle in NE Fennoscandia: a nearly continuous recordfrom Sokli (Finnish Lapland). Quaternary Science Reviews 19, 1605e1623.

Helmens, K., Väliranta, M., Engels, S., Luoto, T., Self, A., 2011. Dramatic shifts inEarly Glacial (MIS 5d-c) climate regimes recorded by multi-proxy data fromthe high latitude site Sokli (N Finland). http://www.inqua2011.ch/?a¼programme&subnavi¼abstract&id¼1971&sessionid¼7 2011.

Huntley, B., Birks, H.J.B., 1983. An Atlas of Past and Present Pollen Maps for Europe0e13,000 Years Ago. Cambridge University Press.

Isarin, R.F.B., Bohncke, S.J.P., 1999. Mean July temperatures during the YoungerDryas in Northwestern and Central Europe as inferred from climate indicatorplant species. Quaternary Research 51, 158e173.

Iversen, J., 1944. Viscum, Hedera and Ilex as climate indicators. Geologiska Före-ningens I Stockholm Förhandlingar 66, 463e483.

Janczyk-Kopikowa, Z., 1987. Uwagi na temat palinostratygrafii czwartorzedu(Remarks of palynostratigraphy of the Quaternary). Kwartalnik Geologiczny 31,155e162 (in Polish with English summary).

Jastrzebska-Mame1ka, M., 1985. Interglacja1 eemski i wczesny Vistulian w Zgierzu-Rudunkach na Wy _zynie qódzkiej (The Eemian Interglacial and early Vistulian atZgierz-Rudunki in the qód�z Plateau). Acta Geografica Lodziensia 53, 1e75 (inPolish with English summary).

Karlsdóttir, L., Hallsdóttir, M., Thórsson, Ae. T., Anamthawat-Jónsson, K., 2008.Characteristics of pollen from natural triploid Betula hybrids. Grana 47, 52e59.

Klatkowa, H., Za1oba, M., 1991. Kszta1towanie budowy geologicznej i rze�zbypo1udniowego obrze _zenia Basenu Uniejowskiego (The formation of thegeological structure and relief of the southern margin of the Uniejów Basin). In:Stankowski, W. (Ed.), Przemiany �srodowiska geograficznego obszaru Konin-Turek. Inst. Bad. Czwart. UAM, Pozna�n, pp. 33e44 (in Polish with Englishsummary).

Klotz, S., Guiot, J., Mosbrugger, V., 2003. Continental European Eemian and early Wür-mian climate evolution: comparing signals using different quantitative reconstruc-tion approaches based on pollen. Global and Planetary Change 36, 277e294.

Ko1aczek, P., Fia1kiewicz-Kozie1, B., Karpi�nska-Ko1aczek, M., Ga1ka, M., 2010. The lasttwo millennia of vegetation development and human activity in the Orawa-Nowy Targ Basin (south-eastern Poland). Acta Palaeobotanica 50, 133e148.

Krupi�nski, K.M., Nory�skiewicz, A.M., Nalepka, D., 2004. Taxus baccata L. e Yew. In:Ralska-Jasiewiczowa, M., Wasylikowa, K., Tobolski, K., Madeyska, E.,Wright Jr., H.E., Turner, C. (Eds.), Late Glacial and Holocene History of Vegetationin Poland Based on Isopollen Maps. W. Szafer Institute of Botany, PolishAcademy of Sciences, Kraków, pp. 209e215.

Krupi�nski, K.M., Kotarbi�nski, J., Skompski, S., 2006. Osady jeziorne interglacja1ueemskiego w Leszczynie eWysoczyzna P1ocka (Lacustrine sediments of Eemianinterglacial at Leszczyno (P1ock Upland e Central Poland)). Przeglad Geo-logiczny 54, 632e638 (in Polish with English summary).

Kühl, N., Litt, T., 2003. Quantitative time series reconstruction of Eemian temper-ature at three European sites using pollen data. Vegetation History andArchaeobotany 12, 205e214.

Kupryjanowicz, M., Nalepka, D., Walanus, A., Balwierz, Z., Bi�nka, K.,Granoszewski, W., Krupi�nski, K.M., Malkiewicz, M., Nita, M., Nory�skiewicz, B.,Pidek, I.A., Tobolski, K., Winter, H., 2011. Eemska historia ro�slinno�sci Polski napodstawie map izopolowych e pierwsza informacja o projekcie. In:Karasiewicz, T.M., Nory�skiewicz, A.M., Hulisz, P., Winter, H. (Eds.), V PolskaKonferencja Paleobotaniki Czwartorzedu, Cz1owiek i jego wp1yw na �srodowiskoprzyrodnicze w przesz1o�sci i czasach historycznych. Górzno, 13e17 czerwca2011, pp. 30e31 (in Polish).

Kupryjanowicz, M., 2008. Vegetation and climate of the Eemian and Early Vistulianlakeland in northern Podlasie. Acta Palaeobotanica 48, 3e130.

Lata1owa, M., Tobolski, K., Nalepka, D., 2004. Cyperaceae e Sedge family. In: Ralska-Jasiewiczowa, M., Wasylikowa, K., Tobolski, K., Madeyska, E., Wright Jr., H.E.,Turner, C. (Eds.), Late Glacial and Holocene History of Vegetation in PolandBased on Isopollen Maps. W. Szafer Institute of Botany, Polish Academy ofSciences, Kraków, pp. 283e292.

Lindner, L., 2005. Nowe spojrzenie na liczbe, wiek i zasiegi zlodowace�n�srodkowopolskich w po1udniowej cze�sci �srodkowo-wschodniej Polski (A newlook at the number, age and extent of the Middle Polish Glaciations in thesouthern part of central-eastern Poland). Przeglad Geologiczny 53, 145e150 (inPolish with English summary).

Litt, T., Junge, F.W., Böttger, T., 1996. Climate during the Eemian in north-centralEurope e a critical review of the palaeobotanical and stable isotope datafrom central Germany. Vegetation History and Archaeobotany 5, 247e256.

Litt, T., 1994. Paläoökologie, Paläobotanik und Stratigraphie des Jungquartärs imnordmitteleuropäischen Tiefland. Dissertationes Botanicae 227, 1e185.

Lord, T.R., Travis, H.J., 2011. Ferns and Fern Allies of Pennsylvania. Pinelands Press.Makra, L., Juhász, M., Béczi, R., Borsos, E., 2005. The history and impacts of airborne

Ambrosia (Asteraceae) pollen in Hungary. Grana 44, 57e64.Malkiewicz, M., 2003. Palynology of biogenic sediments of the Eemian Interglacial

at Bieganin near Kalisz, Central Poland. Geological Quarterly 47, 367e372.

P. Kołaczek et al. / Quaternary International 268 (2012) 9e2020

Malkiewicz, M., 2010. Early Vistulian vegetation history and climate change atGutów (Wielkopolska Lowland) from pollen analysis. Geological Quarterly 54(3), 357e366.

Mamakowa, K., 1988. Pollen stratigraphy of the Eemian and adjoing glacial depositsbased on continuous sequences in Poland. Bulletin, Polish Academy of Sciences,Earth Science 36 (3e4), 299e307.

Mamakowa, K., 1989. Late Middle Polish Glaciation, Eemian and Early Vistulianvegetation at Imbramowice near Wroc1aw and the pollen stratigraphy of thispart of the Pleistocene in Poland. Acta Palaeobotanica 29 (1), 11e176.

Matuszkiewicz, W., 2005. Przewodnik do oznaczania zbiorowisk ro�slinnych Polski.Wydawnictwo Naukowe PWN (in Polish).

Menke, B., Tynni, R., 1984. Das Eeminterglazial und das Weichselfrühglazial vonRederstall/Dithmarschen und ihre Bedeutung für die mitteleuropäische Jung-pleistozän-Gliederung. Geologische Jahrbuch, Ser. A 76, 3e120.

Michalska-Hejduk, D., Kope�c, D., 2002. Lemno minoris-Salvinietum natantis iHydrocharitetum morsus-ranae z udzia1em Salvinia natans w starorzeczach Sanui propozycje ich ochrony (Lemno minoris-Salvinietum natantis and Hydro-charitetum morsus-ranae with Salvinia natans in old river-beds of the river San(Poland) and proposals of their protection). Fragmenta Floristica et GeobotanicaPolonica 9, 319e328 (in Polish).

Miros1aw-Grabowska, J., 2009. Evolution of palaeolake environment in Polandduring the Eemian Interglacial based on oxygen and carbon isotope data fromlacustrine carbonates. Quaternary International 207, 145e156.

Moore, P.D., Webb, J.A., Collinson, M.E., 1991. Pollen Analysis. Blackwell ScientificPublications, Oxford.

Müller, U.C., Sánchez-Go�ni, M.F., 2007. Vegetation dynamics in Southern Germanyduring Marine Isotope Stage 5 (w130 to 70 kyr ago). In: Sirocko, F.,Claussen, M., Sánchez-Go�ni, M.F., Litt, T. (Eds.), 2007. The Climate of PastInterglacials, Developments in Quaternary Science, vol. 7. Elsevier, Amsterdam,pp. 277e287.

Müller, U.C., Pross, J., Bibus, E., 2003. Vegetation response to rapid climate change inCentral Europe during the past 140,000 yr based on evidence from the Für-amoos pollen record. Quaternary Research 59, 235e245.

Müller, U.C., Klotz, S., Geyh, M.A., Pross, J., Bond, G.C., 2005. Cyclic climate fluctu-ations during the Last Interglacial in Central Europe. Geology 33, 449e452.

Nalepka, D., Walanus, A., 2003. Data processing in pollen analysis. Acta Palae-obotanica 43, 125e134.

Nory�skiewicz, A.M., 2001. Historia cisa we Wierzchlesie na tle rozwoju post-glacjalnej ro�slinno�sci (The history of yew in Wierzchlas against the postglacialhistory of vegetation). Ph.D. thesis. Archive of Department of Biology and EarthSciences, N. Copernicus University in Toru�n, Poland. (in Polish).

Novenko, E.Y., Seifert-Eulen, M., Boettger, T., Junge, F.W., 2008. Eemian and EarlyWeichselian vegetation and climate history in Central Europe: a case studyfrom the Klinge section (Lusatia, eastern Germany). Review of Palaeobotany andPalynology 151, 72e78.

Punt, W., Hoen, P.P., 2009. The Northwest European Pollen Flora, 70, Asteraceae e

Asteroideae. Review of Palaeobotany and Palynology 157, 22e183.Reille, M., Andrieu, V., de Beaulieu, J.-L., Guenet, P., Goeury, C., 1998. A long pollen

record from Lac du Bouchet, Massif, central France: for the period ca. 325 to100 ka BP (OIS 9c to OIS 5e). Quaternary Science Reviews 17, 1107e1124.

Reille, M., 1992. Pollen et spores d’Europe et d’Afrique du Nord. Laboratoire deBotanique Historique et Palynologie, Marseille (in French).

Roman, M., Balwierz, Z., 2010. Eemian and Vistulian pollen sequence at Kub1owo(Central Poland): implications for the limit of the Last Glacial Maximum.Geological Quarterly 54, 55e68.

�Srodo�n, A., 1990. Buk w historii lasów Polski (Beech in the forest history of Poland).In: Bia1obok, S. (Ed.), Buk zwyczajny e Fagus sylvatica L. Polska Akademia Nauk,Instytut Dendrologii, PWN, Warszawa-Pozna�n (in Polish).

Stachowicz-Rybka, R., Granoszewski, W., Hrynowiecka-Czmielewska, A., 2009.Quaternary environmental changes at Starunia palaeontological site andvicinity (Carpathian region, Ukraine) based on palaeobotanical studies. AnnalesSocietatis Geologicorum Poloniae 79, 279e288.

Stankowski, W., Nita, M., 2004. Stratigraphy of Late Quaternary deposits and theirneotectonic record in the Konin area, Central Poland. Geological Quarterly 48,23e34.

Stockmarr, J., 1971. Tablets with spores used in absolute pollen analysis. Pollen etSpores 13, 615e621.

Tobolski, K., 1991. Biostratygrafia i paleoekologia interglacja1u eemskiego i zlodo-wacenia Wis1y regionu koni�nskiego (Biostratigraphy and palaeoecology of theEemian Interglacial and the Vistulian Glaciation of the Konin Region (Poland)).In: Stankowski, W. (Ed.), Przemiany �srodowiska geograficznego obszaru Konin-Turek. Instytut Bada�n Czwartorzedu, Pozna�n, pp. 45e87 (in Polish).

Turner, C., 2002. Formal status and vegetational development of the Eemian inter-glacial in Northwestern and Southern Europe. Quaternary Research 58, 41e44.

van Geel, B., Bohncke, S.J.P., Dee, H., 1981. A palaeoecological study of an upper LateGlacial and Holocene sequence from “De Borchert”, The Netherlands. Review ofPalaeobotany and Palynology 31, 367e448.

Velichko, A.A., Novenko, E.Y., Zelikson, E.M., Boettger, T., Junge, F.W., 2007.Comparative analysis of vegetation and climate changes during the EemianInterglacial in Central and Eastern Europe. In: Sirocko, F., Claussen, M., Sánchez-Go�ni, M.F., Litt, T. (Eds.), The Climate of Past Interglacials. Developments inQuaternary Science, vol. 7. Elsevier, Amsterdam, pp. 255e264.

Walkling, A., Coope, G.R., 1996. Climatic reconstructions from the Eemian/EarlyWeichselian transition in Central Europe based on coleopteran record fromGröbern, Germany. Boreas 25, 145e159.

Wohlfarth, B., Skog, G., Possnert, G., Holmquist, B., 1998. Pitfalls in the AMSradiocarbon-dating of terrestrial macrofossils. Journal of Quaternary Science 13(2), 137e145.

Zagwijn, W.H., 1996. An analysis of Eemian climate in Western and Central Europe.Quaternary Science Reviews 15, 451e469.