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Quaternary International 292 (2013) 124e135

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Quaternary International

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Late Glacial and Early Holocene development of lakes in northeastern Poland inview of plant macrofossil analyses

Mariusz Ga1ka a,*, Marcin Sznel b

aAdam Mickiewicz University, Department of Biogeography and Palaeoecology, ul. Dziegielowa 27, 61-680 Pozna�n, Polandb Suwalski Landscape Park, MalesowiznaeTurtul, 16-404 Jeleniewo, Poland

a r t i c l e i n f o

Article history:Available online 21 November 2012

* Corresponding author.E-mail addresses: galka@amu.edu.pl (M. Ga1ka), m

1040-6182/$ e see front matter � 2012 Elsevier Ltd ahttp://dx.doi.org/10.1016/j.quaint.2012.11.014

a b s t r a c t

The Late Glacial history of the development of seven extant or former, now sediment-filled, lakes innortheastern Poland was investigated based on high-resolution plant macrofossil analyses and AMSradiocarbon dating. According to palaeobotanical data and AMS dating, the accumulation of organicsediments began during the Bölling period. The bottom sediments of all studied lakes contain plantmacrofossils typical of Late Glacial vegetation, such as Betula nana, Dryas octopetala, Potamogetonfiliformis, and Potamogeton alpinus. The first plants to occur in the lakes were Chara sp., P. filiformis andBatrachium sp. The Late Glacial sediments included rarely encountered macrofossils of Sphagnum teres. Atthe beginning of the Holocene, the water level in the studied northeastern Polish lakes decreasedsignificantly. This was manifested in a change in the type of sediment, transformation of small lakes intopeatland in places, and expansion of typical (indicator) plants of shallow waters.

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1. Introduction

The contemporary landscape of northeastern Polandwasmainlyformed by the Scandinavian Ice Sheet (Ber, 1974, 2000, 2006;Krzywicki, 2002; Marks, 2002). The ice sheet retreated from NEPoland at approximately 14 600 cal. BP (Rinterknecht et al., 2006). Italso contributed to the development of numerous lakes. After theretreat of the ice sheet and with the melting out of blocks of deadice, hundreds of basins of various shapes and size contained water.The process of biogenic sediment accumulation began in the basins.The changing climate at the turn of the Pleistocene caused varia-tions in temperature and humidity, as well as the development ofsoil processes. One result of such changes was plant migration.In accordance with the vegetation succession transformationsobserved during consecutive glacialeinterglacial cycles (Iversen,1964; Tobolski, 1976; Birks, 1986; Lang, 1994), pioneer speciestypical of shrub tundra were the first to appear, followed by Borealelements and deciduous forests during the Holocene climaticoptimum.

The accumulation of biogenic sediments in lakes and the sedi-ments’ ability to preserve plant and animal fossils permits palae-oecological reconstructions (Tobolski, 2000; Birks and Birks, 2006).One of the most important methods applied in palaeoecological

arcinsznel@wp.pl (M. Sznel).

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studies is plant macrofossil analysis. Identification of such remainsenables estimates of the age of biogenic sediments (Velichkevichand Zastawniak, 2006, 2008; Ammann et al., 2007; Koff andTerasmaa, 2011). Analysis of bottom layers of sediments in termsof the content of plant macro- and micro-fossils permits recon-struction of the first vegetation to appear in areas uncovered byretreating ice sheets (van Geel et al., 1989; Birks, 1993, 2003; Allenand Huntley, 1999; Bennike, 2000; Wohlfarth et al., 2006; Vasariet al., 2007; Heikkilä et al., 2009) or mountain glaciers (Lang,1993; Wick, 1994; Tobolski and Ammann, 2000). Due to theirindicator properties, the aquatic, peatland, and terrestrial plantmacrofossils included in biogenic sediments enable the recon-struction of palaeoclimate, water level variations, and the rate ofexpansion of individual plant species (Hannon and Gaillard, 1997;Birks and Ammann, 2000; Birks, 2003, 2007; Bennike et al., 2004;Väliranta, 2006; Bos et al., 2007; Wohlfarth et al., 2007; Binneyet al., 2009). An additional advantage of the application of plantmacrofossil analysis is the possibility of the identification of a givenplant at the species level, which is rarely possible through pollenanalysis (Tobolski, 2000; Tobolski and Ammann, 2000). Numerousspecies belonging to the genera Potamogeton or Carex are particu-larly useful for interpretation of issues discussed in this paper. Themajor objectives of the present study were to reconstruct the initialstage of development of local vegetation and to determine the timeof the commencement of the deposition of organic sedimentswithin seven selected water bodies in northeastern Poland.Undertaking such investigations was encouraged by the lack of

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existing analyses of Late Glacial and Early Holocene sediments interms of the presence of plant macrofossils. The data included inthis paper constitute part of the results of palaeoecological studiesfocusing on the postglacial history of the vegetation and climate inthis region of Europe. These data represent the first results ofmacroscopic plant fossil analyses of Late Glacial sediments con-ducted in this area. Previous palaeobotanical studies in north-eastern Poland have been restricted to palynological analyses(O1tuszewski, 1937; Kupryjanowicz, 2007; Lauterbach et al., 2010).Studies including analysis of plant macrofossils in Late Glacial andEarly Holocene sediments have been conducted in Lithuania(Stan�cikait _e et al., 2008, 2009; Gaidamavi�cius et al., 2011) andwestern Belarus (Stan�cikait _e et al., 2011).

2. Study area

The study area is located in Suwalski Landscape Park (NE Poland,Fig. 1), in the Polish section of the Lithuanian Lakeland of theWschodniosuwalskie Lakeland mesoregion (Kondracki, 1998). Thelandscape of northeastern Poland developed as a result of thepresence of the Pleistocene ice sheet in this area. This region’scontemporary land relief developed in the final stage of the Pleis-tocene and during the Holocene (Ber, 2000; Krzywicki, 2002). It isdistinguished by high hills (highest elevation 275 m a.s.l.), deep

Fig. 1. Stud

river valleys, and lake channels surrounded by steep scarps ofplateaus. Local relief reaches 100 m. The lowest point is LakePowstawelek at 146 m a.s.l. The study area is located in the Suwa1kiclimatic region of Podlasie, which is distinguished by a transitionaltemperate climate with a clear influence of continental climate,with a mean July temperature of 17 �C and a mean Januarytemperature of 5 �C. Themean annual air temperature is lower thanthat of western Poland by 2 �C (Lorenc, 2005). This results in anextended presence of ice cover on lakes, although winters withoutfull ice cover have been reported on Lake Hancza (Górniak andPekala, 2001). The total mean annual precipitation is more than650 mm, with the maximum in July and the minimum in February.Consequently this region is characterized by short summers, a shortvegetation period of less than 200 days, and the shortest periodwith no frost in lowland Poland (Stopa-Boryczka andMartyn,1985).The climatic specificity of northeastern Poland translates intoa specific character of its vegetation. The study area is located in theCentral European LowlandeUpland Province, Northern Division,Suwa1kieAugustów Region (Szafer and Paw1owski, 1972). Themajority of the forest complexes in the landscape park are mesicmixed Coryleto-Piceatum forests and spruce mixed coniferousCalamagrostio arundinaceae-Piceetum forests (Soko1owski, 1973).Sediments for the analyses were sampled from peatlands. Two ofthese peatlands, constituting sites I and II, developed at lake shores.

y site.

M. Gałka, M. Sznel / Quaternary International 292 (2013) 124e135126

A fen developed at the southern shore of Lake Kojle with a depth of33 m. This fen is currently overgrown with Alnus glutinosa andFrangula alnus. The rush zone of the lake includes Cladiummariscus,Thelypteris palustris, and Carex paniculata. A transitional bogdeveloped around Lake Linówek with a maximum depth of 4.2 m.At the sampling site, the occurrence of Sphagnum teres, Oxycoccuspalustris, and Scheuchzeria palustris, among other species, wasobserved. The studied peatland lakes (sites IIIeVII) are located tothe east of Lake Ha�ncza. These lakes are channel-shaped anddistributed parallel to each other and to the significantly largerbasin of Lake Boczniel (Fig. 1). Four sites (III, IV, V, VI) are peatlandswith Sphagnum mosses, including Sphagnum magellanicum,Sphagnum fallax, Sphagnum palustre, and S. teres, and the vascularplant species O. palustris, Drosera rotundifolia, and Andromedapolifolia. One site (VII) does not currently include plants with peat-forming abilities. The peatlands constituting sites III, IV, V, and VIwere meliorated and subjected to peat exploitation, as demon-strated by the presence of numerous peat pits, which have beencolonised by aquatic wetland vegetation, including Sphagnummosses. The drilling and sampling sites are designated by thefollowing coordinates:

I) 54�16021.3600N, 22�53021.8000E; II) 54�13024.3000N,22�50029.9900E; IIIA) 54�15030.900N, 22�49030.400E; IIIB)54�15026.7700N, 22�49023.2200E; IV) 54�15012.900N, 22�49020.900E;V) 54�15016.700N, 22�49048.900E; VI) 54�15028.8200N, 22�49050.6700E;VII) 54�15034.8800E, 22�5007.1800E.

3. Methods

The field research was conducted in the period extending fromJune to August 2010. For the drilling, a manual Instorf corer witha diameter of 5 cm and length of 50 cm was used. The location of

Fig. 2. Diagram of plant macrofossils. Site I. 1 e sand with silt, 2 e fine detritus gyttja. Del e leaf, n e needle, fs e fruits scale, o e oospore, per e periderm, end e endocarp.

the sampling sites is presented in Fig. 1. At all investigated sites thedrilling reached mineral sediments, demonstrating the bottom ofbiogenic sediments and the beginning of the existence of waterbodies in the area. The oldest sediments of seven cores weresampled for detailed laboratory analyses. Limnic-peat sedimentwas placed in PVC tubes. In the laboratory, sediments wereunpacked, cleaned, and further divided into slices of 1 cm thickness,using a surgical scalpel. The samples were washed with warmrunning water on sieves with a 0.25-mm mesh. Generativemacrofossils (seeds, fruits, fruit scales) were identified usinga Nikon stereoscopic microscope under 10e200� magnification.Vegetative parts (leaves of mosses) were identified with a ZeissAMPLIVAL light microscope under 400� and 1000�magnification.Determination of individual plant fossils was based on thefollowing keys: Beijerinck (1976), Berggren (1968, 1981), Grosse-Brauckmann (1974), Grosse-Brauckmann and Streitz (1992),Tobolski (2000), and Velichkevich and Zastawniak (2006, 2008).The results are based on one-centimetre resolution botanicalcomposition analyses of 705 samples.

The results of the plant macrofossil analyses are illustrated indiagrams (Figs. 2e9), prepared by using C2 graphic software(Juggins, 2003). On the left side of the figures the remains of treesand shrubs are presented. The order of the remaining taxa followsthe vegetation succession from lake to peatland and, finally toupland herb vegetation. Fossil diaspores are indicated in absolutenumbers, and the mosses (e.g., Drepanocladus sp. and S. teres) arepresented as the percentages of the total sediment sample volume(25 cm3).

To determine the beginning of the accumulation process in thestudy area, macrofossils of terrestrial plants from four sampleswere selected for AMS radiocarbon dating. The macrofossils werestored following the recommendations of Walanus and Goslar

scription of plant remains: f e fruit, fb e fruit biconvex, ft e fruit trigonous, s e seed,

M. Gałka, M. Sznel / Quaternary International 292 (2013) 124e135 127

(2009). 14C dating was performed in the Poznan RadiocarbonLaboratory. The resulting conventional radiocarbon dates werecalibrated using OxCal 4.1 software (Bronk-Ramsey, 2009). Meanage values expressed in calendar years are based on tables preparedby Walanus and Goslar (2009). Zonal names such as Allerød orYounger Dryas refer to data defined by Litt et al. (2001).

4. Results

4.1. Lithostratigraphy and chronology

The details of the sediment lithostratigraphy are presented inTable 1. Material for dating was selected from one-centimetre-thickcore slices. Four samples taken from Lake Kojle (Site I) and LakeLinówek (Site II) were dated (Table 2). Two (Lake Kojle, 630e631 cm, and Lake Linówek, 678e679 cm) were selected in orderto date the beginning of detritus gyttja accumulation. The resultsare notably similar: in Lake Kojle 11 650 � 70 14C BP and in LakeLinówek 11 690 � 60 14C BP. Two further samples (Lake Kojle494e495 cm, Lake Linówek 630e631 cm) represent layers of plantremains suggesting significant climate warming. Dating in bothlakes provided the same age: approx. 9510 � 50 14C BP.

Table 1Lithostratigraphic description of the sediment sequence.

Site Description of sediments

I 650e642 cm silt642e495 cm detritus-calcareous gyttja

II 800e768 cm sand with silt768e683 cm silt683e600 cm fine detritus gyttja

III A. 650e646 cm sand646e640 cm silt640e600 cm fine detritus gyttjaB. 1050e1043 cm sand1043e998 cm silt998e950 cm fine detritus gyttja

IV 290e275 cm sand and silt275e260 cm silt,260e206 cm fine detritus gyttja206e166 cm Sphagnum-herbaceous peat166e150 cm fine detritus gyttja

V 700e695 cm sand with silt695e682 cm fine detritus gyttja682e650 cm brown moss-herbaceous peat

VI 700e690 cm sand with silt690e617 cm fine detritus gyttja617e600 cm brown moss-herbaceous peat

VII 300e287 cm brown moss-herbaceous peat287e283 cm fine detritus gyttja283e277 cm brown moss-herbaceous peat277e258 cm fine detritus gyttja258e250 cm brown moss-herbaceous peat

Table 2Radiocarbons dates.

Site/depth(cm)

Dated material Lab. No. 14C date Calibratedrange 94.5%

I 494e495 Fruits and fruitsscales of Betula sp.

Poz-38823 9510 � 50 BP 11 086e10 600

I 630e631 Seed of Pinussylvestris andfruits of Betula sp.

Poz-39560 11 650 � 70 BP 13 706e13 326

II 607e608 Seed of Pinussylvestris andfruits and fruitsscales of Betula sp

Poz-35958 9510 � 60 BP 11 090e10 592

II 678e679 Seed ofArctostaphylosuva-ursi

Poz-35959 11 690 � 60 BP 13 730e13 380

4.2. Plant macrofossil analysis

4.2.1. Site IFour phases of the development of local vegetation were

distinguished (Fig. 2). The first phase (I-1, 650e595 cm) wasdominated by tree macrofossils. This phase included fruits of Betulapubescens, Betula sec. Alba and seeds and a needle from Pinus syl-vestris. Aquatic species occurred at this site, including Potamogetonfiliformis, Batrachium sp., and Chara sp. Fruits of sedges were alsofound. In the second phase (I-2, 595e545 cm), only a single Betulasec. Alba fossil was recorded. At a depth of 578 cm, one fruit ofBetula nana was found. Three aquatic species, Batrachium sp.,Potamogeton sp., and Chara sp. (with the highest oospore frequencyin the entire profile), occurred in phase I-2. In the third phase (I-3,545e518 cm), the number of Chara sp. oospores decreased signif-icantly, and B. pubescens fossils appeared. Aquatic species wererepresented byNajas marina and Batrachium sp. Double-sided fruitsof sedges were present. The fourth phase (I-4, 518e495 cm) wasdistinguished by a significant increase in macrofossils. Trees wererepresented by B. pubescens and Pinus sylvestris. In the first part ofthe fourth phase, Potamogeton natans, Potamogeton praelongus, andHippuris vulgaris appeared and were followed somewhat later byCeratophyllum demersum. The middle part included Nymphaea alba,Potamogeton sp., and for the first time, Typha sp. and Carex pseu-docyperus. The uppermost part was dominated by P. natans andH. vulgaris.

4.2.2. Site IIFour phases of the development of local vegetation were

distinguished in the profile of site II (Fig. 3). The first phase (II-1,800e683 cm) included only sporadic macrofossils. The occurrenceof Dryas octopetala and O. palustris was recorded. At a depth of711 cm, a P. filiformis endocarp was found. In the second phase(II-2, 683e653 cm), in the lower part, fruits and fruit scale ofB. nana were present. The aquatic Potamogeton sp. also occurred.The lower part included Arctostaphylos uva-ursi seeds. The thirdphase (II-3, 653e623 cm) was distinguished by the occurrence ofB. nana, Selaginella selaginoides, and the aquatic species Potamo-geton filifomis, Batrachium sp., and Nitella sp. In the fourth phase(II-4, 623e600 cm), Betula sec. Alba, Chara sp. and Potamogetonlucens appeared simultaneously and Typha sp. somewhat later.N. alba, Potamogeton crispus, and P. natans were also observed inthis phase.

4.2.3. Site IIIThe botanical composition at Site III was determined in the

bottom layer of sediments sampled from two cores. In contrast tothe other lakes analysed, the limnic sediments at this site did notinclude a great number of macrofossils. At site three (IIIA) in sedi-ments from a depth of 600e650 cm, the presence of macrophyteswas detected (Fig. 4). In the first phase (IIIA-1, 650e615 cm), Charasp. oospores, Menyanthes trifoliata seeds, and Pinus sylvestrismacrofossils were encountered. In the second phase (IIIA-2, 615e600 cm), Batrachium sp., Potamogeton sp., and Chara sp. occurred.At site two (Fig. 5) in limnic sediments from a depth of 950e1050 cm in the first phase (IIIB-1, 1050e998 cm), Chara sp.oospores and a Betula sec. Alba fruit were found. In the secondphase (IIIB-2, 998e950 cm), B. pubescens, Pinus sylvestris, and B.nana fruits occurred at depths of 982 cm and 966 cm. The upperpart included M. trifoliata.

4.2.4. Site IVBased on the plant macrofossil analysis of sediments from

a depth of 150e290 cm, four phases of the development of localvegetationwere distinguished (Fig. 6). In phase IV-1 (290e258 cm),

Fig. 3. Diagram of plant macrofossils. Site II. Description of plant remains as in Fig. 2.

Fig. 4. Diagram of plant macrofossils. Site IIIA. Description of plant remains as in Fig. 2. Fig. 5. Diagram of plant macrofossils. Site IIIB. Description of plant remains as in Fig. 2.

M. Gałka, M. Sznel / Quaternary International 292 (2013) 124e135128

Fig. 6. Diagram of plant macrofossils. Site IV. Description of plant remains as in Fig. 2.

Fig. 7. Diagram of plant macrofossils. Site V. Description of plant remains as in Fig. 2.

M. Gałka, M. Sznel / Quaternary International 292 (2013) 124e135 129

M. Gałka, M. Sznel / Quaternary International 292 (2013) 124e135130

the occurrence of P. natans endocarps and Chara sp. oospores wasrecorded. Fruits and fruit husks of B. nana were also present. Inphase IV-2 (258e240 cm), numerous macrofossils of macrophytesfrom the genus Potamogeton occurred, which included P. filiformis,P. natans, Potamogeton pusillus, and P. praelongus. The lower partincluded Chara sp. oospores. B. nana also occurred in this phase.Phase IV-3 (200e166 cm) was distinguished by the macrofossiloccurrence of peatland plants, including Comarum palustre, and M.trifoliata, and the sedges Carex rostrata and Carex riparia. Thecontribution of S. teres leaves in the peat reached 100%. Theoccurrence of fruits and fruit scales from B. nana proves the pres-ence of this species in this period. In phase IV-4 (166e150 cm), theoccurrence of the following macrophytes was recorded: P. natans,P. pusillus, Nuphar pumila, and Batrachium sp., in addition to thealgae Chara sp. and Nitella sp. The sediment also included B. nanaand B. pubescens macrofossils.

4.2.5. Site VBased on the occurrence of macrofossils in sediments from

a depth of 700e650 cm, three phases of the development of localvegetation were distinguished (Fig. 7). In the first phase V-1 (700e693 cm) in limnic sediment, B. nana fruits and seeds were found. Inthe second phase V-2 (693e682 cm) in coarse detritus gyttja, N.alba seeds and a P. natans endocarp, as well as B. pubescensmacrofossils and a Pinus sylvestris needle, were recorded. The thirdphase V-3 (682e650 cm) is characterized by the mass appearanceof the moss Drepanocladus sp.

Fig. 8. Diagram of plant macrofossils. Site VI.

4.2.6. Site VISediments sampled from a depth of 600e700 cmwere analysed

at this site. Three phases of the development of local vegetationwere distinguished (Fig. 8). In phase VI-1 (700e676 cm), Chara sp.,as well as Potamogeton sp. and Myriophyllum sp., occurred. Thesediment also included B. nana fruits and fruit scales. In phase VI-2(676e618 cm), macrofossils of Pinus sylvestris (needles) and Betulasec. Alba (fruits) were found. Macrophytes were represented byP. natans and Chara sp. The peat sediment representing phase IV-3(618e600 cm) was dominated by the moss Drepanocladus sp.

4.2.7. Site VIIBased on the occurrence of plant macrofossils found in the

bottom layers of sediments at site V, five phases of the developmentof local vegetationwere distinguished (Fig. 9). The first phase, VII-1(300e287 cm), contained the peatland plants M. trifoliata andC. palustre and sedges, including C. rostrata. In the uppermost partof the phase, the occurrence of leaves of brown mosses wasrecorded. Trees were represented by Pinus sylvestris and Betula sec.Alba fossils. In the second phase, VII-2 (287e283 cm), Potamogetonalpinus and Batrachium sp. appeared. B. nana was also present. Inthe third phase, VII-3 (283e276 cm), Arctostaphylos uva-ursi, Stel-laria sp., and brownmosses occurred. In phase VII-4 (276e258 cm),numerous B. nana fossils were found. Endocarps of P. alpinus andP. pusillus and fruits of Batrachium sp. also occurred. Severalsamples in the middle part of the phase included S. teres leaves.Additionally, Arctostaphylos uva-ursi seeds were present. Phase VII-

Description of plant remains as in Fig. 2.

Fig. 9. Diagram of plant macrofossils. Site VII. Description of plant remains as in Fig. 2.

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5 (258e250 cm) was poor in macrofossils. Only one C. palustre fruit,one Betula sec. Alba scale, and a scarce amount of Bryopsida werefound.

5. Discussion

5.1. The first stage of lake development e Bølling

The origin of the studied basins, where lakes existed for a periodin the past, is related to the presence of an ice sheet in this area. Inthe course of the melting out of blocks of dead ice, the basins werefilled with meltwater. This type of lake formation is very commonin recently deglaciated areas (Pasierbski, 1973; Starkel, 1977; Galon,1982; Choi�nski, 1995; Kozarski, 1995). Sediment accumulation inlakes of northeastern Poland began during the Oldest Dryas (Ber,2006). The sediments typical of this period are silts and sands.Organic sediment accumulation began in the Bølling period (Ber,2006). The beginning of fine detrital gyttja accumulation in thisperiod has been confirmed by dating performed in the bottomlayers of sediment from Lakes Kojle and Linówek. Lake Kojle, in thepast connectedwith thewater body of Lake Perty (Ga1ka, 2012), andLake Linówek sediment deposition commenced approx. 13 500 cal.BP (Ga1ka, unpublished data). The bottom layer of sediments, in theform of clay with sand/gravel, in Lake Ha�ncza was dated to12 900 cal. BP (Lauterbach et al., 2010). The later occurrence of themelting out of ice in Lake Ha�ncza resulted from its geometry(depth: 108.5 m, water surface area: 303 ha). Bottom sedimentsfrom Lake Wigry, which is located approx. 20 km from the studiedsites, were dated to approx. 13 700 cal. BP (Zawisza andSzeroczy�nska, 2007). A similar time of the commencement ofsediment accumulation was determined for lakes in Latvia(Stan�cikait _e et al., 2008, 2009) and Estonia (Saarse et al., 2009;Amon et al., 2010). These data suggest that the timing of the initi-ation of the lake development in northwestern Poland was

correlated with the BøllingeAllerød. Data concerning the durationof the deposition of organicmatter on a lake bottom is not sufficientto reach a firm conclusion regarding the beginning of the devel-opment of the basin. Development of the landform in which a lakeoriginates could occur at a time different than the filling of thebasinwith water and the beginning of sediment accumulation. Thisis particularly important in the case of small water bodies. A two-stage character of biogenic sediment accumulation in lakes andpeatlands in northern Poland was observed by Tobolski (2006). Ina number of these sites, the first accumulation occurred during theBøllingeAllerød and in others at the beginning of the Holocene.

The lack of radiocarbon dating data for the bottom layers ofsediments of lakes located to the east of Lake Ha�ncza does not allowdetailed determination of the commencement of biogenic sedi-ment accumulation. The form of the bottom sections of sedimentsand the results of plant macrofossil analyses are useful for thispurpose. According to such data, the bottom layers of sediments inthe five studied lakes clearly developed during the Late Glacial. Inthree of these sites (III, V, and VI), the bottom section of the sedi-ments included a layer of silts underlying detrital gyttja. Sucha distribution of the layers in the bottom sections of sediments istypical of lake development determined to have begun during theBølling (Ber, 2006). Commencement of sediment deposition in theLate Glacial is suggested by numerous plant fossils. The sediments’common occurrence is typical of the period. In the bottom layers ofsediments in all seven lakes, B. nana fruits and fruit scales werefound. B. nana currently occurs commonly in boreal and subarcticzones, e.g. in northern Europe and in the Alpine zone of mountains(Pieko�s-Mirkowa and Mirek, 2003). In Central Europe, this speciesis therefore commonly recognized as a glacial relic (Korna�s andMedwecka-Korna�s, 2002). B. nana is a good palaeoclimate indi-cator, and its presence in sediments usually represents a coldperiod preceding the Holocene climate warming (Godwin, 1975;Lang, 1994). However, macrofossils of this species are sometimes

M. Gałka, M. Sznel / Quaternary International 292 (2013) 124e135132

found at sites outside the area of its contemporary occurrence inthe layer of sediments deposited in the Holocene. For example, B.nana macrofossils have been found at sites in northern Germany(Overbeck, 1975) and central Poland in the Tuchola Forest-peatlandin the Wierzchlas reserve (Ga1ka, unpublished data) and thevicinity of Brodnica (Stachowicz-Rybka, unpublished data). B. nanawas one of the main components of tundra assemblages occurringin this part of Europe in the Late Glacial (Kupryjanowicz, 2007;Stan�cikait _e et al., 2008;Wacnik, 2009). This species is recognised asa plant growing near the ice sheet terminal zone in this period(Binney et al., 2009). The presence of B. nana fossils in Late Glacialsediments has also been recorded in other parts of Poland (e.g.,Kubiak-Martens, 1998; Schubert, 2003; Drzymulska, 2006). Addi-tionally, macrofossils of this species are found quite frequently inLate Glacial sediments in other parts of Europe (Lang, 1994;Wohlfarth et al., 2002). The beginning of sediment accumulation inthe Late Glacial is also demonstrated by presence (in sediments) ofthe macrofossils of several macrophytes. The studied lakes con-tained endocarps of P. filiformis, P. alpinus, P. pusillus, and N. pumila.The presence of macrofossils of these plants in sediments is anindicator of cool climate (Szafer, 1954; Velichkevich and Zastawniak,2006, 2008; Mortensen et al., 2011). Their contemporary distribu-tion is mainly concentrated in northern Europe (Czubi�nski, 1950;Walter and Straka, 1970; Hultén and Fries, 1986; Zaleska-Ga1osz,2001; Pieko�s-Mirkowa and Mirek, 2003; K1osowski et al., 2011).The Late Glacial development of the studied lakes is also docu-mented by the common occurrence of Chara sp. oospores. Analysisof Late Glacial sediments sampled from lakes in other parts ofPoland has also revealed the common occurrence of stonewortoospores (Kubiak-Martens, 1998; Schubert, 2003; Lamentowicz,2005; Drzymulska, 2008; Milecka et al., 2011). Numerous occur-rences of stonewort oospores in Late Glacial sediments have alsobeen recorded at other sites in Europe, including Switzerland (Haas,1999; Tobolski and Ammann, 2000), the Czech Republic (Pokornýand Jankovska, 2000), Great Britain (Walker et al., 2003), France(Guiter et al., 2005), western Russia (Wohlfarth et al., 2006),Lithuania (Stan�cikait _e et al., 2008), Estonia (Saarse et al., 2009;Amon et al., 2010), Latvia (Koff and Terasmaa, 2011), and Denmark(Mortensen et al., 2011). In Greenland, Chara sp. oospores occurredat the initial stage of the development of lakes in the Holocene(Fredskild, 1992). This finding suggests that stoneworts were amongthe first plants to colonise the bottoms of newly developed lakes. Asimilar phenomenon is also observed today. Stoneworts belong topioneer elements, which are usually the first species to colonisea lake’s bottom during its development (Dambska, 1966; Crawford,1977; Podbielkowski and Tomaszewicz, 1996). Pioneer macro-phytes that are often found in Late Glacial sediments also includeBatrachium (Birks, 2000). Within the study area, Batrachium sp.fruits were found in four lakes.

5.2. Climate cooling e Younger Dryas

The Younger Dryas and the climate cooling that occurredduring this period can be observed in the macrofossil record forsite I (Fig. 2, zone I-2). Between depths of 595 and 545 cm, theoccurrence of the birch trees B. pubescens and Betula sec. Alba, aswell as that of Pinus sylvestris, decreased significantly. A single B.nana fruit scale was recorded in this zone. During the period ofclimate cooling, Chara sp. expanded. Climate cooling during theLate Glacial is also visible at site II in zone II-3 (Fig. 3). B. nanaexpanded again in the vicinity of the lake, and S. selaginoidesappeared. The presence of S. selaginoides in sediments is typical ofcold periods, such as that of the Younger Dryas (Tobolski, 1998,2006; Stan�cikait _e et al., 2009; Gaidamavi�cius et al., 2011; Mortensenet al., 2011). Climate changes resulted in the reappearance of

P. filiformis in the lake. The correlation of the Late Dryas with thisphase in the overall pattern detected in this study was confirmed inthe palynological record, as manifested in a rapid decrease in thePinus and Betula curves (tree forms) and an increase in Juniperus (upto 15%) and Artemisa (approx. 10%), as well as B. nana (up to 10%)(Tobolski, unpublished data).

A highly valuable finding at sites IV and VII associated with theLate Glacial was S. teres leaves. S. teres currently belongs to theSphagnum species that frequently occur in eutrophic habitats(Daniels and Eddy, 1990; Hölzer, 2010; Laine et al., 2011). Thepresence of S. teres macrofossils was determined in Late Glacialsediments from southeastern Germany (Hölzer and Hölzer, 1994)and in sediments from the Late Pleistocene in northwesternGermany (Behre et al., 2005). This instead of these macrofossils’occurrence in the Late Glacial can be correlated with the presenceof a more fertile substratum, similar to the case of the presence ofstoneworts and P. filiformis. The presence of these plants in LateGlacial sediments demonstrates that during the initial stage ofthe development of the lakes, the water was rich in calcium(Samuelsson, 1934; Marciniak, 1979). This result is related to thepartial redeposition of calcium carbonate accumulated in post-glacial sediments in depressions (Nowaczyk and Tobolski, 1980;Borówka, 1992; Bukowska-Jania, 2003).

5.3. Holocene

The Holocene climate warming detected in the area basing onmulti-aspect studies performed on sediments of Lake Ha�ncza,dated to 11600 cal. BP (Lauterbach et al., 2010), is clearly observablealso in the results of the macroscopic analyses performed in thepresent study, for example, at site I (Fig. 2, zone I-3, I-4) and site II(Fig. 3, zone II-3 and II-4). The Early Holocene timing of sedimentaccumulation at those sites was confirmed by radiocarbon dating.In both cases, a date of approx. 10 745 cal. BP was obtained. At site Iin zone I-3, B. pubescens and Betula sec. Alba fruits and a N. marinaseed occurred. The latter macrophyte is commonly recognised as anindicator of a warm climate (Backman, 1941; Godwin, 1975; Lang,1994). In Denmark, seeds of N. marina the Early Holocene around10 300 cal. BP were found (Bennike et al., 2001). In the subsequentphase, I-4, Typha sp., C. pseudocyperus, and C. demersum seedsappeared for the first time at this site. The expansion of these threeplants in the area provides additional evidence that the sedimentdeveloped during the phase of climate warming at the beginning ofthe Holocene. The presence of Typha sp. macrofossils in the sedi-ment suggests that the temperature in July oscillated approxi-mately around 15 �C (Wasylikowa, 1964) or between 13 and 16 �C,depending on the Typha species (Kolstrup, 1979). C. pseudocyperusis an indicator of warm climate (Backman, 1935). Its occurrenceis correlated with a mean July temperature of approx. 13 �C(Brinkkemper et al., 1987). The presence of C. demersum can indi-cate that the temperature in July oscillated approximately around15 �C (Litt, 1994) or even 18 �C (Mai, 1985).

5.4. Early Holocene decrease in water levels

During the Early Holocene climate warming in northeasternPoland, the water level in lakes decreased significantly. Theoccurrence of a decreasing water level can be observed in a changein sediment type. Among the limnic sediments that developed inthe Early Holocene, a tens of centimeter peat thick layer occurs(Ga1ka, in preparation). Examples of decreasing lake level andsimultaneous development of peat layers on gyttjas have been re-ported from other part of Poland (Niewiarowski, 1995; Marks,1996), Eastern Belarus (Novik et al., 2010) and Scandinavia(Gaillard, 1985; Digerfeldt, 1986).

M. Gałka, M. Sznel / Quaternary International 292 (2013) 124e135 133

Additional evidence for a decrease in the water level in lakes inthe area is provided by the disappearance of smaller water bodiesand the development of peatlands on their limnic sediments. Twoof the water bodies analysed (V, VI), at the beginning of the Holo-cene changed into peatlands. At site V (Fig. 7), peatland develop-ment during zone VeIII was preceded by a decline of B. nana (zoneVeI) occurring in the Late Glacial and expansion of B. pubescens andPinus sylvestris (zone VeII). A decrease in the water level duringzone VeII is suggested by the presence of P. natans, N. alba andTypha sp., which are plants that usually grow in the zone of shallowsilting up lakes. Disappearance of the water body at site VII (Fig. 8)also occurred at the beginning of the Holocene. In zone VII-5, whichwas interpreted as a sediment layer that developed during the LateGlacial, the following macrophytes occurred: P. alpinus, P. pusillus,and Batrachium sp. Zone VII-6 is characterised by strongly decom-posed peat. A decrease in the water level is indicated by a change inthe vegetation in the lakes themselves, as manifested in Lake Kojle(site I, zone I-4), in the development of assemblages includingcontributions of P. natans and H. vulgaris, i.e., plants associated withthe shallow zones of lakes.

The decrease in water levels in the studied lakes is correlatedwith decreasing water levels in other lakes (with a surface area ofseveral hundred hectares), occurring in the Early Holocene innortheastern and eastern Poland, such as Lake qukcze (Ba1aga,1990), Lake Miko1ajki (Ralska-Jasiewiczowa and Lata1owa, 1996),and Lake Wigry (Zawisza and Szeroczy�nska, 2007). A lake leveldecrease in the area of North and Central Poland is visible in theresults of studies on fossil Cladocera at site Lake Sierzywk (Mileckaet al., 2011) and in Lake Go�scia _z (Starkel et al., 1998). The lake leveldecrease in the Early Holocene in NE Poland also corresponds withother sites in Europe, as in Estonia (Hang et al., 2008) and TheNetherlands (Bos et al., 2007).

6. Summary

The palaeoecological study presented here focused on the LateGlacial and Early Holocene history of the development of vegeta-tion in selected lakes in northeastern Poland. Within the scope ofthe study, the bottom layers of sediments from seven lakes wereanalysed in terms of their plant macrofossil content. The results ofthe study suggest the following conclusions:

1. High-resolution plant macrofossil analysis provided valuableinformation regarding climate changes, vegetation, and waterlevel variations in lakes in northeastern Poland.

2. The formation of the fossil lakes is dated to the BøllingeAllerød.According to radiocarbon dating, organic sediment accumula-tion began approx. 13 500 cal. BP (in Lake Kojle 11 650 � 7014C BP and in Lake Linówek 11 690 � 60 14C BP). The initialdevelopment of the studied lakes is demonstrated by thecommon occurrence of plant fossils that are typical of the LateGlacial. The analysed sediments included numerous B. nanafruits and fruit scales, P. filiformis and, P. alpinus endocarps, andN. pumila seeds.

3. Stoneworts (Characeae), P. filiformis and Batrachium sp. wereamong the first plants to appear in the lakes, which is inaccordance with other sites in Europe.

4. The Late Glacial sediments included rarely encountered fossilsof the moss S. teres.

5. At the beginning of the Holocene, the water level in lakes innortheastern Poland decreased. In the studied peatlands, thisdecrease is documented by- a change in the vegetation in the lakes, with species growingin shallow water beginning to appear, e.g., P. natans andH. vulgaris;

- the occurrence of a peat layer between gyttjas; and- the disappearance of smaller lakes and the development ofpeatlands on limnic sediments.

Acknowledgments

This scientific work, financed by the Ministry of Science andHigher Education as research project N N305 325933, was per-formed in the years 2007e2010 (head M. Ga1ka). We express ourgratitude to Krystyna Milecka and three anonymous reviews forproviding constructive comments on the earlier version of thispaper.

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