MAC RO FOS SIL RE CON STRUC TION OF PRE BO REAL WET LAND FORMED ON DEAD ICE BLOCK: A CASE STUDY OF THE BOR ZECHOWO MIRE IN EAST POM ERA NIA, PO LAND

Studia Quaternaria, vol. 27 (2010): 3 10. MAC RO FOS SIL RE CON STRUC TION OF PRE BO REAL WET LAND FORMED ON DEAD ICE BLOCK: A CASE STUDY OF THE BOR ZECHOWO MIRE IN EAST POM ERA NIA, PO LAND Mi cha³ S³owi ñski De part ment of Geo mor phol ogy and Hy drol ogy of Low lands, In sti tute of Ge og ra phy and Spa tial Or gani za tion, Pol ish Acad emy of Sci ences, ul. Ko per nika 19, 87-100 To ruñ, Po land, e- mail: mi chal@geo pan.to run.pl Ab stract In or der to re con struct en vi ron men tal changes in the Borzechowo mire, a sed i ment core was sub jected to macrofossil and strati graphic anal y ses. The mire is lo cated in the east ern part of the Pom er a nian Lakeland (Tuchola For est, north - ern Po land). It is a limnogenic mire, formed as a re sult of terrestrialisation of a wa ter body. The time of basal peat ac - cu mu la tion was es ti mated by ra dio car bon dat ing as 9860±130 14 C BP (Gd-12393) and by palynological anal y sis as Preboreal. The anal y sis of macrofossils shows that in that pe riod, con sid er able hy dro log i cal changes took place in the study area. These hy dro log i cal changes were caused by melt ing of dead ice blocks that was com mon place in the Late Gla cial and the Early Ho lo cene. Key words: Pre bo real, wet land, mac ro fos sils, ba sal peat, bur ied dead ice, N Po land INTRODUCTION A peat layer found at the base of limnic de pos its, over ly - ing min eral bot toms of biogenic sed i men tary bas ins, is gen - er ally as so ci ated with rapid hy dro log i cal changes at the be gin ning of lake for ma tion. In the early post-gla cial land - scape of Po land (Liberacki 1958, Kozarski 1963, Stasiak 1963, 1971, Wiêckowski 1966, 1993, urek 1990, B³aszkiewicz, Krzymiñska 1992, Nowaczyk 1994a, 1994b, 2006, Wojciechowski 2000, B³aszkiewicz 2003, 2005, 2007), Netherlands (Hoek et al. 1999) and North Amer ica (Kovanen, Easterbrook 2001), such fea tures were formed as a re - sult of melt ing of dead-ice blocks bur ied within gla cial de - pos its. De pend ing on var i ous con di tions, both re gional and lo cal, the melt ing of dead-ice blocks lasted from the be gin - ning of the Late Gla cial till the Preboreal pe riod (B³aszkiewicz 2005). Ac cord ing to the hydrologic and ge netic clas si fi ca tion by Succow (1988), mires formed on sandy-grav elly de pos its and fed by wa ter from melt ing dead-ice blocks can be de fined as paludified. Dur ing their for ma tion due to the wan ing of dead-ice blocks, the over ly ing de pos its col lapsed and the telmatic sed i ments were sub merged as a re sult of sub si dence of ket tle holes, and sub se quently cov ered with limnic de pos - its of the emerg ing lakes. Literature of the subject reports relatively large number of sites where peat de pos its de vel oped above bur ied dead-ice blocks, but in only a few such sites a macrofossil anal y sis was con ducted (Marek 1994, Wojciechowski 2000, Kowalewski et al. 2001, Wright, Stefanova 2004, Drzymulska 2006). Such an anal y sis is es pe cially im por tant when lo cal en vi ron - men tal con di tions are to be de fined in a ba sin where biogenic ac cu mu la tion takes place. In the case of fos sil sed i ments, macrofossil analysis is vital to distinguish properly between depositional en vi ron ments, i.e. limnic (a layer at the bot tom of the lake) and telmatic (start ing to ac cu mu late around the lake s per im e ter) (Eslick 2001). Since there are doubts con - cern ing proper clas si fi ca tion of basal peat, macrofossil anal - y sis seems cru cial (Marks 1996, Ga³ka 2007, Tobolski 2007). STUDY SITE The Borzechowo mire is lo cated in the east ern part of the Pom er a nian Lakeland (Pojezierze Pomorskie), at the bor der be tween the Tuchola For est (Bory Tucholskie) and the Kociewie Lakeland (Pojezierze Kociewskie), and within the catch ment of the Wda River (Fig. 1). The mire oc cu pies the bot tom of a subglacial chan nel formed on the outwash plain at the di rect forefield of the gla cier in the max i mum ex tent of the Pom er a nian phase (B³aszkiewicz 2005). The mire is an ex am ple of a de graded fen. In the 1980s, the area was drained, which in flu enced sig nif i cantly the lo cal eco sys tem. In terms of hy drol ogy and or i gin, the Borzechowo mire is limnogenic, i.e. formed by the pro cess of terrestrialisation of a wa ter body (Succow 1988). The lithofacial and chronostratigraphic char ac ter is tics of sed i ments fill ing the fos sil lake ba sin near Borzechowo were de scribed by B³aszkiewicz (2005). Ac cord ing to that au thor, the base of the biogenic de pos its lies di rectly on the min eral basal com plex. A con tin u ous layer of the Preboreal peat of var i ous thick ness (3 10 cm), was found at the depth of 11.42 m. The above layer un der lies a thick stra tum of cal - car e ous gyttja, which in the top sec tion is cov ered by peat.

4 M. S OWIÑSKI MATERIAL AND METHODS The sed i ment core was ob tained with the use of 110 cm-long (5 cm di am e ter) Living stone corer in Wiêckowski s mod i fi ca tion (Wiêckowski 1956). The sed i ment for macrofos sil anal y sis was col lected at the cen tral sec tion of the subglacial chan nel, at the con tact zone with the Wda River val ley (Fig. 1). In the cen tral sec tion of the chan nel, for the pur pose of this study, we took the com plete pro file of biogenic sed i ments, whose to tal length was 11.42 m. Li thol ogy of the sed i ments was de scribed fol low ing West (1977). Prep a ra tion of sam ples for macrofossil anal y sis fol - lowed widely ac cepted meth ods (Tobolski 2000). Be fore the anal y sis the cores were frag mented into 1-cm sec tions. The bot tom de pos its, highly de com posed, were boiled in the so lu - tion of po tas sium hy drox ide (KOH) for about 10 min utes. Next, the sludge was sieved through mesh sizes of 0.1 mm, 0.25 mm and 0.5 mm. Each sam ple was used to pre pare four mi cro scope slides. De ter mi na tion of both fos sil and subfossil re mains was based on the lit er a ture (Bertsch 1941, Landwehr 1966, Katz et al. 1977, Grosse-Brauckmann 1972, Nilsson 1972, Beijerinck 1976, Daniels, Eddy 1985, Smith 2004, Tobolski 2000, Ga³ka 2006, Velichkevich, Zastawniak 2006) and the ref er ence col lec tion of the De part ment of Biogeography and Paleoecology. The re sults ob tained from the macrofossil study were ana lysed and pre sented in the form of macrofossil di a grams plot ted by the C2 soft ware (Jug gins 2003). The un count able macrofossils are pre sented in the di a gram on a five-level scale. Count able items, such as seeds and nee dles, are given in ab so lute val ues. The lo cal macrofossil as sem blage zones (LMAZ) were de lim ited. The dig i tal ter rain model for the study area has been gen - er ated through the near est neigh bour al go rithm from points ob tained as a re sult of vectorisation of con tour lines from a top o graphic map on a scale of 1:10 000 (sheet num ber and name: 335.113 Borzechowo). The ESRI ArcMap9 soft ware was used. Fig. 1. Lo ca tion of the study site and the geo log i cal cross-sec tion the Borzechowo mire. 1 dig i tal ter rain model (DTM) with lo ca tion of a core as well as the course of the geo log i cal sec tion; 2 li thol ogy of the pro file; 3 geo log i cal sec tion A-B across the Borzechowo mire (af ter B³aszkiewicz 2005).

MACROFOSSIL RE CON STRUC TION OF PREBOREAL WETLAND 5 RE SULTS Re sults of the macrofossil anal y sis are shown in Fig. 2 and Ta ble 1. The di a gram in Fig. 2 in cludes six cat e go ries (brown mosses and Sphag num mosses, trees and shrubs, aquatic plants, telmatic vegetation, invertebrates and others). The anal y sis of biogenic de pos its in di cates that a 14-cm basal peat, sep a rated by a layer of ae olian sands (4.5 cm) is un der - lain by sands of var i ous grain sizes (depth 11.42 m). The peat layer is over lain by a grey ish-cr me calcareous gyttja. The be gin ning of basal peat ac cu mu la tion was dated on 9860± 130 years BP (Gd-12393). On the ba sis of the macrofossil anal y sis of the core sec tion, two stages of de vel op ment of the study ob ject can be dis tin guished: telmatic and limnic. Telmatic phase Telmatophytes ini ti ated the de vel op ment of the mire. They in cluded Thelypteris palustris and Equisetum sp. Ac - cord ing to the eco log i cal in di ca tor val ues (Zarzycki et al. 2002) those spe cies pre fer me dium shad ing and fer tile (eutrophic) en vi ron ments with neu tral re ac tion (6 ph<7) and form dense patches (K³osowski S., K³osowski K. 2001). They were accompanied by Phragmites aus tra lis and Carex spp. The ex ist ing ba sin with biogenic sed i ments was then over grown by shrubs of Betula spp. This point of view is sup - ported by the subfossil find ings linked with this spe cies, such as fruit and periderm. Betula spp. was ac com pa nied by the dwarf shrub of Betula nana, whose fruit was found in the bot - tom de pos its. In the Preboreal pe riod this spe cies was abun - dant in the veg e ta tion of the Pom er a nian re gion (Lata³owa 2003). The other spe cies pres ent in the area in cluded trees: Populus tremula and Pinus sylvestris. Dur ing this phase, fires oc curred in the mire area. This is in di cated by nu mer ous sharp-edged pieces of char coal of up to 17 mm in di am e ter. Two peaks of char coal con tent were re corded: the first one at the depth from 11.38 m to 11.35 m, and the other one at 11.31 m. The fires on that mire, prob a bly in clud ing the neigh - bour ing ar eas, burnt the veg e ta tion. This ini ti ated short-last - ing ae olian and de nu da tion pro cesses, lead ing to the accumulation of a min eral layer. The larg est stock of the sandy frac - tion in the ana lysed sec tion of the pro file is re corded at the depth of 11.31 m. Due to the fire, the ph value in the sur face layer in creased, while the con tent of the to tal ni trate de - creased. This brought about a change in both the soil and veg - e ta tion cover. The wet land sur face low ered down to the wa ter ta ble, re sult ing in sec ond ary plant suc ces sion fol lowed by the paludification pro cess (Kania et al. 2006). The fire caused edaphic changes within the mire sub stra - tum, which al tered the veg e ta tion. New spe cies ap peared, such as Lycopus europaeus, Filipendula ulmaria, Calamagrostis sp., Scirpus lacustris, Eriophorum angustifolium and Epipactis sp. More over, the den sity of emer gent veg e ta tion in creased it in cluded Thelypteris sp. and Carex sp. It must be stressed that the larg est stock of seeds of Lycopus europaeus is cor re lated with the wan ing of char coal con tent. This is a re sult of the fast ad ap ta tion of this spe cies a pi o neer of secondary succession (Kania et al. 2006). Lycopus europaeus tol er ates shad ing in the eutrophic en vi ron ment with neu - tral reaction (6 ph<7) (Zarzycki et al. 2002). It is a dif fer en - tial spe cies of the phytosociological class Phragmitetea (Podbielkowski, Tomaszewicz 1982). Filipendula ulmaria, whose seeds were found at a sim i lar depth, also in di cates such edaphic con di tions. Mosses are im por tant bioindicators of the nat u ral en vi - ron ment as well as com po nents of de pos its (Dick son 1986, Tobolski 2006, Janssens 1990). Brown mosses are ex cel lent in di ca tors of minerotrophic hab i tats. Meesia triquetra is the dom i nant brown moss re corded in the ana lysed de posit. This spe cies is treated as a relic of the rich con ti nen tal fens (Jannsens 1990, Lamentowicz 2005, Swinehart 1995), in some pub li ca tions also de scribed as a gla cial relic (Tobolski 2003, Ga³ka 2007). They played an im por tant role in the peat-form ing pro cesses at the late Pleis to cene and early Ho - lo cene (Jasnowski 1957a, 1957b, 1959). In this phase, peat mosses were less abun dant than brown mosses. Testae amoebae are sensitive indicators instantly react - ing to changes in the nat u ral en vi ron ment, such as eutrophicat ion, acid i fi ca tion, drain age or waterlogging (Tolonen 1986, Charman 2002, Lamentowicz 2006, 2007a, 2007b). The appearance of Centropyxis aculeata indicates the change of wa ter con di tions in the mire. It im plies the chang - ing hy dro log i cal con di tions in the ba sin of biogenic ac cu mu - la tion, which is proved by the pres ence of the amoeba Arcella discoides (Lamentowicz 2005, 2008). This spe cies pre fers wet hab i tats (Warner 1990a), but it tol er ates a wide range of ph (Lamentowicz 2005). Nearly the en tire sec tion of the core un der ques tion con tains head cap sules of Chironomidae as well the re mains of Cladocera and Oribatida. At the end of Table 1 Char ac ter is tics of the lo cal macrofossil as sem blage zones (LMAZ) from the bot tom de pos its of the Borzechowo mire Zone Depth (m) Characteristics BorzM1 Carex-Lycopus BorzM2 Nymphaea- Ceratophyllum 11.42 11.24 11.24 11.18 This zone in cludes mire subfossil plant re mains: Carex sp., Phragmites aus tra lis, Thelypteris sp., Equisetum sp., Calamagrostis sp., Filipendula ulmaria, Lycopus europaeus, Ranuculus sp., Eriophorum angustifolium, Pinus sylvestris, Betula sp. and Calluna vulgaris. Other find ings in clude moss (Sphag num spp., Polytrichum com - mune, Messia triquetra, Drepanocladus) and re mains of Pinus sylvestris, Populus tremula and fruits of Betula nana. This zone also in cludes wood, char coal and sand. In the fi nal sec tion of this zone, wa ter plant re mains ap - pear (Ceratophyllum demersum and Nymphaea sp). A char ac ter is tic fea ture of this zone is the pres ence of aquatic plants, such as Ceratophyllum demersum, Nymphaea sp., Najas ma rina and Chara sp. Large amounts of Mollusca ap pear, whose abun dance de creases when mov ing up. Aquatic fauna: Arcella discoides, Chironomidae and Cladocera.

6 M. S OWIÑSKI F ig. 2. Plant macrofossil diagram of the basal peat of the Borzechowskie mire.

MACROFOSSIL RE CON STRUC TION OF PREBOREAL WETLAND 7 the telmatic phase, Thelypteris palustris and Scirpus lacustris in creased in abun dance. Their max i mum is re corded at the depth of 11.26 m. Other plant re mains that be came more abun dant at that time in clude utricles of Carex sp. (rostrata?), nee dles of Pinus sylvestris, fruits of Betula sp. and epi der mis of Calamagrostis sp. Limnic phase The be gin ning of this phase in di cates a rad i cal trans for - ma tion of the con di tions of biogenic ac cu mu la tion. The semi-aquatic hab i tat evolves into an aquatic one. The palynolog i cal anal y sis made by B. Noryœkiewicz (un pub lished data) in di cates that those changes took place at the turn from the Preboreal to the Bo real pe ri ods. The ex ist ing then wa ter body was sur rounded by pine-birch for ests with a quickly spread ing ha zel. The limnic phase doc u ments the first stage of lake de vel - op ment. Be sides the change of the de posit from a peaty to limnic one (cal car e ous gyttja), also other changes were re - corded, e.g. in the subfossil plant re mains and the ap pear ance of Mollusca. Ceratophyllum demersum was found at the depth 11.27 m. It is a free-float ing aquatic plant, which pre - fers fer tile or very fer tile small astatic wa ter bod ies of neu tral or even alkaline reaction (6 ph>7). Ceratophyllum demersum reaches its max i mum abun dance in shal low wa ter bod ies heated in ten sively in sum mer. En vi ron men tal tol er ance en - ables this spe cies to grow at the depth from 0.5 to 10 metres (Hannon, Gaillard 1997). It cre ates dense patches (K³osowski S., K³osowski K. 2001, Zarzycki et al. 2002). The larg est amount of Mollusca is found in the third centi metre of the core in the limnic phase, at the depth of 11.24 m. The lake was oc cu pied by spe cies of the ge nus Nymphaea with free-float ing leaves. This is proved by its re mains, such as the epi der mis and idioblasts. Plants of the ge nus Nymphaea pre - fer meso-eutrophic wa ter bod ies, whose re ac tion ranges from slightly acidic to al ka line (5 ph>7) (Zarzycki et al. 2002). They also tol er ate a small depth of wa ter, and rarely live be - low 3 metres in depth (Hannon, Gaillard 1997). Due to their fast growth and large pro duc tion of plant bio mass, those plants con trib ute to shallowing and over grow ing of wa ter bod ies (Janecki 1999). Nymphaeids were ac com pa nied by Ceratophyllum demersum, whose nee dle-like leaves were found at the top of the ana lysed sec tion. The pres ence of oo - spores in di cates that Charales also grew there. Lake con di - tions are also con firmed by the pres ence of head cap sules of Chironomidae and the de creas ing abun dance of Oribatida. In this en vi ron ment, large com mu ni ties of Cladocera de vel - oped. The lake shores were over grown by sedge com mu ni - ties with brown mosses of the ge nus Drepanocladus, as well as Thelypteris palustris, Phragmites aus tra lis and Scirpus lacustris. The top sec tion of the core also in cluded the seeds of Najas ma rina. This macrophyte is an an nual plant and forms thick un der wa ter mead ows. Najas ma rina pre fers shal low meso-eutrophic lakes up to 3 m deep (op ti mum depth is about 1 m) (Hannon, Gaillard 1997), and al ka line re - ac tion of ph>7 (Zarzycki et al. 2002). Such con di tions are found in both fresh wa ter and salty wa ter bod ies (sa lin ity up to 10 ). Najas ma rina was also re corded in de pos its of the Bal tic Sea (Bennike et al. 2001). Plant in di ca tors found in the ana lysed sec tion of the core, such as Ceratophyllum demersum, Lycopus europaeus and Scirpus lacustris, indicate that mean minimum temperature for July at the base of biogenic ac cu mu la tion ranged be tween 13 C and 16 C (Isarin, Bohncke 1999, Bos et al. 2007). The en tire sec tion of the core in cluded an ad mix ture of un iden ti - fied or ganic mat ter, which is plot ted in the di a gram as UOM. DISCUSSION High res o lu tion anal y sis of the bot tom sec tion of the core en abled re con struc tion of the evo lu tion of the mire, which had de vel oped on a bur ied block of dead ice. Ac cord ing to the macrofossil anal y sis for the bot tom sec tion of the core, the de posit found di rectly on the min eral base is thought to be fen. It was ac cu mu lated in telmatic con di tions, which are ev i - denced by subfossil plant re mains. On the ba sis of the pres - ence and rel a tive abun dance of spe cific plant spe cies, the so-called in di ca tor plants, it was pos si ble to in fer the type of fen. It was Limno-Phragmitioni (reed peat) (Tobolski 2000, To³pa et al. 1967). In in ves ti ga tions on palaeohydrological changes of wetlands and cor re la tions be tween that peat and en vi ron men tal fac tors, urek (1990, 1993) proved that peat was de vel op ing on sandy de pos its in the Late Gla cial pe riod. Ac cord ing to him, this phe nom e non is linked with melt ing out of dead ice blocks and de vel op ment of peat de pos its on top of it. In creas ing rel a tive wa ter ta ble fi nally sub merged the peat. Sum ma riz ing all the re search, it may be con cluded that the basal peat came into be ing as a re sult of in ten si fied paludification dur ing an early phase of the dead ice melt-out. Wa ter logged basal sed i men tary com plex, found on top of the melt ing block, con trib uted to the de vel op ment of minerotrophic peat-form ing veg e ta tion. Ground wa ter played a cru cial role in the de vel op ment of the mire. It stim u lated sed i men ta - tion of biogenic mat ter di rectly on sandy-gravely de pos its. The rate of peat ac cu mu la tion de pends mostly on cli ma tic and hy dro log i cal con di tions in the place where biogenic ac - cu mu la tion oc curs. Those con di tions in flu ence the biogenic de com po si tion and ac cu mu la tion. Plant com mu ni ties that take part in peat sed i men ta tion are also of great sig nif i cance. How ever, be sides cli ma tic and hy dro log i cal con di tions they are also de pend ent on edaphic con di tions. The macrofossil anal y sis, es pe cially of the subfossil artefacts re corded at the bot tom of the Borzechowo mire, made it pos si ble to re con - struct palaeohydrological con di tions in this lo cal ity. The basal de pos its that fill up the bed of the subglacial chan nel in - di cate two phases of de vel op ment: telmatic and limnic. Both plant and an i mal spe cies found in the de pos its in di cate fre - quent os cil la tions of the wa ter ta ble. The telmatic phase in di - cates a ris ing wa ter ta ble. At that time a shal low astatic wa ter body ex isted. This was the ef fect of sys tem atic melt ing out of dead ice blocks, which fi nally caused the deep en ing of the ba sin and in tro duc tion of new el e ments in limnic flora. A sig - nif i cant wa ter rise, which brought about a change from a telmatic to a limnic en vi ron ment, is in di cated by the pres ence of plant macrofossils of Najas ma rina, Nymphaea sp. (Hannon, Gaillard 1997), al gae Chara sp. (Podbielkowski 1978, Pe³echaty et al. 2007) and an i mal macrofossils of Tardigrada (Cro mer 2008), Cladocera and Chironomidae.

8 M. S OWIÑSKI The basal peat was rarely ana lysed in re spect of palaeo - ec ol ogy and macrofossils. A few pa pers dis cuss the floristic com po si tion and evo lu tion of wet land de vel op ment on bur - ied blocks of dead ice (Marek 1994, Wojciechowski 2000, Kowalewski et al. 2001, Wright, Stefanova 2004, Drzymulska 2006). Both floristic and faunistic com po si tion of the bot tom sec tion of the cores, which in cluded basal peat, in di - cate palaeohydrological changes in the sed i men tary bas ins. The thresh old was the first win ter sea son when the wa ter above the peat bog sur face did not freeze to the bot tom (B³aszkiewicz 2005). This trig gered a sud den dis in te gra tion of dead ice, and a quick sub merg ing of the mire. The macrofos sil com po si tion in di cates quick changes in the wa ter ta ble (Hannon, Gaillard 1997). The or i gin of the lake ba sin of the Borzechowo mire cor re lates with the re corded palaeohydrolog i cal changes in the early post-gla cial land scape ar eas in Cen tral Eu rope. On the ba sis of the re search by Niewiarowski (1990) and Ralska-Jasiewiczowa (1987), urek (1990) de lim ited two pe ri ods of low wa ter ta ble. The first one was con nected with the fi nal stage of the dead ice melt-out, correlated with the palaeoecological investigations in the bot tom de pos its of the Borzechowo mire. The macrofossil anal y sis is a key el e ment in an un am big - u ous def i ni tion of biogenic sed i ments and of the char ac ter of peat de pos its (Tobolski 2000, 2006). Along with other palaeoecological analyses (analyses of pollen, testate amoe - bae, cladocerans, di a toms, chi rono mids, etc.) it re flects a wider spec trum of en vi ron men tal changes and en hances pos - sibilities for their more accurate interpretation, as well as the processes that stimulated them. Acknowledgements I thank Ass. Pro fes sor Miros³aw B³aszkiewicz and Pro fes sor Tomasz Goslar, Dr Jaros³aw Kordowski for sup port ing our work and care ful re vi sion of the manu script. I also would like to thank re - view ers for their valu able re marks on the manu script, Dr Bo ena Noryœkiewicz for shar ing her un pub lished data, and Sebastian Tyszkowski, MSc, for co op er a tion in the field work. This work is part of a re search pro ject funded by the Min is try of Sci ence and Higher Ed u ca tion: Cli mate and en vi ron ment change in the Late Gla cial and early Ho lo cene in the area Tuchola For est in the light of high-res o lu tion palaeo eco logi cal anal y sis (no. N N306 085037) (Prin ci pal In ves ti ga tor Michal S³owiñski). REFERENCES Bennike O., Jensen J.B., Lemke W. 2001. Late Qua ter nary re cords of Najas spp. (Najadaceae) from the south west ern Bal tic re - gion. Re view of Palaeobotany and Palynology 114, 259 267. Beijerinck W. 1976. 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