NON-POL LEN PALYNOMORPHS CHAR AC TER IS TIC FOR THE DYSTROPHIC STAGE OF HUMIC LAKES IN THE WIGRY NA TIONAL PARK, NE PO LAND

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1 Studia Quaternaria, vol. 32, no. 1 (2015): DOI: /squa NON-POL LEN PALYNOMORPHS CHAR AC TER IS TIC FOR THE DYSTROPHIC STAGE OF HUMIC LAKES IN THE WIGRY NA TIONAL PARK, NE PO LAND Magdalena Fi³oc, Miros³awa Kupryjanowicz De part ment of Bot any, In sti tute of Bi ol ogy, Uni ver sity of Bia³ystok, K. Cio³kowskiego 1J, Bia³ystok, Po land; mfiloc@op.pl, m.kupryjanowicz@uwb.edu.pl Ab stract The nu mer ous dystrophic (humic) lakes are a very im por tant fea ture of Wigry Na tional Park, NE Po land. As the most re cent palaeo eco logi cal data in di cate, at the be gin ning of its de vel op ment (in the Late Gla cial and Early and Mid dle Ho lo cene) these wa ter bod ies func tioned as har mo ni ous lakes, and their trans for ma tion into dystrophic lakes and the sta bi li za tion of the trophic state took place at the be gin ning of the Subboreal. Palynological anal y sis of sed i ments from two such lakes (Lake Œlepe and Lake Suchar II), with spe cial em pha sis on non-pol len palynomorphs (NPPs), was aimed at a de tailed bi o log i cal char ac ter iza tion of dystrophic lakes dur ing their long-last ing ex is tence. The ob tained re - sults al lowed for the des ig na tion of or gan isms char ac ter istic for dystrophic lakes, of which representatives appeared with the de creas ing ph of the wa ter and the for ma tion of Sphagnum peat around lakes. These or gan isms were di vided into four groups: al gae, fungi, tes tate amoe bas, and an i mals. Their representatives appear in various developmental stages of dystrophic lakes. Key words: NPPs, palaeo eco logi cal re con struc tion, humic lake, pol len anal y sis, NE Po land Manuscript received 30 September 2014, accepted 11 May 2015 INTRODUCTION Dystrophic lakes are an im por tant com po nent of land - scapes in nu mer ous parts of the world, al though op ti mum con di tions for form ing these lakes are most abun dant in the north ern re gions with a cool, hu mid cli mate (Salonen et al., 1983; Kankaala et al., 2006). The nu mer ous dystrophic lakes are a unique fea ture of the Wigry Na tional Park, but in other re gions of Po land they are rel a tively rare (Kraska et al., 2001; Wilk-WoŸniak et al., 2012). In the Wigry Na tional Park area there are twenty-one dystrophic lakes. They are small, shal low wa ter bod ies with - out out flow. The wa ter of this type of lakes is char ac ter ized by spe cific en vi ron men tal con di tions, such as a def i cit in nu - tri ents, higher amount of humic sub stances (it gives wa ter a brown color), low ph, low level of ox y gen and low con cen - tra tion of cal cium in the wa ter and sed i ments (Kamiñski at al. 2001; Górniak 1995). Humic ac ids in the wa ter re act with many microelements and chem i cal com pounds nec es sary of life for plants and an i mals, ty ing them up and as a re sult make them un avail able in the wa ter. All these fac tors cause the poor biodiversity and small phytoplankton bio mass (Górniak et al., 1999; G¹bka and Owsianny 2006). Typ i cal for dystrophic lakes is the oc cur rence of the float ing mats with mire plants such as Menyanthes trifoliata, Ledum palustre, Eriophorum vaginatum, Andromeda polifolia, Drosera rotundifolia, Scheuchzeria palustris, Carex limosa, Carex rostrata, Sphag num angustifolium, Sphagnum fallax and Sphagnum magellanicum. The other characteristic fea ture of dystrophic lakes is that their catch ment ar eas are cov ered by co nif er ous for ests with a large pro por tion of spruce (Kamiñski 2002). In ves ti ga tions of life in dystrophic lakes has been lim ited mainly to con tem po rary flora and fauna (Czeczuga 1995, Zawiska et al., 2013). Also the de gree of knowl edge on the past changes in the trophic sta tus of this type lakes is not sat - isfactory and seem a very interesting scientific problem. Un - til re cently, dystrophic lakes were de scribed as not chang ing dur ing their de vel op ment (Górniak 1996). The lat est stud ies in di cate that a tran si tion from a typ i cal humic state to an other state and in versely to that pro cess proves to have been likely in the past as ap pears from geo chem i cal stud ies and anal y sis of macrofossil plant re mains (Drzymulska and Zieliñski 2013; Drzymulska et al., 2013). The next step is to com ple - ment of these stud ies by re search of postglacial suc ces sion of veg e ta tion and other or gan isms ex ist ing in these lakes. Ac cord ingly, in ter dis ci plin ary palaeo eco logi cal re - search of sev eral dystrophic lakes lo cated within the Wigry Na tional Park, which has been car ried out for a few years in the De part ment of Bot any at the Uni ver sity of Bia³ystok, has an im por tant role in stud ies of this type of lakes (Drzymulska 2012; Drzymulska and Kupryjanowicz 2012; Fi³oc and Ku-

2 32 M. FI OC & M. KUPRYJANOWICZ max. depth, 54 05'14" N, 23 01'03" E) are lo cated in north-east ern Po land, in Wigry Na tional Park (WNP), near the shore of Lake Wigry, the larg est lake in the na tional park (Fig. 1). The sur face of this area was shaped by the Vistula Gla ci ation, i.e. Weischelian (Marks 2002). WNP lies on the bor der two phys i cal-geo graph ical mesoregions, the East Suwa³ki Lakeland and the Augustów Up land, which are in - cluded to the Lith u a nian Lakeland (Kondracki 1994). The cli mate of this area is tem per ate tran si tional with a ten dency to ward con ti nen tal. It is most se vere across the low land parts of the coun try (Krzysztofiak and Olszewski 1999). The in ves ti gated lakes are char ac ter ized by the zon ing of veg e ta tion with mire plants typ i cal for humic lakes along the sublittoral to lit to ral. The last veg e ta tion zone con sists of marshy co nif er ous for est with Pinus sylvestris, Picea abies and Betula pubescens grow ing on a peat sub stra tum on the mar gin of the lakes. Fig. 1. Core Lo ca tion of stud ied lakes. * places of the corings. Table 1 Ra dio car bon dat ing of the an a lyzed sed i ments Depth [m] Dated material Age 14 C (BP) pryjanowicz 2013a, b; Drzymulska et al., 2014, 2015). The study in cludes a lot of as pects of the his tory of the ex am ined lakes in clud ing changes in trophic state. A part of the pro ject is a pol len anal y sis of sed i ments of Lake Œlepe and Lake Suchar II. Its main pur pose is to iden tify or gan isms lim ited to dystrophic lakes. Par tic u lar em pha sis is placed on the iden ti - fi ca tion of non-pol len palynomorphs, which al lows for a de - tailed re con struc tion of the changes tak ing place in these wa ter bod ies dur ing dystrophic state. STUDY AREA Cal i brated age (cal. years BP) range 68.2% range 95.4% S stems of 9560± SII mosses 3080± SII ± SII sediment 4120± SII ± S 1 pro file from shore of Lake Œlepe, S 2 pro file from cen tral part of Lake Œlepe, SII 1 pro file from shore of Lake Suchar II, SII 2 pro file from cen tral part of Lake Suchar II. The humic Lake Œlepe (0.6 ha, 5.5 m max. depth, 54 00'35" N, 23 06'46" E) and Lake Suchar II (2.6 ha, 9.5 m Field works METHODS Four sed i ment cores were col lected from the lakes se - lected for study. The cores from mar gin parts of lakes (S 1 and SII 1 ) were col lected with a Rus sian corer (Belokopytov and Beresnevich 1955; Jowsey 1965; Aaby and Digerfeldt 1986) with a length of 50 cm and a di am e ter of 8 cm. The length of the cores was 4.38 m for Lake Œlepe and 7.00 m for Lake Suchar II. The drillings in cen tral parts of lakes (S 2 and SII 2 ) were car ried out us ing the Wiêckowski s probe (Wiêckowski 1989) with a length of 110 cm and a di am e ter of 5 cm. The length of the cores was 5.18 m for Lake Œlepe and 6.10 m for Lake Suchar II. It was nec es sary to sup ple ment the col lected pro files with top lay ers of highly liq ue fied sed i ments that could not be col lected with a Wiêckowski s probe. The miss - ing sed i ments from cen tral part Lake Œlepe 0.23 m were col lected us ing the Kajak probe. The sed i ments from cen tral part Lake Suchar II were not col lected yet. Age of sed i ments The age of sed i ments from Lake Suchar Wielki was de - ter mined by AMS ra dio car bon method in the Gliwice Ra dio - car bon Lab o ra tory, Poznañ Lab o ra tory and Ab so lute Dat ing Lab o ra tory in Ska³a (Tab. 1). OxCal on line soft ware (Bronk Ramsey 2009) was used to cal i brate the ra dio car bon age of the sam ples. Due to a small num ber of ra dio car bon age de ter mi na - tions in the stud ied pro files, the chro nol ogy of events re - corded in these pro files has been de ter mined also in di rectly, based on a sim i lar ity be tween pol len spec tra with the ra dio - met ri cally well-dated pro file from the nearby Lake Wigry (Kupryjanowicz 2007) and it was pub lished (Drzymulska et al., 2014, Fi³oc et al., 2014). The lo cal pol len as sem blage zones from the mar ginal and cen tral parts of lakes were cor - re lated in the ear lier pa per (Drzymulska et al., 2014). Pol len anal y sis Sam ples for pol len anal y sis, 1 cm 3 in size were taken from cores ev ery 2 cm. The prep a ra tion of the sam ples and

3 NON-POL LEN PALYNOMORPHS CHARACTERISTIC 33 their mi cro scopic anal y sis were car ried out in ac cor dance with the stan dard pro ce dure (Berglund and Ralska-Jasiewiczowa 1986). Each sam ple was boiled with 5% KOH, and next treated by Erdtman s acetolysis method (Faegri and Iversen 1975). The ma te rial was then mounted in glyc er ine. In each sam ple, at least 500 pol len grains of trees and shrubs (AP) and ter res trial herbs (NAP) were counted as well as all other ac com pa ny ing palynomorphs. Pol len and spores were iden ti fied us ing sev eral keys (e.g. Moore et al., 1991; Beug 2004) and the ref er ence col lec tion of mod ern pol len slides from the De part ment of Bot any, Uni ver sity of Bia³ystok. The non-pol len palynomorphs (NPPs) were ana lysed us - ing sev eral keys (e.g. Van Geel 1978; Van Geel et al., 1981; Komárek and Jankovská 2001; The per cent age value of each pol len and non-pol len taxon has been cal cu lated in re la tion to the to tal sum of tree and shrub (AP), and her ba ceous plant pol len (NAP). The re sults are pre sented as per cent age pol len di a grams pre pared with POLPAL 2004 ver soft ware (Walanus and Nalepka 1999; Nalepka and Walanus 2003). The di a grams were di vided into spe cial pol len as sem blage zones (S PAZ) (Figs 2, 3) il lus trat ing the changes in the lo cal plant or other or gan ism com mu ni ties with the use of CONISS ap pli ca tion (Grimm 1987). RESULTS The an a lyzed cores had been shortly de scribed dur ing the field works, and then com pleted af ter clean ing them in the lab o ra tory (Tab. 2). In the sim pli fied pol len di a grams 2 spe cial pol len as sem - blage zones based on changes in wa ter and mire taxa (in clud - ing NPPs) were dis tin guished for mar ginal part Lake Œlepe Lake (S 1 pro file) and 4 for cen tral part Lake Œlepe (S 2 pro file) (Fig. 2) and 4 for mar ginal part Lake Suchar II (SII 1 pro file) and 4 for cen tral part Lake Suchar II (SII 2 pro file) (Fig. 3). Their short char ac ter is tics are showed in Ta ble 3. Non-pol len palynomorphs (Fig. 4) had a very im por tant role in de ter min ing the spe cial pol len as sem blage zones and subzones. These or gan isms were grouped into four ma jor tax o nomic groups (al gae, fungi, tes tate amoeba, an i mals), which made it pos si ble to trace the changes in trophic sta tus of in ves ti gated lakes and de vel op ment of Sphag num peat. CHANGES IN THE TROPHIC STATE OF STUDIED LAKES INTERPRETATION AND DIS CUS SION OF RE SULTS Depth [m] Lake Œlepe mar ginal part (S 1 ) Li thol ogy of an a lyzed pro files Sphag num peat Sediment description lack of sed i ment; there is wa ter lens sapropel Lake Œlepe cen tral part (S 2 ) sapropel Lake Suchar II mar ginal part (SII 1 ) Sphag num peat lack of sed i ment; there is wa ter lens Sphag num peat lack of sed i ment; there is wa ter lens Sphag num peat Sphag num peat with Pinus sapropel Lake Suchar II cen tral part (SII 2 ) sapropel Table 2 In pro files from cen tral parts of lakes, depths are counted from the wa ter sur face Drzymulska et al. (2015) proved that the trans for ma tion to the dystrophic state in Lake Suchar II took place at the be - gin ning of the Subboreal. This trans for ma tion was made pos - si ble by the changes in the en vi ron ment that oc curred at this time, e.g. cli mate change to colder and drier. A de crease in av er age tem per a tures and an in crease in the amount of pre - cip i ta tion led to the for ma tion of pine for ests with a large share of spruce in this re gion. As we know, the co nif er ous for ests grow ing in the catch ment area are a source of large quan ti ties of humic sub stances (HS) flow ing into the lake (Hagedorn et al., 2000; Górniak and Zieliñski 2000). Their sig nif i cant sup ply was just one rea son for the sub stan tial shift in the trophic state of the lake stud ied, i.e. the shift to humotro phy (Drzymulska et al., 2015). Based on the re sults for Lake Suchar II, we con cluded that in Lake Œlepe the dystrophic state was also reached in the Subboreal. How ever, as our data show, in the older and mid dle part of the Subat lan tic (S 2 -III S PAZs Fig. 2) this lake tem po rarily re turned to mesotrophy, and then it was fi nally trans formed to the humotrophic state in the youn ger part of the Subatlantic (S 1 -I, S 1 -II, S 2 -IV S PAZs Fig. 2). Tran si tion into the state of dys - tro phy and the deep en ing of this state were ac com pa nied by changes in var i ous non-pol len palynomorphs in stud ied sed i - ments. Dur ing the dys tro phy stage of both stud ied lakes, al gae (mainly Botryococcus) were the group of or gan isms with the highest percentage share. Botryococcus is a ge nus that in - cludes many spe cies of al gae char ac ter is tic for small dystrophic, oligotrophic or mezotrophic lakes, and some spe cies are char ac ter is tic for eutrophic lakes and bogs (Jankovská and Komárek 2000). This taxon showed a high share, even in the mar ginal parts of the in ves ti gated lakes, where float ing mats lim ited ac cess to light. An other char ac ter is tic group of or gan isms for the dystrophic stage of the stud ied lakes is Pediastrum angulosum var. angulosum (Fig. 4A). This cos - mo pol i tan alkaliphilous taxon is as so ci ated with eutrophic wa ter bod ies and metaphyton of the shal low and lit to ral parts of var i ous types of lakes (Jankovská and Komárek 2000; Komárek and Jankovská 2001). Our re search sug gests that Pediastrum angulosum var. angulosum is the best adapted Pediastrum to low trophic con di tions. This is con firmed by research on contemporary subarctic lakes in Finland (Weckström et al., 2009), which are also sur rounded by spruce-pine for ests. Sim i larly, the study car ried out on pres ent-day lakes in Po land in di cate that this taxon oc curs in low land lakes with lower trophic sta tus (Lenarczyk 2014). Fungi such as Tillietia sphagni, Entoplyctis lobata, Helicoon pluriseptatum (Fig. 4B) and the other un iden ti fied taxa, characteristic for mire communities, are the next discussed group of or gan isms. Tillietia sphagni (Fig. 4C) is a par a site

4 34 M. FI OC & M. KUPRYJANOWICZ Table 3 Char ac ter is tics of the spe cial pol len as sem blage zones (S PAZs) il lus trat ing changes in the lo cal plants or other or gan ism dis tin guished in an a lyzed pro files Num ber Depth [m] Description Lake Œlepe mar ginal part (S 1 ) S 1 -I S 1 -II Lake Œlepe cen tral part (S 2 ) S 2 -I Max i mum of Cyperaceae (4.7%); con tin u ous curve of Equisetum, Entoplyctis lobata, Assulina muscorum/seminulum, Centropyxis, Hyalosphenia papilio, Helicoon pluriseptatum, Tillietia sphagni and fun gal re mains; high val ues of Sphag num (17.5%); fall of Botryococcus (5 35.5%). Maxima of Sphag num (32.5%), Entoplyctis lobata (10%), Helicoon pluriseptatum (4.8%), Tillietia sphagni (2.6%), Assulina muscorum/seminulum (1.7%) and fun gal re mains (31.5%). Pro por tion of Botryococcus be tween 2 and 20.5%; val ues of Filicales monolete and Turbellaria very low (to 0.6% and 1%, respectively). S 2 -II Rel a tively low val ues of Botryococcus with peak of 38%. S 2 -III Max i mum of Botryococcus (6 69%). S 2 -IV Increase of Sphag num to 4.2%; high val ues of Cyperaceae with max i mum 1.3%; fall of Botryococcus (3 46.5%). Lake Suchar II mar ginal part (SII 1 ) SII 1 -I SII 1 -II SII 1 -III SII 1 -IV Max i mum of Botryococcus (20%); be gin ning of con tin u ous curve of Cyperaceae; cul mi na tion of Pediastrum angulosum var. angulosum with max i mum 0.5%; low val ues of Amphitrema flavum (0.5%). Decrease of Botryococcus (3 0.6%); con tin u ous curve of Filicales monolete ( %); max i mum of Scheuchzueria palustris (2.7%); low val ues of Sphag num ( %). Max ima of Cyperaceae (5.5%), Amphitrema flavum (43%) and Assulina muscorum/seminulum (8.0%); rises of Sphag - num to 14%. Maxima of Sphag num (25.5%), Tillietia sphagni (12%), Entoplyctis lobata (4%) and Hyalosphenia subflava (6.5%). The zone was di vided into tree subzones: SII 1 -IVa val ues of Assulina muscorum/seminulum higher than other subzones; SII 1 -IVb peak of Botryococcus and de pres sion of Sphag num; SII 1 -IVc depression of Cyperaceae. Lake Suchar II cen tral part (SII 2 ) SII 2 -I SII 2 -II Max i mum of Botryococcus (24%); con tin u ous curve of Cyperaceae; de crease of Bryales (0 2.5%) and Filicales monolete (1 4%); high val ues of Pediastrum angulosum var. angulosum (to 4.5%). Maxima of Sphag num (1.7%) and Pediastrum (5.5%); high val ues of Pediastrum angulosum var. angulosum with max i mum 4.5%; pro por tion of Botryococcus be tween 2 and 21%. The zone was di vided into two subzones: SII 2 -IIa val ues of Botryococcus very higher than up per subzone; SII 2 -IIb de pres sions of Botryococcus and Filicales monolete; high peak of Sphag num. SII 2 -III High val ues of Botryococcus ( %); fall of Filicales monolete; cul mi na tion of Pediastrum angulosum var. angulosum in the top part of the zone. SII 2 -IV Rel a tively low val ues of Botryococcus (1 4%); cul mi na tion of Cyperaceae with max i mum 1.5%. of var i ous spe cies of Sphag num (Dick son 1973; Van Geel 1978), whose spores are rep re sented in large num bers in the sed i ments from the shores of both ex am ined lakes. In the case of Lake Suchar II, the Sphag num ge nus was rep re sented by S. magellanicum and S. fallax (Drzymulska et al., 2015). Entoplyctis lobata (Fig. 4E) oc curs com monly on Oxycoccus palustris, Polytrichum sp., Erica tetralix and other Ericaceae grow ing on the shores of dystrophic lakes (Van Geel 1978). These fungi were also found in side the cells of the aquatic leaves of Sphag num imbricatum (Van Geel 1978), which forms spread ing float ing mats. Based on the re sults ob tained from both lakes (Fig. 2 and 3), it can be con cluded that Entoplyctis lobata fre quently ap pears in the fi nal stage of the for ma tion of the float ing mats, when the pro por tion of the Sphag num is very high. Other re mains of fungi iden ti fied as Type 96A/B (Fig. 4F) were found on liv ing leaves of Oxycoccus palustris (Van Geel 1978), sim i lar to the pre vi ous. They can be rep re sented by the fun gal re mains of one or more spe - cies of the dark-col oured gen era Beltrania, Beltraniopsis, Beltraniella, Ellisiopsis, Pseudobeltrania and Hemibeltrania (Ellis 1971). On the di a gram from the shore of Lake Suchar II Type 96A/B has not been counted yet, al though it was pres ent in all sam ples be tween 0 and 300 cm SII 1. The presence of Helicoon pluriseptatum conidia may be due to the in tense in flow from the catch ment area (Van Geel 1978). This fun gus was found grow ing on de cay ing leaves of birch and nee dles of pine and spruce (Van Beverwijk 1954), and the oc cur rence of its re mains in sed i ments is usu ally linked to an in crease in the share of Pinus and Picea pol len (Yeloff et al., 2007). More over, in the shore sed i ments of both stud ied lakes, es pe cially in Lake Suchar II, the high share of pine and spruce pol len cor re lates very well with the oc cur rence of Helicoon pluriseptatum conidia (Drzymulska et al., 2015). This fun gus was also re cently re corded in an other dystrophic lake in Wigry Na tional Park (Czeczuga 1995). Tes tate amoeba like Amphitrema flavum (Fig. 4L), Assulina muscorum/seminulum (Fig. 4G, 4H), Centropyxis, Hyalosphenia subflava (Fig. 4J), Hyalosphenia papilio (Fig. 4K), Arcella, Nebela (Fig. 4N), Heleopera (Fig. 4M), and Difflugia are con nected with dif fer ent spe cies of Sphag num.

5 NON-POL LEN PALYNOMORPHS CHARACTERISTIC 35 Fig. 2. Lake Suchar II. Sim pli fied pol len di a grams il lus trat ing changes of the lo cal plants and other or gan isms.

6 36 M. FI OC & M. KUPRYJANOWICZ Fig. 3. Lake Œlepe. Sim pli fied pol len di a grams il lus trat ing changes of the lo cal plants and other or gan isms.

7 NON-POL LEN PALYNOMORPHS CHARACTERISTIC 37 The shell-form ing Arcella oc curred com monly in bryophyte peat, par tic u larly among peat mosses (Hoogenraad and Groot 1979; Chardez and Beyens 1987), for ex am ple, S. fuscum (Van Geel et al., 1981) or S. fallax (Lamentowicz et al., 2007a). There are many dif fer ent spe cies of this ge nus, and they are dif fi cult to dif fer entiate in the subfossil ma te rial. Most likely, at least one of the spe cies of this ge nus was A. artocrea (Fig. 4 ), as ev i denced by its size, µm, and appearance ( Arcella and Centropyxis are the most com mon tes tate amoe bae among epiphytic bryophytes (Glime 2013). At least one of the spe cies of Centropyxis was of the Centropyxis aculeata type (Fig. 4I), which ap peared on the Sphag num mat dom i nated by S. fallax, and pre ferred higher ground wa ter lev els (Lamentowicz et al., 2007a). Some spe cies of Centropyxis are as so ci - ated with high ph, e.g. Centropyxis aculeata, C. hirsuta, C. aerophila and C. ecornis, but Centropyxis aculeata has a wide range of tol er ance for en vi ron men tal con di tions (Lamentowicz et al., 2005). Arcella artocrea and Centropyxis aculeata type were found in Lake Œlepe and Lake Suchar II, for ex am ple, on S. fallax (Drzymulska et al., 2015, and un - pub lished data). Hyalosphenia subflava (Fig. 4J) is sup posed to give an in di ca tion of se ri ous dis tur bances in peat growth (Van Geel 1978), and pre fers rel a tively dry con di tions com - pared to other spe cies (Booth 2001). In our re search this taxon was found only in Lake Suchar II af ter a lon ger pe riod of for ma tion of the float ing mats. In high hu mid ity, Hyalosphenia papilio (Fig. 4K) is one of the most nu mer ous spe - cies of peat bog (Lamentowicz et al., 2007a, Glime 2013), and is as so ci ated with acid hab i tats (Lamentowicz et al., 2007a). H. papilio was pres ent only in Lake Œlepe. Assulina muscorum/seminulum are closely re lated to Sphag num (Van Geel 1978). Assulina muscorum ap pears in spring, and A. seminulum in sum mer (Glime 2013). These oligotraphentous thecamoeba are an ad di tional in di ca tor of low nu tri ent con - tent, such as in the drier poor fens (Van Geel et al., 1989). Assulina muscorum/seminulum in habit both com mu ni ties de scribed by Mazei and Tsyganov (2007/08) in the Sphag - num peatlands of Rus sia: bot tom sed i ments of the drain age and the Sphag num quag mire (acc. Glime 2013). The re sults from Lake Suchar II (Fig. 3) con firmed that the early stages of Sphag num peatlands were char ac ter ized by wide spread spe cies such as Assulina muscorum and Arcella arenaria, whereas the sphagnobionts, such as Hyalosphenia, were ab - sent (Mazei and Bubnova 2007; Glime 2013). An other amoeba spe cies, Amphitrema flavum, is characteristic for young Sphag num peat, for ex am ple, Sphag num cf. rubellum peat formed in the Subatlantic (Van Geel 1978). This amoeba is an in di ca tor of a low-nu tri ent sub stra tum (Van Geel et al., 1989) and wet con di tions (Schnitchen et al., 2003). In our study this spe cies reaches its max i mum oc cur rence in the early phase de vel op ment of Sphag num peat, when the wa ter level in creases, as also dem on strated by a study on Jelenia Wyspa mire (Lamentowicz et al., 2007b). Low ph is a char - ac ter is tic fea ture of dystrophic lakes, and in our in ves ti ga - tions such con di tions were in di cated by acidophilic taxa such as Arcella artocrea, Assulina muscorum, Amphitrema flavum and Hyalosphenia sp.. This is con sis tent with other stud - ies (Lamentowicz and Mitch ell 2005; Glime 2013). It has to be pointed out, how ever, that gen era of amoeba such as Arcella, Assulina, Amphitrema and Hyalosphenia are dom i - nant for peatland com mu ni ties. Amphitrema flavum, Assulina muscorum, Assulina seminulum and Hyalosphenia papilio are as so ci ated with the up per parts of Sphag num spec i - mens, as de scribed by Mazei and Tsyganov (2007/08). This is in di cated by other stud ies in Po land on peatlands at dif fer - ent stages of suc ces sion, where the dom i nant spe cies were Amphitrema flavum, Assulina muscorum, Arcella discoides type, and Hyalosphenia papilio, which to gether rep re sented around 60% of the to tal com mu nity count (Lamentowicz and Mitch ell 2005). Sin gly, in the an a lyzed mar ginal pro files (S 1 and SII 1 ), Nebela (Fig. 4N), Heleopera (Fig. 4M) and Difflugia oc curred. These taxa are char ac ter is tic for Sphag numdom i nated peatlands (Lamentowicz and Mitch ell 2005). How ever, their des ig na tion only to the ge nus does not give enough in for ma tion. Microremains, such as eggs and lorica of Rotatoria, co - coons of Turbellaria, spermatophore of Copepoda, and eggs of Tardigrada were non-pol len pallynomorphs of an i mal or i - gin. The re mains of Turbellaria were more nu mer ous in sed i - ments from the cen tral parts of the stud ied lakes than in sed i ments from the mar ginal parts. Eggs of the rotifer Fillinia, rec og nized only to the ge nus, do not pro vide rel e vant in - for ma tion. Lorica of the rotifer Habrotrocha angusticollis (Fig. 4R) oc curred fre quently in the sed i ments of the mar - ginal parts of the in ves ti gated lakes (not in cluded in the di a - grams). They are as so ci ated with mosses in Sphag num bogs, marshes, and with float ing mats and an other emer gent veg e - ta tion along the shores of lakes and ponds through out the world (Murray 1906; Bartos 1951; Haigh 1963; Chengalath and Koste 1983; Francez 1986; Koste and Shiel 1986; Warner and Chengalath 1988, Lamentowicz et al., 2007b). The re sults of our study also in di cate that this rotifer is most char - acteristic for Sphag num hab i tats. Also sper ma to phores of Copepoda ap peared in Sphag num peat from the stud ied pro - files (Fig. 4O). These re mains prob a bly rep re sent dif fer ent spe cies (Van Geel 1978). Pres ent-day copepods have been found, among other places, in the wa ter logged moss layer on peatlands (Rybak and B³êdzki 2005). The re sults of our re - search sug gest that they are sphagnobionts, and ap pear in the late-stage of development of Sphagnum bogs. Macrobiotus harmsworti/richtersi (Fig. 4P), clas si fied by Jankovská (1991) (acc. Montoya et al., 2010), ap peared only in a few sam ples. Eggs of this Tardigrada have been found in moss and foliose li chen (Mayer 2013). SUMMARY AND CONCLUSIONS The trans for ma tion of both stud ied lakes, Lake Œlepe and Lake Suchar II, into a dystrophic state at the be gin ning of the Subboreal re sulted in the ap pear ance of or gan isms typ i cal for wa ter with low ph and poor ac cess to light. Dur ing this stage, Sphag num peat started to ac cu mu late in the mar ginal parts of the lakes. This has given rise to the for ma tion of nu - mer ous moss com mu ni ties form ing float ing mats, which cre - ate a hab i tat for many new or gan isms. In the first stage of dys tro phy, be fore the de vel op ment of Sphag num bog, mainly al gae (Botryococcus and Pediastrum angulosum var. angulosum) and some Rotatoria and Turbellaria were ob served.

8 38 M. FI OC & M. KUPRYJANOWICZ

9 NON-POL LEN PALYNOMORPHS CHARACTERISTIC 39 The sec ond stage be gins with an in crease in the share of Sphag num. Where the Sphag num peat was formed, the share and di ver sity of non-pol len palynomorphs in the mar ginal parts of both in ves ti gated lakes be come higher than in their cen tral parts. In the early part of the stage, Sphag num bogs were char ac ter ized by the spread of tes tate amoeba spe cies such as Amphitrema flavum, Assulina muscorum and Arcella arenaria, whereas the sphagnobionts were ab sent. In a later part of these stage, or gan isms as so ci ated with Sphag num float ing mat, such as spe cies of tes tate amoeba: Assulina seminulum, Hyalosphenia subflava, Hyalosphenia papilio, Arcella, Nebela, Heleopera, Difflugia, single representatives of Rotatoria: Habrotrocha angusticollis, and Tardigrada: Macrobiotus harmsworti/richtersi, some Copepoda and Tillietia sphagni, a par a site fun gus of Sphag num ap peared. At this time, on float ing mats dif fer ent mire plants and the Entoplyctis lobata fungi as so ci ated with them also ex isted. The pres ence of Helicoon pluriseptatum conidia in the an a - lyzed sed i ments sug gests an in ten si fi ca tion of in flow from the catch ment area, where, since the be gin ning of the Subbo real, the share of co nif er ous trees has in creased, and this fun gus was found on the nee dles of pine and spruce. Acknowledgements We thank Danuta Drzymulska, Ph.D., for use ful sug ges tions con cern ing the lake shift to the humic con di tion. Magdalena Fi³oc is a ben e fi ciary of the pro ject Schol ar ships for PhD stu dents of Podlaskie Voivodeship. The pro ject is co-fi nanced by Eu ro pean So cial Fund, Pol ish Gov ern ment and Podlaskie Voivodeship. 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