Studia Quaternaria, vol. 30, no. 1 (2013): 5–17. DOI: 10.2478/squa-2013-0001

LATE GLA CIAL AND HO LO CENE MOL LUS CAN AS SEM BLAGES IN DE POSITS FILLING PALAEOLAKES IN NORTHERN POLAND

Witold Pawe³ Alexandrowicz AGH Uni ver sity of Sci ence and Tech nol ogy, Fac ulty of Ge ol ogy, Geo phys ics and En vi ron ment Pro tec tion, Chair of Geol ogy and Geoturism, Al Mickiewicza 30, 30-059 Cra cow; e.mail: [email protected]

Ab stract Late Glacial and Holo cene carbon ate lacus trine depos its devel oped as lacus trine chalk and calcar e ous gyttja are fairly wide spread across north ern Po land. They form fill ings of palaeolakes which de vel oped dur ing the deglaciation. These for mations are usu ally cov ered by peat. Rich and di ver sified malacofauna has been found in the afore mentioned sed i - ments. The pro files from 154 sites de scribed and pub lished by var i ous au thors were sub jected to malacological anal y - sis. In the whole ma te rial, 18 mol luscan as semblages were dis tin guished, rep re sent ing three types of hab i tats: terres- trial, of tem po rary wa ter bod ies, and of per manent wa ter bod ies. The compo si tion and struc ture of these assem blages al lows char ac ter iz ing cli mate and dif fer en ti at ing hab i tats. The time-se quences of malacological as sem blages pro vided possi bil ity to de fine three types of malacological sequences. On the basis of these succes sions, a scheme of the lake wa ter bod ies evo lu tion dur ing Late Gla cial and in north ern Po land was elab o rated.

Key words: lac us trine chalk, cal car e ous gyttja, palaeolakes, mol lus can as sem blages, Late Gla cial, Ho lo cene, north ern Po land

tic u larly in ten sive in the wa ter bod ies with abun dant veg e ta- IN TRO DUC TION tion which can be asso ci ated with uptake of carbon diox ide The lac us trine chalk and cal car eous gyttja form the fill- by plants dur ing pho to syn the sis. The cal cium car bon ate con- ings of palaeolakes which devel oped in high num bers during tent ex ceeds 80%. The sed iments are white, grey, some times the deglaciation. These may also be found on the coasts of yel lowish or brown ish. Some of them contain small lumps of present-day lakes where they mark their previ ous wider ex- cal cium carbon ate, and sometimes oolith or stromatolith- tents. The ac cu mu lation of these lake sedi ments oc curred type structures (e.g. Rzepecki, 1983; Rutkowski et al., 2002; chiefly dur ing tran si tional pe ri ods be tween cold gla cial pha- Freytet and Verrecchia, 2002). Very of ten, the lac us trine ses and warm interglacials when dead ice blocks, trapped chalk contain remnants of molluscs and ostracods. ear lier in moraine or glaciofluvial sed iments have melted. The name of cal car eous gyttja is given to these lake sedi - Drainage-less depres sions of vari ous sizes, depths and shapes ments that contain from 50 to 80% of cal cium carbon ate. De- were formed, and became stag nant bod ies of freshwaters, sub- pending on the propor tions of other compo nents e.g.: mineral jected to complex evolu tion during the Late Glacial and Ho - or organic substances, or organic detri tus, a num ber of types locene. Some of them transformed into perma nent lakes, are dis tin guished among cal car e ous gyttja. The clas si fi ca tion others were filled with depos its and became peat bogs, and fi- of theses sedi ments were taken up by many studies (e.g. Mar- nally, ter res trial hab i tats with var i ous mois ture lev els. The kowski, 1980; Rzepecki, 1983; Freytet and Verrecchia, 2002). course of sedi ment-fill ing and disap pear ance of these de- The car bon ate de pos its are ac com pa nied by min eral de - pressions was recon structed and described by numer ous au- posits: sands and silts, and more rarely – gravels. Among the thors. The age of great major ity of the sites with carbon ate or grav els, cob bles of Scandi na vian rocks are rep re sented most mineral de pos its fill ing palaeolakes is linked to de clin ing often by granites, gneisses or porphy ries. In terms of their or- phase of the last glaci ation and with the Holo cene. Also igin these are either mo raine or glaciofluvial forma tions. In known are the oc cur rences of these sedi ments of the ear lier many sites these depos its occur at the bot tom of palaeolake interglacials, mainly Eemian and Mazovian (Holsteinian) se quences. (e.g. Skompski, 1996; S. W. Alexandrowicz and W. P. Peats are of es sen tial impor tance. They are as so ciated Alexandrowicz, 2010). with boggy envi ron ments and usually develop in the final Palaeolakes are filled with var ious types of sed iments. stage of evolu tion of lake water bodies. They therefore occur Com mon, and the most im portant from the viewpoint of at the top of depositional sequences and testify to overgrow - malacological stud ies, are lac us trine chalk and cal car e ous ing of lakes and their trans for mation into moist ter res trial en- gyttja. viron ments. Thin interlayers or lenticles of peats are some- Lac us trine chalk is a car bon ate sed i ment oc cur ring as a times found also within the lake sedi ments. Their presence re sult of pre cip i ta tion of CaCO3 in water. This process is par- proves tempo rary shallowing of the lakes. 6 W. P. ALEXANDROWICZ

Fig. 1. Lo cal iza tion of mol lusc-bear ing cal car e ous lake sed i ments in North ern Po land. For ex pla na tions of num bers of lo cal i ties (1–91) see Table 1.

The inter nal struc tures of the palaeolake fillings is of ten and palaeo geo graphi cal re con struc tion based on such data. very com pli cated due to vary ing propor tions and mu tual rela - There are rather few more gen eral stud ies na ture, pre sent ing tion ships be tween the afore men tioned prin ci pal ge netic changes and evo lution of mollus can assem blages over larger types of sedi ments. A total thickness of forma tions of the areas (S. W. Alexandrowicz, 1989; W. P. Alexandrowicz, palaeolakes is from 0.5 to over 10 m. 1999, 2007). Lac ustrine chalk and calcar e ous gyttja often abound in shells of snails and bi valves. It is prin cipally caused by high MA TE RIAL AND METHOD content of CaCO3 in the sedi ments. Remnants of mollusk shells, usu ally less preserved and often bearing the traces of In order to describe the proper ties of changes in se- chem i cal dis so lu tion may some times ap pear also within quences of mollus can assem blages and to carry out re con- peats that ac com pany the car bon ate sed i ments. Pres ence of struc tion of the evolu tion of sedi men tary in lake carbon ate mol lus can shells in the lake sed iments of Late Gla cial and in sed i ments, 91 sites were se lected. Ad di tion ally, the data from northern Poland was recorded as early as in the 19th cen tury 32 sites with fragmen tary malacological data were used as and the early 20th by German geol o gists (e.g. Keilhack, 1888; well as those from 31 sites de scribed in the early 20th cen tury Menzel, 1911). More detailed analy ses of mollus can fauna in by German geol o gists. This last group of sites can be treated lacus trine chalk were under taken by Dembiñska (1924) dur- as a sup ple ment only, as they con tain very gen eral data, and ing inter-war pe riod. After WWII, a great progress was made lo ca tions of most of these sites can not be prac ti cally iden ti- in research of carbon ate lake sedi ments. During the wide- fied at present. Owing to this large number of sites, however, range mapping and deposit docu men ta tion works, more than it was pos si ble to char ac ter ise par tic u lar types of as sem- ten thou sand sites with lac ustrine chalk and calcar e ous gyttja blages of the fauna as well as their com po sitions and struc- were lo cated and cat alogued. Some of them became the top - tures. Re sults of these stud ies were published in a num ber of ics of detailed and inter dis ci plin ary studies, but huge major - scien tific ar ticles (e.g. S. W. Alexandrowicz, 1989; W. P. ity still awaits a more elabo rated re search. De spite the fact Alexandrowicz, 1999, 2007). The study pre sented here rep- that shells of snails and bi valves com monly occur in lac us- re sents an at tempt to gen er alise re sults of stud ies completed trine chalk and calcar e ous gyttja, the state of rec og nition of at var ious sites and to pres ent char acter istics of main fau- malacocoenoses is far for be ing suffi cient. In volu mi nous nistic as semblages and their re la tion ship with cli mate chan- ref er ences there are many men tions on pres ence of malaco- ges and evolu tion of water bodies during Late Glacial and fauna but with out spe cies iden ti fica tion. One can even as- Holo cene. The most im portant sites which provided the basis sume that in many cases the shells of molluscs have been for the presented analy sis are compiled (Table 1, Fig. 1). omitted or gone completely unno ticed. The studies listing On the sites of lac us trine chalk and cal car eous gyttja fau nas, some times with num ber of in di vid u als of par tic u lar (Ta ble 1), rich and di ver sified malacofauna was iden tified. spe cies are rare, and only ex cep tion ally a full malacological The hab i tat and cli ma tic re quire ments of par tic u lar spe cies anal y sis has been done, ac com pa nied by palaeo eco logi cal were discussed in detail in many pub li cations. Sev eral papers LATE GLA CIAL AND HO LO CENE MOL LUS CAN AS SEM BLAGES 7

Ta ble 1 Main lo cal i ties of mol lusc-bear ing cal car e ous lake sed i ments in North ern Po land. For loca tion of partic u lar profiles see Fig. 1

No Local ity Refer ences 43 Lubicz Urbañski (1952) Kopczyñska-Lamparska et al. (1984); 44 Malinowo 1 Niechorze S.W. Alexandrowicz (1989) S.W. Alexandrowicz (1989) 45 Florczaki 2 D¹bki S.W. Alexandrowicz (1991) 46 ¯uchowo Urbañski (1957) Brodniewicz (1979); S.W. Alexandrowicz 47 Kurzêtnik Brodniewicz (1966) 3 Ustka et al. (1989, 1990); S.W. Alexandrowicz D¹browski ed. (1981); (1998); Wojciechowski (2012) 48 Woryty W.P. Alexandrowicz (1999) Krzyszkowski et al. (1998); S.W. 4 Podd¹bie Alexandrowicz (1999b) 49 T³okowo W.P. Alexandrowicz (1999) 50 Bez³awki Œwierczyñski (1958) 5 Kluki Wojciechowski (1995, 2011); S.W. 6 Jezioro £ebsko Alexandrowicz (1999b) 51 Szestno Skompski (1996) 52 Rudkowo 7 Karwia S.W. Alexandrowicz (1999b) S.W. Alexandrowicz (1989) 8 Skowarcz W.P. Alexandrowicz (1999, 2002, 2005) 53 So³dany 9 Chojna Czepiec (1998) 54 Kruklin Stasiak (1963); Czepiec (1997) 10 Brzesko Kowalkowski and Berger (1966) 55 Kulwasy ¯urek and Dziêczkowski (1971) 11 P³oñ S.W. Alexandrowicz (1989) 56 Jezioro G³êbokie W.P. Alexandrowicz (1999) 12 Okrzeszyce 57 Jezioro Wigry W.P. Alexandrowicz (2000, 2009) Kowalkowski and Berger (1971/72) 13 ¯abów 58 Jezioro Gajlik W.P. Alexandrowicz and ¯urek (2010) 14 Zelewo 59 Kurowo S.W. Alexandrowicz and ¯urek (2005) S.W. Alexandrowicz (1989) 15 Lubiatowo 60 Zaczopki W.P. Alexandrowicz and Kusznerczuk (2012) 16 Osiek S.W. Alexandrowicz (1980, 1989) 61 Woroblin S.W. Alexandrowicz (1980); S.W. 62 Radzików Dobrowolski et al. (2012) 17 Prostynia Alexandrowicz and Tchórzewska (1981) 63 Olszewickie B³oto S.W. Alexandrowicz (1983b) S.W. Alexandrowicz and Tchórzewska 64 Cisowe 18 Zdbice S.W. Alexandrowicz and ¯urek (1998) (1981) 65 Zamoœæ 19 Zacisze S.W. Alexandrowicz (1989) 66 G³owina Brykczyñski and Skompski (1979) S.W. Alexandrowicz and Tchórzewska 20 Marcelin 67 Os³onki (1981) S.W. Alexandrowicz (2005, 2008) 68 Zabór 21 Jezioro Du¿e G³uche S.W. Alexandrowicz (1980); S.W. 69 Pomorsko 22 Jezioro Ma³e G³uche Nowaczyk et al. (1999); Nowaczyk and Alexandrowicz and Nowaczyk (1982) 23 Jezioro Charzykowskie S.W. Alexandrowicz (2008) 70 Szumi¹ca S.W. Alexandrowicz (1980) 24 Jezioro Ostrowite 71 Zacisze S.W. Alexandrowicz and Tchórzewska 72 Kie³kowo S.W. Alexandrowicz and ¯urek (1991) (1981); Filonowicz and Krzymiñska 25 Grabowo 73 Niepruszewo-Cieœle Apolinarska and Ciszewska (2006) (1989); S.W. Alexandrowicz (1995b, 1999b) 74 Bogdanowo Skompski (1994) 26 S³upsk S³owiañska 75 Miêtkowo Kasprzak and Berger (1978) 27 S³upsk Rac³awicka 76 Oz Budzyñski Dziêczkowski (1982) 28 S³upsk Po³udnie 77 Smuszewo S.W. Alexandrowicz et al. (1989, 1990) 29 Sto¿ek GlaŸny 78 Jezioro Bnin Wojciechowski (1999, 2000) 30 Dêbica Kaszubska 79 Jezioro Jeziory Wielkie 31 Ga³êŸnia Ma³a 80 Jezioro Kurnickie Wojciechowski (1999, 2000) S.W. Alexandrowicz and Tchórzewska 81 Jezioro Raczyñskie 32 Borzytuchom (1981) 82 Imio³ki Apolinarska and Ciszewska (2006) 33 Jezioro Jasieñ 83 Rybitwy 34 Kozin Florek et al. (1999) 84 JóŸwin Ciszewska (1996) 35 Zawiaty 85 Krucz 36 Orle S.W. Alexandrowicz (1988, 1999b) 86 Chodzie¿ 37 Dol ina Wierzycy B³aszkiewicz and Krzymiñska (1992) 87 Strzelce Dembiñska (1924) 38 Nowe Polaszki S.W. Alexandrowicz (1989) 88 Byszkowice S.W. Alexandrowicz, Tchórzewska 39 Sulêczyno 89 Szarlej (1981) 90 Bytyñ 40 Siwia³ka S.W. Alexandrowicz (1989) 91 Bo¿ejowice Makohonienko et al. (1998) S.W. Alexandrowicz, Tchórzewska 41 £ubiana (1981) 42 Fordon Skompski (1961) 8 W. P. ALEXANDROWICZ

Fig. 2. Eco log i cal struc ture of mol lus can as sem blages. TI . Ter res trial hab i tats: SL – shady en vi ron ments, OP – open en vi ron ments, HG – moist en vi ron ments: TII. Tem po rary bod ies of wa ter: TR – ter res trial en vi ron ments, TM – tem po rary bod ies of wa ter, LA – per manent bod ies of water, TIII. Per manent bod ies of wa ter (lakes): TM – tempo rary bodies of wa ter, LA – per manent bod ies of wa ter (lakes), RE – flow ing wa - ter; ex pla na tions of sym bols mol lusc as sem blages in text.

were used for this re view of bivalves (Piechocki and Dy- distin guish ing the assem blages was described by W. P. Ale- duch-Falniowska, 1993), aquatic (Piechocki, 1979) and ter- xandrowicz (2004) and S. W. Alexandrowicz and W. P. res trial snails (Wiktor, 2004). The spe cies were clas sified Alexandrowicz (2011). into ecolog i cal groups defined and described in de tail by Lozek (1964), S. W. Alexandrowicz (1987), and S. W. Ale- RE SULTS xandrowicz and W. P. Alexandrowicz (2011). The last of the cited papers gave also the basic princi ples for grouping spe- Owing to the abundant mate rial it has been possi ble to cies and defin ing assem blages. Stratigraphic setting of par- distin guish faunistic assem blages. Their structure and com- ticu lar as semblages was de ter mined on the ba sis of their posi tion closely match the condi tions under which the depo - loca tion in the sections, results of other published studies and sition of sedi ments filling the palaeolakes occurred. It anal y ses (palynological, diatomological, and ra dio car bon should, never the less, be stressed that the malacological as so - dat ing) as well as strati graphic char ac ter is tics of in di vid ual ci a tions de fined be low can be much di ver si fied. This di ver si- taxa of molluscs (Lozek, 1964; S. W. Alexandrowicz, 1987; fica tion is caused by spe cific regional condi tions as well as W. P. Alexandrowicz, 2004). Index spe cies for partic u lar as- by vari ous local factors. The assem blages of molluscs identi - sem blages rep re sent ei ther the most typ i cal forms (char ac ter - fied in Late Gla cial and Holo cene lake car bonate sed iments is tic of cer tain types of hab i tats or cer tain cli matic phases) or can be allo cated to one of three groups: assem blages with the taxa oc curring in the high est num bers. This method for pre dom i nant ter res trial spe cies, as sem blages with pre dom i- LATE GLA CIAL AND HO LO CENE MOL LUS CAN AS SEM BLAGES 9

Fig. 3. Stratigraphic range of mollus can assem blages from calcar e ous lake sedi ments. ST. stra tigra phy: PL – Pleniglacial, BL+OD – Older Dryas and Brlling AL – Alleröd, YD – Youn ger Dryas, PB – Preboreal Phase, BO – Boreal Phase, AT – At lan tic Phase, SB – Subboreal Phase, SA – Subatlantic Phase (subdi vi sion of Holo cene based on Starkel (1977)); AS. mol luscan as semblages, TR – ter res trial hab i tats, TM – tem po rary bod ies of wa ter, LA – per manent bod ies of wa ter (lakes).

nant spe cies typ i cal of tem po rary or over grow ing wa ter bod - rica (Müll.), Euconulus fulvus (Müll.). Other ecolog i cal ies, and as sem blages with pre dom i nant spe cies typ i cal of groups usually occur in minor propor tions (Fig. 2TI). The per ma nent, open wa ter bod ies. mentioned as soci a tion has been found in few sites only and occurs within the sedi ments above the lake sequence, thus in- As sem blages with pre dom i nant ter res trial spe cies dicat ing a final phase of fill ing of lake basins. This asso ci a - tion can be corre lated with the period from a decline of (TR) Bo real Phase untill the pres ent (Fig. 3). They are charac ter ised by great di ver sity in both struc - The as sem blage with Pupilla loessica (Pl) is a fauna of ture and compo si tion. In the fillings of palaeolakes these as- very poor spe cies com po si tion. The nom i nal taxon, Pupilla semblages are of lit tle im por tance. They ap pear rarely and muscorum (L.) are accom pa nied by Vallonia tenuilabris were recorded only in some sites. Usually, they occur in the (Brown) and occa sion ally, by the oligothermic bivalve spe- bottom or at the top of sequences, indi cat ing the prelim inary cies of (West.). Such fau nas are typ ical or final phases of lake evolu tion. for a very cold gla cial climate and hab itats of a po lar steppe The assem blage with shade-loving species (Sl) is char- type with many small and shallow water bodies (Fig. 2TI). acter ised by great species diver sity. The domi nant group is Simi lar malacocenoses are commonly re corded in loess for - com posed of shade-lov ing forms, among them both forest mations wide spread throughout Central Europe (S. W. Ale- taxa: Dis cus ruderatus (Fér.), Cochlodina laminata (Mont.), xandrowicz, 1995a; W. P. Alexandrowicz et al., 2002). In and snails prefer ring slightly more open hab itats (open for- northern Poland, this assem blage was found at Kurzêtnik ests, brush zones) – Cepaea hortensis (Müll.), Bradybaena (Brodniewicz, 1966; W. P. Alexandrowicz, 1999, 2007), and fruticum (Müll.). A propor tion of forms of shaded habi tats in sed i ments un der ly ing lake for ma tions only. The as so ci a- usually reaches up to 50%. The sup plement ing el ements in- tion with Pupilla loessica repre sents a declin ing part of clude taxa with wide eco log ical toler ance: Cochlicopa lub- Pleniglacial and the ear liest part of Late Gla cial (Fig. 3). 10 W. P. ALEXANDROWICZ

The as sem blage with Vallonia pulchella (Vpu) is mar- portance (Fig. 2TI). The sec ond es sen tial com po nent in- ked by a predom inant pro portion of snails of open grass land cludes euryecological mesophilous forms: Euconulus fulvus hab i tats with vari able mois ture con tents: Vallonia pulchella (Müll.) and Nesovitrea hammonis (Ström). The as semblage (Müll.), Vallonia costata (Müll.), and Pupilla muscorum (L.) can be sup ple mented by oligothermic taxa char acter istic for (Fig. 2TI). They are sup plemented by mesophilous taxa: shaded hab i tats: Semilimax kotulae (West.) and Arianta ar- Cochlicopa lubrica (Müll.), Euconulus fulvus (Müll.), and bustorum (L.). Oc cur rence of aquatic molluscs such as: hygrophilous taxa: Succinea putris (L.), Carychium mini - Gyraulus laevis (Ald.), lapponicum mum (Müll.). At some sites, an es sen tial compo nent of the (Cless.) and Galba truncatula (Müll.) in di cates pres ence of afore men tioned as so ci a tion is formed by aquatic molluscs small, much overgrown or tem po rary water bod ies. The liv ing in small and tem po rarily dis ap pear ing wa ter bod ies: malacocoenosis with Ver tigo genesii is in dic a tive of cold cli - Planorbis planorbis (L.), Galba truncatula (Müll.). The mate and dom ina tion of moist, usually open habi tats of tun- shade-lov ing species are prac tically absent. At many sites, dra type, and is char ac ter istic for Late Gla cial, and par- the fauna with Vallonia pulchella appears at the top of lake ticularly the Younger Dryas. The upper limit of its occur - de pos its, in di cat ing the phases of dis ap pear ance of wa ter rence is marked by the phase of rapid warm ing, mani fested in bod ies and their transfor ma tion into moist grass lands. This the Preboreal Phase of the Holo cene (Fig. 3). malacocoenosis is char ac ter istic for mid-and late Ho lo cene The as so ci a tions of sim i lar com po si tions were de scribed (Fig. 3). in many sec tions of ge net i cally di verse sed i ments through out The as semblage with mesophilous spe cies (Me) is mar- Europe (e.g. Lozek, 1964; S. W. Alexandrowicz, 1987; ked by presence of all eco logi cal groups, al though the forest Limondin-Lozouet, 1992; Krolopp and Sümegi, 1993; S. W. and brush species, and aquatic forms are of sec ond ary im por- Alexandrowicz and W. P. Alexandrowicz, 1995; W. P. tance (Fig. 2TI). The most impor tant, however, are meso- Alexandrowicz, 1997, 1999, 2004). philous forms: Cochlicopa lubrica (Müll.), Euconulus fulvus (Müll.), Nesovitrea hammonis (Ström), and Ver tigo angu- As sem blages with pre dom i nant spe cies typ i cal for stior (Jeffr.). These are supple mented by taxa of open: Vallo- tem po rary or over grow ing wa ter bod ies (TM) nia pulchella (Müll.), Vallonia costata (Müll.), and of moist en vi ron ments: Succinea putris (L.). The fauna hereby de- The as sem blage with Valvata cristata (Vcr) is a rich and scribed rep re sents the phases of dis ap pear ance of wa ter bod - diver si fied malacocoenosis in which two groups of aquatic ies and is typi cal for the whole Holo cene, and it occurs molluscs predom i nate. The first, and usually more numer ous par tic u larly of ten dur ing Bo real and At lan tic Phases of the group consists of species that are typ ical for shal low, small Ho lo cene (Fig. 3). and much overgrown water bodies: Valvata cristata (Müll.), The as sem blage with Ver tigo parcedentata (Vpa) is a Valvata macrostoma (Mörch), Acroloxus lacustris (L.), and very rare malacocoenosis, marked by presence of oligother- other species. The sec ond group includes forms charac ter is- mic hygrophilous species: nom inal taxon, Ver tigo genesii tic for tem po rarily dis ap pear ing wa ter bod ies: Galba trunca- (Gredl.), and mesophilous species: Columella columella (G. tula (Müll.) and Anisus leucostomus (Mill.). There are often Mart.). Also pres ent are aquatic spe cies with wide ther mal tol - also euryecological aquatic molluscs: er ance – Gyraulus laevis (Ald.) (Fig. 2TI). The taxa of open (Poli.), Radix baltica (L.), and even rheophilous taxa as and shaded envi ron ments are ab sent. This fauna repre sents the (Müll.). The ad mixture of ter res trial spe - tundra with moist substrate, very cold, po lar cli mate and is cor- cies, chiefly hygrophilous, is usually fairly sig nifi cant. More re lated with the Pleniglacial and early Late Gla cial (Fig. 3). open biotopes are in dicated by occur rence of Succinea putris The as sem blage with Perforatella bidentata (Pb) is a (L.) and Zonitoides nitidus (Müll.) while in more shaded hab- malacocoenosis poor in terms of species compo si tion, mar- itats, mesophilous species are com mon: Ver tigo angustior ked by pre dom i nance of nom i nal spe cies. Perforatella bi- (Jeffr.) and Ver tigo substriata (Jeffr.) along with forms typ i- dentata (Gmel.) is a shade-loving taxon, in habit ing forests of cal of water logged woods as Perforatella bidentata (Gmel.) alder swamp type, growing on wa ter logged sites (Fig. 2TI). (Fig. 2TII). The pre sented malacocoenosis is typ ical for a Besides the nom inal species, hygrophilous forms: Succinea biotope with a small, shal low wa ter body, grad u ally filled putris (L.), Zonitoides nitidus (Müll.), and aquatic molluscs with sed iments. The char acter istic feature of such small lake liv ing in tem po rary wa ter bod ies: Galba truncatula (Müll.), is an abun dant de vel op ment of veg e ta tion, com posed mostly Anisus leucostomus (Mill.) are also impor tant. Re maining of immersed plants. The compo si tion of terres trial fauna ac- eco log i cal groups are rep re sented spo rad i cally only. The as- com pany ing the aquatic spe cies depends closely on features sem blage with Perforatella bidentata is typ i cal for early Ho- of the envi ron ment surround ing the water body. The malaco- locene, and par ticu larly for Boreal Phase (Fig. 3). This coenosis is com posed of species with great ther mal toler ance as so ci a tion de vel oped most of ten on shores of ma jor lakes and can appear in various climatic stages of the Holocene, during low water level peri ods, in sedi ments that form inserts and even in the Late Glacial sediments (Fig. 3). in gyttja and lacus trine chalk. The as sem blage with Pisidium stewarti (Pst) is indi cated The as sem blage with Ver tigo genesii (Vg) is common by domi nance of bivalves. Along with the nomi nal species, and, at the same time, one of the most char acter istic. The spe - there are also: (Cless.), Pisidium obtu- cies prefer ring moist, sometimes wa ter logged biotopes, de- sale lapponicum (Cless.), and some times Pisidium hiber- vel op ing un der cold con ti nen tal cli mate: Ver tigo genesii nicum (West.). Snails are rep re sented by oligothermic aqua- (Gredl.), Ver tigo geyeri (Lindh.), Columella columella (G. tic forms as Gyraulus laevis (Ald.) as well as by hygrophilous Mart.) and Ver tigo parcedentata (Braun.) are of primary im - spe cies: Ver tigo genesii (Gredl.) and Ver tigo geyeri (Lindh.). LATE GLA CIAL AND HO LO CENE MOL LUS CAN AS SEM BLAGES 11

This fauna devel oped in shallow and inten sively overgrown, char acter ized by predom i nance of opercula over shells and is perma nent lakes (Fig. 2TII). It is char ac ter is tic of cold cli mate in dic a tive of pres ence of compact reeds. The sec ond type in - andcorresponds to the Late Glacial (Fig. 3). dicates preva lence of shell over opercula and corre sponds to The as sem blage with Pisidium obtusale lapponicum the lit toral zone of the lake (Steenberg, 1917; S. W. Alexan- (Pol) is a malacocoenosis poor in spe cies com po sition, con- drowicz, 1999a). The admix ture of terres trial fauna is usually sisting chiefly of aquatic species, among which the forms liv- minor. This assem blage is often found in early and mid-Ho- ing in small, very shal low, in tensively overgrown and tem- locene depos its. It occurs only occasionally in the Late porarily disap pear ing water bodies, typi cal for a cold climate. Glacial formations (Fig. 3). Apart from the nomi nal taxon, there are also oligothermic The as sem blage with Valvata piscinalis (Vpi) has a poor forms: Pisidium lilljeborgii (Cless.) and Pisidium stewarti com po si tion in di cated by pre dom i nance of nom i nal spe cies, (Pat.) as well as spe cies of wide ther mal toler ance: Galba ac com pa nied by euryecological bi valves: Pisidium caserta- truncatula (Müll.) and Ra dix baltica (L.). Ter res trial spe cies num (Poli.), (Malm), and snails – are few and they are most often repre sented by hygrophilous Bithynia tentaculata (L.) (both shells and opercula). Terres - and oligothermic snails as Ver tigo genesii (Gredl.) (Fig. trial spe cies are prac tically absent (Fig. 2TIII). This as so ci a- 2TII). This malacocoenosis is typi cal of Late Gla cial, par ticu - tion usually devel ops in deeper parts of the lakes, on a muddy larly of the Younger Dryas. It also occurs rarely at the begin - bottom without rooted vege ta tion. The fauna with Valvata ning of the Holo cene. It does not appear in its ypunger part, piscinalis oc curs both in Late Glacial and Ho lo cene sed i- starting from the Boreal Phase (Fig. 3). ments. In the lat ter case, it is usu ally more di ver si fied in its The as sem blage with Gyraulus laevis (Gl) is char ac ter - species composition (Fig. 3). ised for its abundant occur rence of oligothermic aquatic The as sem blage with (Pm) is predom i- snails. Apart from the most char ac ter is tic nomi nal taxon, nated by bivalves of the Sphaeridae family. Apart from the there are also Pisidium obtusale lapponicum (Cless.) as well nom inal taxon, the most fre quently are: Pisidium casertanum as other spe cies of wide ther mal toler ance such as: Ra dix (Poli.), Pisidium subtruncatum (Malm), and Sphaerium cor- balthica (L.), Galba truncatula (Müll.) and others. The ter- neum (L.) as well as (Shep.) and res trial fauna with preva lence of oligothermic spe cies: Ver- (A. Schmidt). Warmer phases are in di - tigo genesii (Gredl.), Ver tigo geyeri (Lindh.) and mesophi- cated by pres ence of (Pal.). Aqua- lous species usually ap pears as ad mixture, al though even up tic snails are more rare and are repre sented by lake forms: to 40% (Fig. 2TII). This as sem blage is char ac ter is tic for Bithynia tentaculata (L.) and Stagnicola turricula (Held), small, shal low but not dry ing lakes of a cold cli mate. The as- while ter res trial spe cies are ab sent (Fig. 2TIII). This as sem - sem blage with Gyraulus laevis is char ac ter is tic for warmer blage is typi cal of open lake water bodies where it lives in the phases of the Late Glacial or the begin ning of the Holo cene. zones charac ter ized by poor veg eta tion, both above the water In the up per part of the Holo cene, the nom inal species occurs sur face and im mersed. The malacocoenosis with Pisidium extraordinarily rarely (Fig. 3). milium occurs exclu sively in the Holo cene sedi ments (Fig. 3). The as sem blage with (Pn) is indi cated As sem blages with pre dom i nant of spe cies typ i cal by major propor tion of bivalves among which a major role is played by Pisidium nitidum (Jen.), Pisidium amnicum for per ma nent open wa ter bod ies (LA) (Müll.), Pisidium casertanum (Poli.) and Pisidium subtrun- The as semblage with Lymneaidae (Ly) is a rich and di - catum (Malm). Aquatic snails are rare while the ad mix ture of versi fied malacocoenosis, consist ing chiefly of aquatic spe- terres trial forms does not exceed 20% (Fig. 2TIII). This cies of wide eco log i cal tol er ance, in hab it ing per ma nent malacocoenosis is distinct for its large contents of species wa ter bod ies: Ra dix baltica (L.), Stagnicola turricula (Held), that require running wa ter. Such fauna devel ops in lowland Stagnicola palustris (Müll.), Pisidium casertanum (Poli.), rivers with slow current, or in bot tom parts of large lakes, in and many other species. Forms typi cal of small water bodies: the zones in which the currents occur. The assem blage with Acroloxus lacustris (L.), Gyraulus albus (Müll.), and some- Pisidium nitidum was found in the sedi ments allo cated to times even forms typi cal for tempo rarily drying zones: Galba Late Gla cial, and to early and mid-Holocene (Fig. 3). truncatula (Müll.) ap pear as ad mix ture. Ter res trial spe cies The as sem blage with Dreissena polymorpha (Dp) is pre- are either absent or appear only as single shells (Fig. 2TIII). domi nated by the nomi nal species up to 95%. This assem - The com po si tion and struc ture of this as so ci a tion are very blage oc curs in lakes, char ac ter istic for its dif fer ent na ture variable and depend closely on nature of the water body. The and tro phy. The nom inal species is remark ably euryecolo- assem blage with Lymnaeidae de velops during the whole Ho- gical, but it prefers partic u larly to colo nise hard surface locene, both in small, shallow and inten sively overgrown zones (cobbles, shells of large bivalves as Unio or Anodonta lakes and in larger lakes of different trophism (Fig. 3). genera, anthropogenic hydrotechnical struc tures) where it The as sem blage with Bithynia tentaculata (Bt) is one of de vel ops huge pop u la tions (Stañczykowska, 1977) (Fig. the most char ac ter is tic and most com monly oc cur ring as so ci- 2TIII). Pres ence of Dreissena polymorpha (Pall.) in the sed i - ations in lake carbon ate sedi ments. In this malacocoenosis, ments indi cates their young age. This form was in troduced the most impor tant spe cies is Bithynia tentaculata (L.) which acci den tally into Poland and has been recorded there since is some times ac com pa nied by nu mer ous forms char ac ter is tic the be ginning of the 19th cen tury (W.P. Alexandrowicz and for perma nent water bodies (Fig. 2TIII). The fauna with S.W. Alexandrowicz, 2010) (Fig. 3). Bithynia tentaculata is typ ical for semi-open lakes, partly The as sem blage with Potamopyrgus antipodarum (Pa) overgrown with reeds. There are 2 types of it. The first one is forms a poor as so ci a tion, pre dom i nated by the nom i nal spe- 12 W. P. ALEXANDROWICZ cies, ac compa nied by other aquatic forms typi cal for lakes: fauna with Ver tigo genesii) be came in creas ingly im por tant Ra dix baltica (L.), Stagnicola turricula (Held), Bithynia as well as as so cia tions typ ical for small, shal low and densely tentaculata (L.), as well as spe cies typi cal for tempo rarily over grown wa ter bod ies (as sem blage with Pisidium stewarti dried-up zones as Galba truncatula (Müll.) (Fig. 2TIII). and with Pisidium obtusale lapponicum). Starting from the Potamopyrgus antipodarum (Gray) is a young im migrant, early Holo cene, the propor tion of aquatic and swamp taxa recorded in Poland since 1933 (W.P. Alexandrowicz and declines gradu ally and shadow-loving, open-country and S.W. Alexandrowicz, 2010) and it is therefore a good age mesophilous snails take over (Fig. 4). indicator (Fig. 3). The aforemen tioned sequence is a record of evolu tion of small wa ter bodies which emerged at the end of the Late Gla - cial as an ef fect of melting of small dead ice blocks. Small DIS CUS SION and shal low lakes were rapidly filled with sedi ments and The mol lusc as sem blages de velop in def i nite hab i tats. transformed into swamps. Such transfor ma tion took place Their nature depends on two princi pal factors. The first one is during the Younger Dryas or during the early Holo cene. cli mate which de ter mines geo log i cal pro cesses and liv ing From the begin ning of the Atlan tic Phase, progres sive drying condi tions in a regional scale. Within the most recent 15 000 and replace ment of moist swampy habi tats by drier grassland years, two peri ods, utterly differ ent from the viewpoint of cli- and/or forest habitats occurred (Fig. 4). matic condi tions, can be dis tin guished. The older period cov- Type B is the most fre quent one of the malacofauna as- ers a ter mina tion of the last gla ci ation i.e. the Late Gla cial. It semblage in carbon ate lake sed iments (W.P. Alexandrowicz, was a cold phase with two warmer fluctu a tions (BÝlling and 1999, 2007). This sequence begins with a phase pre dom i- Alleröd) which were only very weakly indi cated in malaco- nated by open terres trial habi tats of tundra type. Charac ter is - log i cal suc ces sions of car bon ate lake sed i ments. This pe riod tic features of this inter val include occur rence of oligother- was asso ci ated with progress ing deglaciation, when massive mic as sem blages with Pupilla loessica (on drier sites) or with melting of dead ice blocks led to the emer gence of lakes and Vertigo parcedentata (moist and water logged biotopes). In swamps. The younger period corre sponds with the Holo cene younger Late Glacial, a phase of peat bog devel op ment is when sev eral colder stages are occurred, sepa rated by war- com monly ob served. Pres ence of char ac ter is tic malacoco- mer fluctu a tions. The Holo cene is not only repre sented by enosis with Ver tigo genesii is linked to this period. In water - devel op ing lakes but, in many cases, also by their overgrow - logged areas, there were small, shallow and much overgrown ing, shallowing and gradual disap pear ance (e.g. Starkel, melt-out lakes, in hab ited by as so ci a tions with Pisidium ste- 1977; Ralska-Jasiewiczowa and Starkel, 1988; Ralska- warti and Pisidium obtusale lapponicum. Pro gres sive melt - Jasiewiczowa, 1989; Birks and Birks, 1989). ing of large dead ice blocks during the Younger Dryas and at Local condi tions are the second princi pal factor that de- the be ginning of the Holo cene led to transfor ma tion of these ter mines de vel op ment and vari abil ity of malacocoenoses. habi tats into major lakes. This stage is as so ci ated with oc cur- Great di ver sity of hab itats can re sult in oc cur rence of ut terly rence of rich as semblages of aquatic molluscs, par ticu larly dif fer ent faunistic assem blages in sed iments of the same age. with Bithynia tentaculata. It is also a stage of in ten sive ac cu - The Late Gla cial is asso ciated with oc cur rence of rel a tively mula tion of carbon ate sedi ments, a thickness of which may poor malacoenoses. These are char ac ter ised by sig nif icant even exceed 10 m. Lac us trine chalk and cal car eous gyttja propor tion of oligothermic species and forms of wide eco- gradu ally filled lake basins, leading to their disap pear ance. logi cal toler ance as well as with occur rence of high numbers The final stage of fill ing of the lakes resulted in their trans for - of molluscs living in shallow water bodies, boggy zones and mation into peat bogs. This is confirmed by presence of peat open ter res trial hab itats. The fau nas of the Ho lo cene are usu - lay ers over ly ing the car bon ate sed i ments. The mol lus can ally more di versi fied and include also species with high eco - com mu ni ties show also ev i dent changes. The as sem blages log i cal re quire ments. The taxa as so ci ated with large water with prev alence of spe cies typi cal of lakes are re placed by as - bodies and shaded terrestrial habitats are also common. so ci a tions dom i nated by taxa of shal low and in ten sively It is possi ble to distin guish three types of malacological over grown, some times even ep i sodic wa ter bod ies (as sem - suc ces sions in the ana lysed ma te rial, in di cat ing di ver si fi ca - blage with Valvata cristata). In the late Ho lo cene, these wa- tion of habi tats and stratigraphic sequences of fauna assem - ter logged zones were transformed into ter restrial hab itats, blages. These anal yses of such se quences were at tempted ei ther open (as sem blage with Vallonia pulchella) or shaded only rarely (S. W. Alexandrowicz, 1983a; W. P. Alexan- (assemblage with shadow-loving species) (Fig. 4). drowicz, 1997, 2004), although they provide am ple grounds Type C suc ces sion is char ac ter istic for large lakes. These for in ter pre ta tions and re con struc tions of palaeoenviron- water bodies devel oped from exten sive cave-in lakes, result - ments as well as their changes in time. ing from melt ing dead ice blocks and formed as late as in the Type A is in dicated by dom ina tion of assem blages with Late Gla cial. This stage is as so ciated with oc cur rence of as - major propor tion of terres trial molluscs. They are rare un sem blages with mi nor pro por tion of ter res trial spe cies e.g. lake depos its, but much more often in calcar e ous tufas in fauna with Valvata piscinalis. A domi na tion of aquatic mol- northern Poland (Skompski, 1961; Brykczyñski and Skomp- luscs maintains throughout the whole Holo cene. The chan- ski, 1979) as well as in the south (W. P. Alexandrowicz, ges of wa ter level in lakes (Ralska-Jasiewiczowa and Starkel, 2004). The sequence starts with a malacocoenosis predom i- 1988; Wojciechowski, 1999, 2000) were marked by peri odic nated by oligothermic terres trial species prefer ring open hab- occur rence of assem blages with greater propor tions of taxa i tats (as sem blage with Pupilla loessica). In the final part of typ i cal of shal low wa ter bod ies (as sem blage with Valvata the Late Gla cial, the hygrophilous as semblages (chiefly the cristata) or even water logged terres trial habi tats (assem - LATE GLA CIAL AND HO LO CENE MOL LUS CAN AS SEM BLAGES 13

Fig. 4. Succes sions of mollus can as semblages from lake sed i ments in North Poland. A-type, B-type, C-type – suc ces sions of assem blages described in text: PL – Pleniglacial, LG – Late Glacial, EH – Early Ho locene, MH – Middle Holo cene, LH – Late Holo cene, TR – terres trial en vi ron ments, HG – moist envi ron ments, TM – tem po rary bod ies of wa ter, LA – per manent bod ies of wa ter (lakes). blage with Perforatella bidentata). In the top part of the se - steppe-like hab itats to moist and water logged tun dra bio- quence of lake sed iments, the as semblages with Potamo- topes. The first stage of lake devel op ment is connected with a pyrgus antipodarum and with Dreissena polymorpha occur warming of the Alleröd Interphase. Massive melting of dead and indicate a human impact (Fig. 4). ice blocks led to emergence of numer ous melt-out lakes (e.g. W. P. Alexandrowicz, 1999, 2007; Wojciechowski, 1999, 2000). Thus ex tensive but shallow and inten sively over- CON CLU SIONS growing wa ter bod ies were in habited by rel atively poor mol- On the basis of the anal ysis of features and se quences lus can as sem blages with pre dom i nant oligothermic spe cies. per tain ing to mol lus can as sem blages oc cur ring in lac us trine During the Younger Dryas, many of these wa ter bod ies were chalk and calcarous gyttja, it was possi ble to compile the com pletely filled with sed iments and transformed into peat scheme of evolu tion of lake water bod ies. The old est mollus - bogs. A char ac ter is tic as sem blage with Ver tigo genesii re- can as sem blages doc u ment the pre-lac us trine ter res trial garded as indi ca tor of the afore mentioned phase is asso ci ated phase, corre spond ing to the end of Pleniglacial, and with with these habi tats (Lozek, 1964; S. W. Alexandrowicz, early Late Glacial. The malacoenoses in di cate pres ence of 1987; Limondin-Lozouet, 1992; Krolopp and Sümegi, 1993; open habi tats with varying moisture levels, from dry Arctic S. W. Alexandrowicz and W. P. Alexandrowicz, 1995; W. P. 14 W. P. ALEXANDROWICZ

Fig. 5. Fig. 5. Late Gla cial and Holo cene en vi ronmen tal changes in North Poland in the light of malacological anal ysis of cal car e ous lake sedi ments. ST. stra tig ra phy: BL+OD – Older Dryas and Brlling AL – Alleröd, YD – Younger Dryas, PB – Preboreal Phase, BO – Boreal Phase, AT – At lan tic Phase, SB – Subboreal Phase, SA – Subatlantic Phase (subdi vi sion of Holo cene based on Starkel (1977)); W – high-level wa ter pe ri ods (af ter: Wojciechowski, 1999, 2000), N – low-level wa ter (af ter: Wojciechowski, 1999, 2000).

Alexandrowicz, 1997, 1999, 2004). On the other hand, how- ated. A begin ning of the Holo cene was indi cated by rapid ever, the de clin ing pe riod of the Late Gla cial was in di cated warming that deter mined a subse quent stage of lake water by rising water levels in lakes. This phenom e non, observed bod ies. De creas ing pre cip i ta tion re sulted in a drop of wa ter in almost entire northern Poland (Wojciechowski, 1999, level in lakes (Ralska-Jasiewiczowa and Starkel, 1988; Woj- 2000) can be connected with gradu ally moister climate. Ow- ciechowski, 1999, 2000). This fact, in com bina tion with ing to that, many melt-out lakes were transformed into larger gradual fillings of lake basins with sedi ments resulted in dis- and deeper lakes, in which carbon ate depo si tion was initi - appear ance of many shallow lakes and in peat bogs emerging LATE GLA CIAL AND HO LO CENE MOL LUS CAN AS SEM BLAGES 15 in their places. During mid- and late Holo cene these peat Alexandrowicz, S.W., 1991. The malacofauna of the Ho lo cene lac - bogs were transformed into grass land and forest biotopes. ustrine sed i ments of D¹bki near Dar³owo. Przegl¹d Archeolo- The aforemen tioned drop of water level were much less pro- giczny 28, 19–24. nounced in large lakes, in which depo sition of lacus trine Alexandrowicz S.W., 1995a. Malacofauna of the Vistulian loess in Cra cow Re gion (S Poland). Annales Universitatis Mariae Cu - chalk and calcar e ous gyttja with rich and diverse mollus can rie-Sk³odowska B, 50, 1–28. assem blages has been contin ued throughout early and mid- Alexandrowicz, S.W., 1995b. Malacofauna of the Holo cene sed i - Ho lo cene. Dur ing late Ho lo cene, a re mark able slow ing ments in Grabowo near Koszalin (Pomerania, North Po land). down and in many cases, dis ap pear ance of car bon ate sed i- Questiones Geographicae 4, 13–20. men tation oc curred. Cli mate changes re flected in chang ing Alexandrowicz, S.W., 1998. Ra dio car bon-dated as sem blages of water levels in lakes are also common during this period molluscs in the Pol ish Costal Zone near £eba and Ustka. Field (Ralska-Jasiewiczowa and Starkel, 1988; Wojciechowski, sym posium on glacial geol ogy at the Bal tic See Coast in North - 1999, 2000). During the last one hundred years, human im- ern Po land. Ex cur sion guide, 54–56. pact has become more and more notice able in the envi ron - Alexandrowicz, S.W., 1999a. Bithynia tentaculata (Linnaeus, ment. It is marked in malacocoenoses n lake sediments either 1758) as an in di ca tor of age and de po si tion en vi ron ment of Qua ter nary sed i ments. Folia Malacologica 7, 79–88. by impoverishment of species composition resulting from Alexandrowicz, S.W., 1999b. Malacofauna and stra tig ra phy of progressive eutrophication or by occurrence of expansive Late Vistulian and Holo cene depos its in the Southern Bal tic species alien to Polish fauna that indicate anthropogenic coastal zone. Qua ter nary Stud ies in Po land, special issue, migrations (Fig. 5). 37–50. Changes in mol lus can as sem blages en able dis tin guish- Alexandrowicz, S.W., 2005. Molluscs assem blages of Late Glacial ing of several epi sodes of peat bog devel op ment and disap - and Ho lo cene sed i ments in South ern Kujawy. Prace Komisji pear ance of lake wa ter bod ies. These oc curred in the decli- Paleogeografii Czwartorzêdu PAU 3, 147–152 (in Pol ish with ning part of the Younger Dryas, begin ning of the Atlan tic Eng lish sum mary). Phase and in the Subatlantic Phase (Fig. 5). These ep isodes Alexandrowicz, S.W., 2008. Malacofauna of Late Quater nary lac - evi dently coin cide with phases of lower water levels in lakes ustrine de posits at Os³onki (Kujawy, Cen tral Poalnd. Folia Quaternaria 78, 51–69. (e.g. Starkel, 1977; Ralska-Jasiewiczowa and Starkel, 1988; Alexandrowicz, S.W., Alexandrowicz, W.P., 1995. Mollus can Wojciechowski, 1999, 2000). fauna of the Up per Vistulian and Early Holo cene sed i ments of Recon struc tion of the evolu tion of lake water bod ies in South Poland. Biuletyn Peryglacjalny 34, 5–19. northern Poland, based on anal ysis of mollus can assem - Alexandrowicz, S.W., Alexandrowicz, W.P., 2010. Molluscs of the blages, sup ple ments sim i lar re con struc tions and pat terns de- Eemian Inter glacial in Poland. Annales Societatis Geologorum fined under differ ent methods and published in numer ous Poloniae 80, 69–87. studies, both in Poland (e.g. Goslar et al., 1993; Starkel et al., Alexandrowicz, S.W., Alexandrowicz, W.P., 2011, Malacological 1996; ¯urek and Pazdur, 1999; Wojciechowski, 1999, 2000) anal y sis – method of re search and in ter pre ta tions. Rozprawy and in northern Europe (e.g. Mania, 1973; Mania and Toe- Wydzia³u Przyrodniczego PAU 3, 5–302 (in Pol ish). pfer, 1973; Sunderlau, 1975; Griffiths et al., 1994; Magny Alexandrowicz, S.W., Cichosz-Kostecka, A., Florek, E., Florek, and Ruffaldi, 1995). W., Or³owski, A., R¹czkowski, W., Zachowicz, J., 1989. The evolu tion of the S³upia River Val ley in the Late Vistulian and Ho lo cene. Geologia, Kwartalnik AGH 15, 5–218 (in Pol ish Ac knowl edg ments with English summary). Alexandrowicz, W.P., Florek, W., Zaborowska, K., Zachowicz, J., This study has been spon sored by the AGH Univer sity of Sci - 1990. The S³upia up per floodplain in the vi cin ity of S³upsk, ence and Tech nol ogy through the Uni ver sity grant no. 11.11. Pomerania, Po land. Quaestiones Geographicae 11/12, 5–27. 140.173. Alexandrowicz, S.W., Nowaczyk, B., 1982. Late Gla cial and Holo - REF ER ENCES cene lake sedi ments at Pomorsko near Sulechów. Questiones Alexandrowicz, S.W. 1980. Mol luscan as semblages in lac ustrine Geographicae 8, 5–17. chalk of Lubusz Land. Kreda jeziorna i gytie 2, 24–32 (in Pol - Alexandrowicz, S.W., Tchórzewska, D., 1981. Bog lime in Qua ter - ish with English summary). nary de posits of the Mid dle Pomerania. Geologia, Kwartalnik Alexandrowicz, S.W., 1983a. Malacofauna of the Holo cene calcar - AGH 7, 59–71(in Pol ish with Eng lish summary). e ous sedi ments of the Cra cow Upland. Acta Geologica Polo- Alexandrowicz, S.W., ¯urek, S., 1991. Malacofauna od Ho lo cene nica 33, 117–158. lac ustrine sed i ments and peat-bogs in the vi cin ity of Wolsztyn Alexandrowicz, S.W., 1983b. Malacologic anal y sis of the soil pro - (West ern Po land). Geologia, Kwartalnik AGH 17, 221–234 (in file in the southern strip of swamp in the Kampinos Pri meval Polish with English summary). Forest. In: Wp³yw dzia³alnoœci cz³owieka na œrodowisko gle- Alexandrowicz, S.W., ¯urek, S., 1998. Ho lo cene lac ustrine sed i - bowe w Kampinoskim Parku Narodowym. Wydawnictwa ments in the Mid dle Swamp Strip of the Kampinos Forest. SGGW, 215–227 (in Polish with Eng lish summary). Geologia, Kwartalnik AGH 24, 333–351 (in Pol ish with Eng - Alexandrowicz, S.W., 1987. Malacological anal ysis in Quater nary lish summary). re search. Geologia, Kwartalnik AGH 12, 3–240 (in Pol ish with Alexandrowicz, S.W., ¯urek, S., 2005. Landscape and peatlands of Eng lish sum mary). the Narew National Park. Chroñmy Przyrodê Ojczyst¹ 61, Alexandrowicz, S.W., 1988. Mol luscan as semblages of the lake 5–18 (in Pol ish with Eng lish summary). sedi ments in the ancient melt-lake Orle. Folia Quaternaria 58, Alexandrowicz, W. P., 1997. Malacofauna of Qua ter nary de posits 59–67. and en vi ron mental changes of the Podhale Basin during the Alexandrowicz, S.W., 1989. Molluscs as soci a tions in Late Qua ter - Late Vistulian and Holo cene. Folia Quaternaria 68, 7–132 (in nary lake de posits of north ern Po land. Studia i Materia³y Polish with English summary). Oceanologiczne 56, 267–276 (in Pol ish with Eng lish summary). Alexandrowicz, W.P., 1999. Evo lu tion of the malacological as sem- 16 W. P. ALEXANDROWICZ

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