Mol lusc fau nas of lake de pos its in Gorzów Wielkopolski (NW Po land) as an in di ca tor of en vi ron men tal changes dur ing Eemian and Early Weichselian
Witold Pawe³ ALEXANDROWICZ1, *, Sylwia SKOCZYLAS1, Artur SOBCZYK2, Krzysztof STEFANIAK3, Adam KOTOWSKI3, Bo gus³aw PRZYBYLSKI4, Dariusz CISZEK4,
Janusz BADURA4 and Krzysztof URBAÑSKI4
1 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, Al. Mickiewicza 30, 30-059 Kraków, Po land
2 Uni ver sity of Wroc³aw, In sti tute of Geo log i cal Sci ences, pl. M. Borna 9, 50-024, Wroc³aw, Po land
3 Uni ver sity of Wroc³aw, In sti tute of En vi ron men tal Bi ol ogy, Sienkiewicza 21, 50-335 Wroc³aw, Po land
4 Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, Lower Silesian Branch, al. Jaworowa 19, 53-122, Wroc³aw, Po land
Alexandrowicz, W.A., Skoczylas, S., Sobczyk, S., Stefaniak, K., Kotowski, A., Przybylski, B., Ciszek, D., Badura, J., Urbañski, K., 2021. Mol lusc fau nas of lake de pos its in Gorzów Wielkopolski (NW Po land) as an in di ca tor of en vi ron men tal changes dur ing Eemian and Early Weichselian. Geo log i cal Quar terly, 2021, 65: 36, doi: 10.7306/gq.1605
As so ci ate Ed i tor: Wojciech Granoszewski
Dur ing the con struc tion of the S-3 road near Gorzów Wielkopolski, a sed i men tary suc ces sion of the Eemian Inter gla cial and the older part of the Weichselian Gla ci ation were ex posed. The suc ces sion, ~22 m thick, con sists of lac us trine and flu vio gla - cial de pos its. Lake sed i ments, mainly cal car e ous gyttja with peat in ter ca la tions, rep re sent the infills of two palaeolakes. The al most com plete skel e ton of a for est rhi noc eros, Stephanorhinus kirchbergensis, and a bone of the fal low deer Dama dama were found in the older lake de pos its. Mol lusc shells were nu mer ous in both lake se quences, anal y sis of which re vealed two types of as sem blage, rep re sent ing the coastal, lit to ral zone of a shal low lake with a muddy bot tom. The se quence of mol lusc com mu ni ties ob served in ver ti cal suc ces sion al lowed re con struc tion of en vi ron men tal changes dur ing de po si tion. Sev eral hy dro log i cal changes have been re cog nized within the palaeolake, es pe cially wa ter level fluc tu a tions prob a bly due to cli - mate change.
Key words: palaeolake, malacofauna, en vi ron men tal changes, Eemian Inter gla cial, Early Weichselian, NW Po land.
INTRODUCTION
Eemian de pos its are com mon in Eu rope, par tic u larly in the belt stretch ing from north ern France through the Ben elux coun - tries, north ern Ger many, Po land, to the Bal tic coun tries (e.g., Turner, 2000; Caspers et al., 2002; Behre et al., 2005; Henri - ksen et al., 2008; Helmens, 2013). In this zone, they are usu ally over lain by youn ger de pos its (Weichselian and/or Ho lo cene) and are rel a tively rarely ex posed to the sur face. Most com - monly the Eemian de pos its are of lake or marsh fa cies, with peat as the dom i nant li thol ogy, pro vid ing rich palynological ma - te rial en abling re con struc tion of cli ma tic changes (e.g., Zagwijn,
1996; Cheddadi et al., 1998; Kukla et al., 2002; Klotz et al., 2004; Müller et al., 2005; Brewer et al., 2008). Car bon ate de - pos its – lac us trine chalk and cal car e ous gyttja – are less com - mon. Due to the pres ence of cal cium car bon ate, they cre ate fa - vour able con di tions for the pres er va tion of molluscs. Such suc - ces sions have been de scribed, e.g. in Ger many (Meng et al., 2009a, b; Strahl et al., 2010; Menzel-Harloff and Meng, 2015;
Milano et al., 2020), Po land (Alexandrowicz and Alexandrowicz, 2010; Hrynowiecka et al., 2018), Belarus (Sanko et al., 2011) and Bal tic coun tries (Sanko and Gaigalas, 2007; Sanko et al., 2011).
In north ern Po land, Eemian de pos its are wide spread.
Among these, palaeolake infills are of great im por tance. Many such wa ter bod ies formed af ter the re treat of the Saalian ice sheet. Most of the lo cal i ties rec og nized lie south of the max i - mum range of the Weichselian Gla ci ation (Bruj and Ro man, 2007; Ro man, 2016). North of this line the num ber of lo cal i ties de scribed is much smaller (Urbañski and Win ter, 2005; Brose et al., 2006; Win ter et al., 2008; Niska and Miros³aw- Grabo - wska, 2015).
* Corresponding author, e-mail: wpalex@geol.agh.edu.pl Received: April 20, 2021; accepted: June 10, 2021; first published online: September 3, 2021
few lo cal i ties with Eemian de pos its have been rec og nized and de scribed so far. In ad di tion to lithological de scrip tions, palyno - logical, geo chem i cal and malacological stud ies have been per - formed at sev eral sites (e.g., Skompski, 1994; Urbañski and Win ter, 2005; Win ter et al., 2008; Alexandrowicz and Alexan - drowicz, 2010; Niska and Miros³aw-Grabowska, 2015).
The com po si tion and struc ture of mol lusc com mu ni ties closely re flect the en vi ron men tal con di tions pre vail ing at the time and place of sed i men ta tion. This makes molluscs a well- suited group for the needs of palaeoenvironmental and palaeo - climatic re con struc tions. Lac us trine molusc as sem blages usu - ally con tain ad mix tures of ter res trial spe cies. This cre ates a unique op por tu nity to rec re ate not only the con di tions in the lake it self, but also on its banks.
Malacological anal y ses of Eemian lac us trine de pos its in Po land have so far been re stricted to a rel a tively small group of sev eral dozen lo cal i ties dis persed across the Pol ish Low land (Skompski, 1996; Alexandrowicz and Alexandrowicz, 2010).
From this per spec tive, Eeemian mol lusc as sem blages are poorly un der stood com pared with those from Late Gla cial and Ho lo cene lo cal i ties (Alexandrowicz, 1999a, 2013), or even to those of the Holsteinian Inter gla cial (Skompski, 1996; Szyma - nek, 2012, 2013, 2014). Sim i larly, rel a tively small num bers of Eemian sites con tain ing fresh wa ter mol lusc com mu ni ties have so far been de scribed from other ar eas of the Eu ro pean Low - land (e.g., Sanko and Gaigalas, 2007; Meng et al., 2009a, b;
Strahl et al., 2010; Sanko et al., 2011; Menzel-Harloff and Meng, 2015; Milano et al., 2020).
This study re con structs the en vi ron men tal con di tions and fea tures of an Eemian palaeolake near Gorzów Wielkopolski.
From a re gional point of view, the data col lected will en rich knowl edge of subfossil com mu ni ties of molluscs in palaeolake
mental re con struc tions, es pe cially with re gard to hy dro log i cal and cli ma tic changes in central Eu rope dur ing the Eemian and Early Weichselian. In its com plete ness, the se quence found here is unique and only spo rad i cally ap pears in sites de scribed in the lit er a ture from the en tire area of the Eu ro pean Low land.
SITE DESCRIPTION
The inter gla cial lake-marsh de pos its were ex posed (Badura et al., 2017; Sobczyk et al., 2020) 6 km WSW from the cen tre of Gorzów Wielkopolski in a cut along the S-3 high way (GPS 51°43’N, 15°10’E; Fig. 1). This ar ti fi cial ex ca va tion cuts the mor pho log i cal edge sep a rat ing the mo raine pla teau of the Gorzów Plain (in the north) from the Toruñ-Eberswalde ice- mar ginal val ley (in the south) (Figs. 1 and 2). The re lief of the area is char ac ter ized by heights in the range of 70–90 m a.s.l.
for the mo raine pla teau, and ~18–20 m a.s.l. for the val ley that is cur rently oc cu pied by the Warta River. The south ern slopes of the mo raine pla teau are in many places cut by small val leys de - vel oped af ter the re treat of the Weichselian ice sheet (Fig. 2).
The de pos its ex posed in the cut form a se quence with a to - tal thick ness of ~22 m and were de scribed in de tail by Badura et al. (2017) and Sobczyk et al. (2020). They com prise two lake suc ces sions un der lain by sandy-gravel flu vio gla cial de pos its dated by OSL at 123.6 ±10.1 ka (Sobczyk et al., 2020). These rep re sent the fi nal phase of the Saalian Gla ci ation (MIS 6) (FG-1; Fig. 3). The low est part of the lake de pos its is of brown gyttja. Above, there is grey lac us trine chalk and gyttja with an ad mix ture of sand, bones of Stephanorhinus kirchbergensis and nu mer ous mol lusc shells. The rhi noc eros skel e ton was char ac ter ized by ex cep tional com plete ness (>130 bones).
Fig. 1. Lo ca tion of study area (map base: www.polska.e-mapa.net)
Apart from in di vid ual el e ments, only the right hind limb was miss ing. The bones were prob a bly quickly bur ied by sed i ment, as in di cated by the re mains of plant mat ter pre served in the tooth cav i ties (Sobczyk et al., 2020). The lower lake suc ces sion ends with a layer of peat mixed with sand, where the meta tar sal bone of a fal low deer (Dama dama) was found. The to tal thick - ness of the older lac us trine se quence is ~4.5 m (L-1; Fig. 3).
Palynological anal y ses of these de pos its in di cate that their de - po si tion took place dur ing the Eemian (MIS 5e) (de tailed de - scrip tion: Sobczyk et al., 2020). The lower lake suc ces sion is over lain by light brown lam i nated sand and silty sand ~2 m thick (delta-type de pos its), with an age es ti mated us ing the OSL method of 98.8 ±7.9 ka (MIS 5c; Sobczyk et al., 2020; FG-2;
Fig. 3). Above, there are de pos its be long ing to the up per lake suc ces sion, in clud ing grey chalk and car bon ate gyttja. This suc ces sion ends with peat over lain by a thin layer of gyttja and silty sand. This youn ger cal car e ous unit has a thick ness of
~4.5 m (L-2; Fig. 3) and cor re sponds to the Early Weichselian (MIS 5a–d). Such a strati graphic in ter pre ta tion is sup ported by the re sults of palynological anal y ses of these de pos its (see Sobczyk et al., 2020). Above the lake de pos its, there is an 11 m thick flu vio gla cial unit, its bot tom part rep re sented by 1.5 m of sands and grav els (OSL age 72.0 ±5.2 ka; Sobczyk et al., 2020) up wards giv ing way to gravel diamicton (FG-3; Fig. 3).
MATERIAL AND METHODS
Malacological anal y ses were per formed on 62 sam ples rep - re sent ing the en tire ex posed se quence. Shells were found within the lac us trine se quences (L-1 and L-2), though they were ab sent from the flu vio gla cial de pos its (FG-1, FG-2 and FG-3;
Fig. 3). The fi nal anal y sis was based on 25 sam ples. Each of these rep re sents an in ter val of 15 cm and had a mass of ~3 kg.
The mol lusc fauna in the up per part of the pro file was rel a tively poor, mak ing it nec es sary to com bine ad ja cent sam ples. Each of the three sam ples rep re sent ing this in ter val was made up by com bin ing three sam ples taken di rectly from ex po sure. The sam ples were mac er ated on a 0.5 mm sieve, and af ter dry ing, all com pletely pre served mol lusc shells (both adult and ju ve nile forms) and rec og niz able shell frag ments were picked out. The
shells are very well pre served. There are no signs of chem i cal dis so lu tion, and the small amount of frag ments in di cated a mi - nor role of me chan i cal comminution. Iden ti fi ca tion keys (Wel - ter-Schultes, 2012; Piechocki and Wawrzyniak-Wydrowska, 2016) and ref er ence col lec tions were used. Malacological anal - y sis was per formed us ing stan dard meth ods af ter Alexandro - wicz and Alexandrowicz (2011). In di vid ual spe cies were clas si - fied into eco log i cal groups:
M – mesophilous spe cies, H – hygrophilous spe cies,
WT – spe cies of tem po rary wa ter bod ies, WS – spe cies of per ma nent wa ter bod ies, WM – euryeco logical aquatic spe cies.
Per cent age shares of in di vid ual spe cies and eco log i cal groups were the ba sis for the con struc tion of the malacological di a gram show ing how mol lusc as sem blages vary through the se quence. Within the fauna ana lysed, it was pos si ble to dis tin - guish two types of com mu nity, each of spe cific com po si tion and with dis tinct eco log i cal re quire ments. The data col lected in this way were sup ported by the re sults of lithological, palynological and strati graphi cal stud ies con ducted at the site (Badura et al., 2017; Sobczyk et al., 2020), as well as malacological data from other Eemian lo cal i ties in Po land (e.g., Skompski, 1994, 1996;
Alexandrowicz and Alexandrowicz, 2010; Hrynowiecka et al., 2018) and in neigh bour ing coun tries (e.g., Sanko and Gaigalas, 2007; Meng et al., 2009a, b; Strahl et al., 2010; Sanko et al., 2011; Menzel-Harloff and Meng, 2015; Kenzler et al., 2018;
Milano et al., 2020).
RESULTS
The ma te rial pro cessed con tains a rel a tively rich, though not very di verse malacofauna. The num ber of taxa in in di vid ual sam ples var ied from 1 to 28, while the num ber of spec i mens ranged from 100 to 1672 (Fig. 3 and Ta ble 1). In to tal, all ma te - rial ana lysed in cluded 8641 spec i mens be long ing to 29 taxa (7 ter res trial taxa, 16 wa ter snails and 6 bi valves). The plates of slugs were iden ti fied un der the col lec tive name Limacidae.
There were also shell frag ments in the ma te rial ana lysed (not taken into ac count in the cal cu la tions; Ta ble 1).
Fig. 2. Map of of lo cal i ties with Eemian/Early Weichselian mol lusc-bear ing de pos its (map base: www.polska.e-mapa.net)
Ter res trial molluscs (eco log i cal groups M and H) played a mi nor role. Only taxa of open or par tially shaded, high hu mid ity hab i tats were pres ent. Succinea putris was the most nu mer ous ter res trial snail. It is a hygrophilous taxon in hab it ing open, pe ri - od i cally flooded biotopes, usu ally on the banks of wa ter bod ies.
A sim i lar type of hab i tat is pre ferred by other hygrophilous taxa iden ti fied in the fauna dis cussed, such as Pseudotrichia rubiginosa and Ver tigo antivertigo. Ter res trial spe cies were pres ent only in 8 sam ples. Their share in the as sem blages usu - ally did not ex ceed 10%, though in some sam ples, they were more nu mer ous, con sti tut ing up to 30% of the com mu nity (Fig.
4 and Ta ble 1).
Spe cies of tem po rary wa ter bod ies (eco log i cal group WT) were an im por tant com po nent of the fauna. They oc cur in al - most all sam ples, com pris ing up to 30% of the as sem blage (Fig. 4 and Ta ble 1). The most nu mer ous rep re sen ta tives of this group were Valvata cristata and Valvata macrostoma. Both forms pre fer shal low wa ter bod ies with a muddy bot tom, lush,
es pe cially sub merged veg e ta tion, and to a lesser ex tent reeds (kland, 1990; Piechocki and Wawrzyniak-Wydrowska, 2016).
They also tol er ate pe ri ods of dry ing.
Spe cies of per ma nent wa ter bod ies (eco log i cal group WS) formed the sec ond group of aquatic spe cies. This group was an im por tant com po nent in all sam ples (ex cept for sam ple 1), and its per cent age can reach 50% (Fig. 4 and Ta ble 1). A char ac ter - is tic rep re sen ta tive is Valvata piscinalis – a taxon typ i cal of per - ma nent lakes (and not re sis tant to dry ing out), es pe cially nu - mer ous in shal low, lit to ral zones (at a depth of 3 to 10 m). It has also been found at much greater depths (up to 80 m) (Wel - ter-Schultes, 2012; Piechocki and Wawrzyniak-Wydrowska 2016). Valvata piscinalis is a ben thic snail, liv ing ei ther di rectly on the bot tom or within sed i ments. It pre fers ox y gen-rich wa - ters, a muddy bot tom, and mod er ate plant veg e ta tion (kland, 1990; Piechocki and Wawrzyniak-Wydrowska, 2016). The other taxa in cluded in the WS group played a smaller role (Fig. 4 and Ta ble 1).
Fig. 3. Li thol ogy and sam pling of the Eemian/Early Weichselian mol lusc-bear ing de pos its at Gorzów Wielkopolski
NTAX num ber of molluscs spe cies, NIND – num ber of molluscs in di vid u als, FG-1, FG-3 – flu vio gla - cial de pos its (de scribed in text), FG-2 – delta-type de pos its (de scribed in text), L-1, L-2 – lake de - pos its (de scribed in text); li thol ogy af ter Badura et al. (2017) and Sobczyk et al. (2020)
Other spe cies were forms of high eco log i cal tol er ance that live in dif fer ent types of per ma nent wa ter body (eco log i cal group WM). The most im por tant is Bithynia tentaculata. It is a euryecological, widely-spread, palaearctic taxon liv ing in var i - ous types of wa ter body. In lakes, it pre fers the shal low, ox y - gen-rich part of the lit to ral zone (to a depth of 5 m, usu ally 0.5 to 2 m), and also sur vives short pe ri ods of dry ing (kland, 1990;
Wel ter-Schultes, 2012; Piechocki and Wawrzyniak- Wydro - wska, 2016). It is rarer in deeper zones (up to 20 m), and usu ally lives in zones rich in macrophytes, with a muddy bot tom. It is the most com mon snail spe cies in the en tire ma te rial (~35% of all in di vid u als iden ti fied). Bithynia tentaculata pro duces two hard skel e tal el e ments that pre serve well in sed i ments: the shell and the lid (operculum). Mor pho log i cally these two el e - ments dif fer sig nif i cantly from each other and re act dif fer ently to sed i men ta tion pro cesses. The shells have good buoy ancy and are eas ily dis placed by cur rents and/or waves, while the heavy lids quickly fall to the bot tom. In the ma te rial ana lysed, lids are al ways more nu mer ous than shells (Fig. 4 and Ta ble 1). The lack of chem i cal dis so lu tion of the shells and the rel a tively small num ber of bro ken shell frag ments in di cate that the dif fer ences in the rel a tive fre quency of shells and lids are the re sult of sed i - men ta tion sort ing by cur rents or waves, rather than of se lec tive de struc tion of skel e tal el e ments (chem i cal and/or me chan i cal).
Bithynia tentaculata is a fre quent com po nent of subfossil com - mu ni ties, par tic u larly those as so ci ated with warmer cli ma tic phases (e.g., Alexandrowicz, 1999a, b, 2013; Meng et al., 2009a, b; Menzel-Harloff and Meng, 2015; Milano et al., 2020).
Other spe cies be long ing to the WM eco log i cal group are of lesser im por tance (Fig. 4 and Ta ble 1).
The mol lusc fauna iden ti fied in the palaeolake de pos its of Gorzów Wielkopolski shows ver ti cal vari a tion and it is pos si ble to dis tin guish two types of as sem blage rep re sent ing dif fer ent types of hab i tat.
As sem blage A oc curred only in 4 sam ples (Fig. 4). It was char ac ter ized by the pres ence of ter res trial spe cies (reach ing up to 30% of the as sem blage): mesophilous, and es pe cially hygrophilous (eco log i cal groups M and H). Char ac ter is tic com - po nents of this com mu nity were Succinea putris, Pseudotrichia rubiginosa and Ver tigo antivertigo. The other im por tant group com prised spe cies typ i cal of tem po rary wa ter bod ies (eco log i - cal group WT), es pe cially Valvata macrostoma. Taxa of per ma - nent wa ter bod ies were less nu mer ous. As sem blage A rep re - sents the lake bank en vi ron ment. Spe cies com po si tion in di - cates the pres ence of wet, open hab i tats. Wa ter level fluc tu a - tions led to pe ri odic flood ing or dry ing of the coastal zone.
Hence, both hygrophilous ter res trial taxa and aquatic spe cies typ i cal of tem po rary wa ter bod ies are pres ent, while the share of lake taxa is rel a tively low. Fau nas with sim i lar char ac ter is tics have been de scribed from many lac us trine lo cal i ties rep re sent - ing both the Eemian (e.g., Meng, 2009a, b; Alexandrowicz and Alexandrowicz, 2010; Sanko et al., 2011; Menzel-Harloff and Meng, 2015) and the Ho lo cene (e.g., Alexandrowicz, 1999a, 2013).
As sem blage B was iden ti fied in 21 sam ples (Fig. 4). It is char ac ter ized by the dom i nance of aquatic taxa, es pe cially forms liv ing in per ma nent wa ter bod ies. The share of spe cies of tem po rary wa ter bod ies is rel a tively low and does not ex ceed 30%. Land snails are not pres ent or, at most, are an ac ces sory com po nent of the com mu nity, and their share does not ex ceed 5%. Two spe cies, Valvata piscinalis and Bithynia tentaculata, play the most im por tant role in com mu nity B. Both these forms are char ac ter ized by sig nif i cant eco log i cal tol er ance. The larger share of Valvata piscinalis may (though does not have to) im ply a slightly greater depth of the lakes, as well as a more lim ited
spread of aquatic plants. That re la tion ship is im plied by re sults of anal y ses of molluscs found in inter gla cial de pos its (e.g., Alexandrowicz, 1999a, 2013; Alexandrowicz and Alexandro - wicz, 2011; Szymanek, 2012, 2013, 2014; Hrynowiecka et al., 2018) and of the re cent fauna (Wel ter-Schultes, 2012; Piecho - cki and Wawrzyniak-Wydrowska, 2016).
DISCUSSION
LAKE EVOLUTION
The vari a tion in spe cies com po si tion and eco log i cal struc - ture of mol lusc as sem blages ob served in ver ti cal pro file in di - cates en vi ron men tal changes oc cur ring dur ing lac us trine de po - si tion. The base of the suc ces sion is formed of flu vio gla cial de - pos its rep re sent ing the fi nal phase of the Saalian Gla ci ation (MIS 6) and dated by OSL at 123.6 ±10.1 ka (Sobczyk et al., 2020). Within the older lake de pos its, the malacological se - quence is com plete and con tin u ous. Molluscs oc cur ring in the de pos its form ing the youn ger lake suc ces sion were much poorer. The Gorzów Wielkopolski de pos its ac cu mu lated in a shal low, nearshore part of the lit to ral zone, of a depth pos si bly not ex ceed ing sev eral metres. The lake was char ac ter ized by a muddy bot tom cov ered by macrophytes, nu mer ous re mains of Characeae, Najas ma rina and Najas flexilis be ing found in the de pos its (Sobczyk et al., 2020). There was no reed belt at the lake shore, as in di cated by the scarce pres ence of their re mains in the de pos its (Sobczyk et al., 2020) and by a lack of mol lusc spe cies char ac ter is tic of this type of hab i tat. Bithynia tentaculata opercula are more com mon the re spec tive shell re - flect ing cur rent sort ing given the lack of traces of chem i cal dis - so lu tion. Ac cord ing to palynological data (Sobczyk et al., 2020), the area sur round ing the lake was for est-cov ered. How ever, the mol lusc as sem blages not in clude any for est, or even shade-lov - ing spe cies, in di cat ing a pre dom i na tion of wet, or even pe ri od i - cally flooded open hab i tats. There fore, the lake shore was prob - a bly flat and cov ered by low, hygrophytic veg e ta tion. In these mor pho log i cal sit u a tions, the scale of shell ma te rial trans port is very lim ited. This phe nom e non has been de scribed for the pres ent al lu vial plains of river val leys and flat lake shores (Ilg et al., 2012; Èiliak et al., 2015). This lim i ta tion does not con cern pol len, which can be eas ily trans ported over larger dis tances.
Thus, it is most prob a ble that for est com mu ni ties re con structed on a ba sis of palynological anal y ses grew at a cer tain dis tance (maybe metres or tens of metres) from the lake shore.
The se quence of as sem blages ob served in di cates sig nif i - cant vari a tions in de po si tion con di tions, en ables the dis tinc tion of sev eral phases (Fig. 5).
Phase I (M-1, sam ples 1–4) rep re sents the floor de pos its of the older palaeolake. The type B as sem blage was iden ti fied here. Molluscs are rel a tively scarce, with Bithynia tentaculata play ing the most im por tant role. In the floor sam ple (sam ple 1), Bithynia tentaculata is the only spe cies iden ti fied. Up wards, its share grad u ally de creases. In the en tire in ter val dis cussed, opercula pre dom i nate over shells. The mol lusc fauna in di cates the shal low, nearshore part of the lit to ral zone, with rel a tively abun dant veg e ta tion and a muddy bot tom. The ob served in - crease in the share of Valvata piscinalis up wards may in di cate a slight deep en ing of the lake (Fig. 5).
Phase II (M-2; sam ples 5 and 6) is char ac ter ized by the pres ence of a dif fer ent com mu nity. Ter res trial spe cies, and those of tem po rary wa ter bod ies (as sem blage type A), play the most im por tant role. This fauna in ques tion is typ i cal of flat lake
shores and is char ac ter ized by wet, swampy, pe ri od i cally flooded ter res trial biotopes. The pe ri odic ap pear ance of in ter - ca la tions with ter res trial fauna within suc ces sions con tain ing lake mol lusc as sem blages has been widely de scribed through - out the Eu ro pean Low land (e.g., Alexandrowicz, 1999a, 2013;
Meng et al., 2009a, b; Fig. 5).
Phase III (M-3, sam ples 7–21) en com passes most of the older palaeolake suc ces sion. Spe cies typ i cal of a per ma nent wa ter body (type B as sem blage) pre dom i nate. The taxa rep re - sented are more di verse than those rep re sent ing Phase I.
Bithynia tentaculata and Valvata piscinalis are the most com - mon spe cies. Bithynia tentaculata is par tic u larly com mon at the
base (sam ples 7–13) and top (sam ples 18–21) of this in ter val.
In the mid dle part (sam ples 14–17) the share of Valvata pisci - nalis in creases no tice ably. In this part of the pro file, the mollu - scs are most nu mer ous and di verse. Fur ther more, a nearly com plete skel e ton of a rhi noc eros Stephanorhinus kirchber - gen sis was found here (Badura et al., 2017; Sobczyk et al., 2020). The in crease in the share of Valvata piscinalis in this part of this in ter val may in di cate a slight deep en ing of the lake. At the same time, the main tained pre dom i nance of opercula over shells of Bithynia tentaculata sug gests the pres ence of cur rents or the prox im ity of the wave zone (Fig. 5).
Com po si tion of the mol lusc fauna from the Eemian/Early Weichselian
E Taxon Sam ples
1 2 3 4 5 6 7 8 9 10 11
M
Limacidae Ver tigo substriata (Jeffr.)
Ver tigo angustior Jeffr.
H
Succinea putris (L.) 21 18
Ver tigo antivertigo (Drap.) Zonitoides nitidus (Müll.)
Pseudotrichia rubiginosa (Rossm.) 17 15
WT
Valvata cristata (Müll.) 8 14 21 19 12 1 5 6 7 3
Valvata macrostoma Mörch 17 18 22 26 6 3 13 9 12 6
Galba truncatula (Müll.) 5 1 1
Aplexa hypnorum (L.)
Planorbis planorbis (L.) 3 9 1 5 1 4
Segmentina nitida Müll.
Pisidium obtusale (Lam.) 1 7 8 5 1
WS
Valvata piscinalis (Müll.) 43 69 73 54 21 32 18 18 16 14
Acroloxus lacustris (L.)
Stagnicola palustris (Müll.) 2 2
Lymnaea stagnalis (L.) Physa fontinalis (L.)
Anisus contortus (L.) 9 3 1 1
Gyraulus albus (Müll.) Pisidium nitidum Jen.
Pisidium milium Held
WM
Bithynia tentaculata (L.) (in clud ing operculum)
112 84
37 22
26 22
27 17
44 29
18 10
121 76
74 55
64 40
71 54
85 60 Ra dix peregra (Müll.)
Gyraulus crista (L.) 4 8 1 3
Pisidium casertanum Poli Pisidium henslowanum (Shep.)
Pisidium subtruncatum Malm
To tal spe cies 1 6 7 9 11 11 5 6 8 4 4
To tal in di vid u als 112 109 157 168 198 100 157 114 103 106 108
In de ter mi nate shell frag ments 68 47 31 56 54 18 21 33 23 20 15
Ter res trial snails 6 2
Bithynia cf. tentaculata 34 6 4 9 8 1 5 8 6 5 4
Valvata cf. piscinalis 9 12 5 15 10 2 3 6 4 2 2
Valvata sp. 9 7 7 13 11 3 4 6 5 4 3
Other snails frag ments 14 19 12 18 16 10 8 10 7 7 5
Pisidium sp. 2 3 3 1 3 1 1 3 1 2 1
Eco log i cal group of molluscs (af ter: Alexandrowicz and Alexandrowicz, 2011): M – mesophilous spe cies, H – of per ma nent wa ter bod ies, WM – euryecological aquatic spe cies
Phase IV (M-4; sam ple 22) rep re sents the top of the older lake suc ces sion. The molluscs here in clude rel a tively nu mer - ous ter res trial spe cies (as sem blage type A), while the taxa of pe ri odic wa ter bod ies also play a greater role. The ob served change in the spe cies com po si tion and eco log i cal struc ture of the fauna in di cates a coastal, pe ri od i cally flooded part of the lake and wet land. At the same time, there is a change in the na - ture of the sed i ments: gyttja is re placed by peat (Fig. 5).
Phases I–IV, rep re sent ing the infill of the youn ger palaeo - lake, pos si bly cor re spond to the warm pe riod of the Eemian Inter gla cial. This in ter pre ta tion is sup ported by the re sults of lithological and palynological anal y ses per formed on that in ter -
val (Sobczyk et al., 2020). The com po si tion and struc ture of the mol lus can com mu ni ties, with the pre dom i nance of euryecolo - gical aquatic taxa (Valvata piscinalis and Bithynia tentaculata), do not pro vide firm ev i dence for strati graphi cal con clu sions, only al low ing char ac ter iza tion of changes in eco log i cal con di - tions in the lake and on its shores.
The ab sence of mol lusc shells in the sands (delta-type de - pos its) sep a rat ing the lake suc ces sions is as so ci ated with the very low con tent of car bon ates in these de pos its (Sobczyk et al., 2020). OSL data (98.8 ±7.9 ka) in di cates that these de pos its ac cu mu lated in the ini tial stage of the last gla ci ation, prob a bly dur ing stage MIS5c (Sobczyk et al., 2020).
T a b l e 1 de pos its of Gorzów Wielkopolski
12 13 14 15 16 17 18 19 20 21 22 23 24 25
7 8 5
21 2 10 45 15
17 3 38 19
1 29 12 47 89 34
28 11 12 49 15
18 1 15 38 10
19 5 12 68 4
9 1 29 122 76 87 1 3 4 8 112 10 8 31
7 2 45 158 59 98 7 6 8 14 147 8 14 44
1 39 19 36 1 19 3
8 8 14
36 1 21 1 1
2 6 17 2
4 4 29 28 17 5 1 39 7 5 4
19 14 144 509 269 444 19 9 19 21 487 19 23 24
7 3
10 21 2
9 1 2
3 6 19 1
2 29 10 39 1 7 3 32 9 1
5 19 1 2 2 17 2
29 44 24 4 55 7
33 31 28 5
72 44
86 57
66 41
348 279
190 121
535 421
87 63
79 56
40 28
45 38
285 221
59 47
41 31
93 69
3 2 3 12
8 17 19 22 35 3 4 9 58 8 4 12
8 47 15 39 4 7 10 1 1 2
12 12 17 10
18 21
8 4 9 28 17 28 6 6 11 10 25 7 10 22
122 103 326 1607 811 1585 119 104 102 108 1672 112 113 325
32 16 57 159 120 232 18 18 13 21 168 14 23 59
1 24 16 21 19 7
8 5 8 17 12 17 6 5 2 5 22 3 5 10
6 2 12 19 8 31 3 2 1 3 17 1 3 8
6 2 11 39 27 44 1 1 1 4 34 2 3 12
10 6 18 49 38 86 6 4 7 6 62 5 8 16
2 1 7 11 19 33 2 3 2 3 14 3 4 6
higrophilous spe cies, WT – aquatic spe cies of tem po rary wa ter bod ies, WS – aquatic spe cies
The mol lusc as sem blages pres ent in the youn ger lake suc - ces sion is poorer and al lows less in ter pre ta tion. Sam ples 23 and 24 (Phase V, M-5; Fig. 5) are dom i nated by taxa of per ma - nent wa ter bod ies, in di cat ing a rel a tively shal low, lit to ral zone of the lake. The top of the malacological se quence (Phase VI, M-6, sam ple 25; Fig. 5) rep re sents sig nif i cant en vi ron men tal change. The share of ter res trial spe cies in creases sig nif i cantly, reach ing 25% of the com mu nity. Molluscs of pe ri odic wa ter bod ies also play a greater role. The M-6 phase, there fore, rep - re sents the lake shore. The youn ger lake suc ces sion may well be as so ci ated with the early stage of the last gla cial (prob a bly MIS5b and/or MIS5a; Sobczyk et al., 2020). The re sults of palaeobotanical anal y ses in di cate a cool ing of the cli mate dur - ing the de po si tion of these sed i ments (Klotz et al., 2004; Behre et al., 2006; Helmens, 2013), though the molluscs do not show this trend, with a lack of cold-lov ing spe cies. Early Weichselian lo cal i ties in the Eu ro pean Low land are rel a tively few, and only ex cep tional ones con tain a malacological re cord (Sanko and Gaigalas, 2007; Strahl et al., 2010; Kenzler et al., 2018; Hryno - wiecka et al., 2018). From this per spec tive, the Gorzów Wielko - polski pro file should be con sid ered unique in Eu rope.
REGIONAL IMPORTANCE OF THE MALACOFAUNA
The pro file of lake de pos its re vealed at Gorzów Wielko - polski is an ex cep tion ally com plete malacological se quence cor re spond ing to two cli ma tic stages:
–warmer (Eemian – MIS 5e);
–cooler (Early Weichselian – MIS 5b or MIS 5a).
The malacofauna is char ac ter ized by the dom i nance of aquatic spe cies, while in creases in the share of ter res trial forms is vis i ble only in short sec tions of the se quence. The mol lusc com mu nity found in the older part of the lake se quence does not con tain cold-lov ing spe cies and un doubt edly in di cates the inter gla cial age of the sed i ments. On the other hand, no Belgrandia marginata was found here. This thermophilic spe - cies is in dic a tive of Eemian river and lake de pos its that ac cu mu - lated in the warm est (op ti mal) stage of the inter gla cial. It has been re corded at this strati graphi cal po si tion at many sites through out Eu rope (Sanko and Gaigalas, 2007; Alexandrowicz and Alexandrowicz, 2010; Sanko et al., 2011; Menzel-Harloff and Meng, 2015; Kenzler et al., 2018). The re sults of geo chem i - Fig. 4. Eco log i cal com po si tion of molluscs from the Eemian and Early Weichselian lake
de pos its of Gorzów Wielkopolski For ex pla na tions see Ta ble 1
cal and palynological anal y ses car ried out within the older lake se quence in di cate that it rep re sents a com plete pro file (Badura et al., 2017; Sobczyk et al., 2020). The ab sence of Belgrandia marginata may re sult from the ex is tence of a strati graphi cal gap cov er ing the inter gla cial op ti mum (which is not sup ported by the re sults of other stud ies, es pe cially palynological ones; Sobczyk et al., 2020). It seems more likely that this spe cies did not in - habit the lake and, con se quently, its shells did not ap pear here.
The changes in fau nal com mu ni ties ob served in ver ti cal pro file in di cate fluc tu a tions in the wa ter level in the lake, prob a - bly re lated to cli mate changes, es pe cially with the ap pear ance of colder and warmer phases, as well as drier and wet ter ones (e.g., Cheddadi et al., 1998; Kukla et al., 2002; Koltz et al., 2004; Müller et al., 2005; Brewer et al., 2008). The pres ence of al ter nat ing stages of high and low wa ter lev els in lakes has been thor oughly stud ied for the pe riod of the Late Gla cial and the Ho lo cene (e.g., Goslar et al., 1993; Pun ning et al., 2005;
Ga³ka et al., 2015). Sim i lar anal y ses with re gard to the Eemian Inter gla cial are much less ad vanced.
In the se quence dis cussed, it is pos si ble to dis tin guish two pe ri ods of low wa ter level in the lake. The older one oc curs at
the bot tom of the se quence (phase M-2) and may be as so ci - ated with the end of the Early Eemian. It cor re sponds to the phase dis tin guished in this pro file that is char ac ter ized by a sig - nif i cant share of Corylus pol len (Co-Qe-Al phase; Sobczyk et al., 2020). A de crease in the wa ter level in lakes at that time is re corded at sev eral sites across the Eu ro pean Low land (Kupryja nowicz, 2008; Helmens, 2013), which in di cates the re - gional na ture of this phe nom e non. The later pe riod rep re sents the dis ap pear ance of the lake in the de clin ing stage of the inter - gla cial (phase M-4) and is re corded in many other lake pro files of the Eemian Inter gla cial (e.g., Kupryjanowicz, 2008; Miro - s³aw- Grabowska and G¹siorowski, 2010; Paw³owski, 2011;
Helmens, 2013; Niska, 2015; Hrynowiecka et al., 2018; Salo - nen et al., 2018). In both phases, there is a mol lusc as sem blage with a sig nif i cant share of ter res trial spe cies (as sem blage type A) in di cat ing a lake shore (Fig. 5). A fauna with aquatic spe cies (as sem blage type B) in di cates the lit to ral zone, with a muddy bot tom and more or less rich plant veg e ta tion. This in di cates the rel a tively shal low depth of the lake, though also its sta bil ity (phases: M-1 and M-3; Fig. 5). A tem po rary in crease in the Valvata piscinalis share ob served in phases M-1 and M-3 may Fig. 5. Evo lu tion and en vi ron men tal changes of Eemian/Early Weichselian lakes
near Gorzów Wielkopolski
A, B – mol lus can as sem blages (de scribed in text), M-1–M-6 – phases of lake de vel op ment (de scribed in text), ? – pos si ble higher wa ter level in the lake – in creased share
of Valvata piscinalis
tions in the Gorzów Wielkopolski palaeolakes, re con structed based on changes in mol lusc as sem blages, show sig nif i cant con ver gence with the re con struc tion of hy dro log i cal con di tions car ried out by means of palynological anal y sis and Cladocera stud ies (e.g., Kupryjanowicz, 2008; Miros³aw-Grabowska and G¹siorowski, 2010; Paw³owski, 2011; Helmens, 2013; Niska, 2015; Salonen et al., 2018; Sinopoli et al., 2019), as well as with the con clu sions of palaeoclimatic and iso tope stud ies (Miro - s³aw- Grabowska and G¹siorowski, 2010; Milano et al., 2020).
This site is the first Eemian sed i men tary pro file in Eu rope where an at tempt has been made to re con struct wa ter level fluc tu a - tions us ing mol lusc com mu ni ties. The phases of high wa ter level re con structed in the Gorzów Wielkopolski palaeolake cor - re late with the ma rine trans gres sions de scribed from the Lower Vistula Ba sin (Makowska, 1986).
Sim i larly, com plete se quences of lake de pos its with fresh - wa ter molluscs rep re sent ing an Eemian Inter gla cial are known only from a few sites across the Eu ro pean Low land (Sanko and Gaigalas, 2007; Meng et al., 2009a, b; Alexandrowicz and Alexandrowicz, 2010; Strahl et al., 2010; Sanko et al., 2011;
Menzel-Harloff and Meng, 2015; Kenzler et al., 2018; Milano et al., 2020; Fig. 6 and Ta ble 2). These malacological se quences show many sim i lar i ties, with sim i lar spe cies com po si tions (usu - ally ~20–30 spe cies of aquatic molluscs, and sev eral ter res trial taxa). As at Gorzów Wielkopolski, the pre dom i nant spe cies are most of ten Bithynia tentaculata and Valvata piscinalis. The in - creased share of ter res trial fauna in some pro files is due to the pres ence of cold-lov ing taxa within the gla cial de pos its that un -
found at Gorzów Wielkopolski. There are also no cryophilic spe - cies here, which are pres ent at most other sites. There are gaps within the malacological se quences in all pro files, but typ i cally they cover short seg ments, es pe cially in the in ter vals cor re - spond ing to MIS 5e. De pos its rep re sent ing the end of the Saalian Gla ci ation (MIS 6) are pres ent in only one site, while the suc ces sion cor re spond ing to the Early Weichselian (MIS 5a-d) is usu ally in com plete (Ta ble 2). A com par i son of se lected malacological sites from across the Eu ro pean Low land (Ta ble 2) shows that the Gorzów Wielkopolski pro file may be in cluded among the key malacological pro files of the Eemian Inter gla cial in this part of Eu rope. At these sites, molluscs of sim i lar spe cies com po si tion and eco log i cal struc ture have been iden ti fied, the dom i nant spe cies usu ally be ing Bithynia tentaculata and Valvata piscinalis. These ob ser va tions may sug gest con sid er - able sta bil ity of en vi ron men tal con di tions (cli ma tic, hy dro log i cal) in this re gion. Sim i lar con clu sions are prompted by re gional palynological (e.g., Cheddadi et al., 1998; Behre et al., 2005;
Sinopoli et al., 2019) and cli ma tic (e.g., Kukla et al., 2002;
Brewer et al., 2008; Salonen et al., 2018) re con struc tions. On the other hand, each of these stud ies shows the in flu ence of lo - cal fac tors mod i fy ing re gional trends to a greater or lesser ex - tent. The wa ter level fluc tu a tions ob served at Gorzów Wielko - polski cor re late with cli ma tic phases re con structed on the ba sis of palaeobotanical stud ies (e.g., Zagwin, 1996; Kupryjanowicz, 2008; Helmens, 2013). There fore, it may be in ferred that they are a re cord of re gional cli mate trends.
T a b l e 2 Com par i son of se lected Eemian malacological pro files from the Eu ro pean Low land
Lo ca tion
Gorzów Wielkopolski
(this ar ti cle) (GW)
Pi³a Alexandrowicz
and Alexandrowicz
(2010) (PI)
Peene Val ley Meng et al., (2009a) (PV)
Klein Klütz Höved Menzel-Harloff
and Meng (2015) (KH)
Neumark Nord Strahl et al.
(2010) (NN)
Petrovshchina Sanko et al.
(2011) (PE)
Rumlovka Sanko et al.
(2011) (RU)
Netiesos Sanko and
Gaigalas (2007) (NE) NW
Po land
NW Po land
NE Ger many
NW Ger many
Mid dle
Ger many Belarus Belarus Lith u a nia
Type of sed i ments
lac us trine lit to ral zone
lac us trine lit to ral zone
flu vial/lac us - trine low land river/
lit to ral zone
lac us trine lit to ral zone
lac us trine lit to ral zone
lac us trine lit to ral zone
lac us trine lit to ral zone
lac us trine lit to ral zone Malacofauna
Num ber of spe cies W – wa ter, L – land
29 22(W), 7(L)
21 17(W), 4(L)
29 29(W), 0(L)
43 30(W), 13(L)
25 14(W), 11(T)
24 22(W), 2(L)
39 28(W), 11(L)
33 25(W), 8(L)
Most com mon
spe cies
Bithynia tentaculata;
Valvata piscinalis
Bithynia tentaculata;
Valvata piscinalis
Bithynia tentaculata;
Valvata piscinalis
Bithynia tentaculata
Bithynia tentaculata,
Gyraulus laevis
Valvata piscinalis
Pisidium moitessieria-n
um, Valvata piscinalis
Bithynia tentaculata,
Valvata piscinalis Oc cur rence
of Belgrandia
marginata
lack pres ent lack pres ent lack pres ent pres ent pres ent
Com ments to malacofauna
cold-lov ing, land snails in
MIS 6
cold-lov ing, land snails in MIS 5d
cold-lov ing, land snails in
MIS 4
cold-lov ing, land snails in
MIS 5d Stra tig ra phy
Stra tig ra phy range
MIS 5e;
MIS 5b or/and MIS5a
MIS 5e MIS 5e MIS 6 (up per
part); MIS 5e MIS 5e, d MIS 5e MIS 5e;
MIS 4 MIS 5e, d
Stratigra- phical gaps
lack:
MIS 5d, c
some small gaps dur ing MIS 5e
some gaps dur ing MIS
5e
some small gaps dur ing MIS 5e
some small gaps dur ing
MIS 5e
prob a bly lack of the low er - most part of
MIS 5e
lack: MIS 5d-a
prob a bly lack of the low er - most part of
MIS 5e
CONCLUSIONS
The malacological se quence de scribed from Gorzów Wielkopolski is dis con tin u ous. It cov ers the Eemian Inter gla cial and the older part of the Weichselian Gla ci ation and can be cor - re lated with the iso tope stages MIS 5e (older lake), MIS 5b and/or MIS 5a (youn ger lake) (Sobczyk et al., 2020). While mol - lusc com mu ni ties have been re ported from Eemian palaeo - lakes at many sites through out Eu rope (Sanko and Gaigalas, 2007; Meng et al., 2009a, b; Alexandrowicz and Alexandrowicz, 2010; Sanko et al., 2011; Menzel-Harloff and Meng, 2015;
Kenzler et al., 2018; Milano et al., 2020), Weichselian early phase molluscs are known from only a few sites, mainly within loess (Moine et al., 2005; Alexandrowicz and Dmytruk, 2007;
Sümegi et al., 2016). The mol lus can as sem blages rec og nized have en abled the re con struc tion of the fea tures of a lake and its banks. The mol lusc com mu ni ties in di cate the shal low lit to ral zone of the lake with a muddy bot tom cov ered by less or more abun dant sub merged veg e ta tion, though lacked a reed belt along the shore. The lake was lo cated in a for est area, while the lake shore was flat, wet and cov ered by low, hygrophytic veg e - ta tion. This kind of shore zone made pos si ble pe ri odic flood ing or dry ing, as a con se quence of rel a tively mi nor changes in lake level, and these ep i sodes were re flected in the changes in mol - lus can com mu nity com po si tion and struc ture.
Ver ti cal vari a tion in the fau nal as sem blages in di cates en vi - ron men tal changes in the sed i men ta tion zone. Sev eral fluc tu a -
tions in wa ter level likely re flect cli mate fluc tu a tions. A clear re la - tion ship be tween cli mate change and fluc tu a tions in lake level has been rec og nized for the Late Gla cial and the Ho lo cene (e.g., Goslar et al., 1993; Pun ning et al., 2005; Ga³ka et al., 2015). For the Eemian, such stud ies are fewer. The pro file de - scribed is so far the only one in Cen tral Eu rope where such fluc - tu a tions have been iden ti fied and de fined based on mol lusc as - sem blages. Malacological anal y sis has here al lowed the rec og - ni tion of two pe ri ods of low wa ter in the lake: the older one at the end of the early inter gla cial stage and the youn ger one cor re - spond ing to the pe riod of the lake’s dis ap pear ance at the end of the inter gla cial.
The con clu sions drawn in this study very closely match the re sults ob tained us ing other meth ods (Badura et al., 2017;
Sobczyk et al., 2020). The re sults as a whole, re vealed dur ing ex ca va tion of the S-3 motor way near Gorzów Wielkopolski, in - di cate that this site is one of the most valu able ex po sures of Eemian Inter gla cial de pos its on the Eu ro pean Low land.
Ac knowl edge ments. This re search was sup ported by the grant 0201/2048/18 “Life and death of ex tinct rhino (Stephano - rhinus sp.) from West ern Po land: a multiproxy palaeoenviron - mental ap proach” fi nanced from the funds of the Na tional Sci - ence Cen tre, Po land. We would also like to thank P. Sümegi and M. Szymanek for their con struc tive com ments.
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