Geo log i cal Quar terly, 2016, 60 (1): 225–237 DOI: http://dx.doi.org/10.7306/gq.1271
In sta bil ity of the en vi ron ment at the end of the Eemian Inter gla cial as il lus trated by isopollen maps of Po land
Miros³awa KUPRYJANOWICZ1, *, Wojciech GRANOSZEWSKI2, Dorota NALEPKA3, Irena Agnieszka PIDEK4, Adam WALANUS5, Zofia BALWIERZ6, Magdalena FI£OC1, Piotr KO£ACZEK7, Aleksandra MAJECKA8, Ma³gorzata MALKIEWICZ9, Ma³gorzata NITA10,
Bo¿ena NORYŒKIEWICZ11 and Hanna WIN TER12
1 Uni ver sity of Bia³ystok, De part ment of Bot any, In sti tute of Bi ol ogy, Cio³kowskiego 1J, 15-425 Bia³ystok, Po land
2 Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, Carpathian Branch, Skrzatów 1, 31-560 Kraków, Po land
3 Pol ish Acad emy of Sci ences, W³adys³aw Szafer In sti tute of Bot any, Lubicz 46, 31-512 Kraków, Po land
4 Maria Sk³odowska-Cu rie Uni ver sity, De part ment of Geoecology and Palaeo ge ogra phy, Fac ulty of Earth Sci ences and Spa tial Man age ment, Kraœnicka 2cd, 20-718 Lublin, Po land
5 AGH Uni ver sity of Sci ence and Tech nol ogy, Fac ulty of Ge ol ogy, Geo phys ics and En vi ron men tal Pro tec tion, al. A. Mickiewicza 30, 30-059 Kraków, Po land
6 Uni ver sity of £ódŸ, De part ment of Geo mor phol ogy, In sti tute of Earth Sci ences, Narutowicza 88, 90-139 £ódŸ, Po land
7 Adam Mickiewicz Uni ver sity, De part ment of Biogeography and Palaeo ec ol ogy, In sti tute of Geoecology and Geoinformation, Dziêgielowa 27, 61-680 Poznañ, Po land
8 Uni ver sity of War saw, De part ment of Cli mate Ge ol ogy, In sti tute of Ge ol ogy, ¯wirki i Wigury 93, 02-089 Warszawa, Po land
9 Uni ver sity of Wroc³aw, Lab o ra tory of Palaeobotany, De part ment of Strati graphi cal Ge ol ogy, In sti tute of Geo log i cal Sci - ences, Cybulskiego 30, 50-205 Wroc³aw, Po land
10 Uni ver sity of Silesia, De part ment of Fun da men tal Ge ol ogy, Fac ulty of Earth Sci ences, Bêdziñska 60, 41-200 Sosnowiec, Po land
11 Nicolaus Co per ni cus Uni ver sity, De part ment of Geo mor phol ogy and Palaeo ge ogra phy of the Qua ter nary, Fac ulty of Earth Sci ences, Gagarina 9, 87-100 Toruñ, Po land
12 Pol ish Geo log i cal In sti tute – Na tional Re search In sti tute, Rakowicka 4, 00-975 Warszawa, Po land
Kupryjanowicz, M., Granoszewski, W., Nalepka, D., Pidek, I.A., Walanus, A., Balwierz, Z., Fi³oc, M., Ko³aczek P., Majecka, A., Malkiewicz, M., Nita, M., Noryœkiewicz, M., Win ter, H, H., 2016. In sta bil ity of the en vi ron ment at the end of the Eemian Inter gla cial as il lus trated by isopollen maps of Po land. Geo log i cal Quar terly, 60 (1): 225–237, doi: 10.7306/gq.1271 Many ter res trial pol len pro files from Po land (and a few pol len re cords from other parts of Cen tral Eu rope) show the end of the last inter gla cial (Eemian, MIS 5e) to have been char ac ter ized by cli ma tic and en vi ron men tal in sta bil ity. This is ex pressed by a strong, rapid cool ing in the mid dle part of the pine phase end ing this inter gla cial (E7 re gional pol len as sem blage zone), and then a re-warm ing at the very end of this phase, im me di ately be fore the tran si tion to the gla cial con di tions of the last gla ci - ation (Vistulian, Weichselian, MIS 5d). We have char ac ter ized the re gional dis tri bu tion of these cli ma tic fluc tu a tions in Po - land on the ba sis of isopollen maps pre pared for the Eemian Inter gla cial based on palynological data from 31 Pol ish pol len pro files. These maps show un equiv o cally that the intra-inter gla cial cool ing at the end of the Eemian Inter gla cial was a transregional phe nom e non, which was re flected very clearly by a tem po rary open ness of veg e ta tion across the whole of Po - land. It was as so ci ated with a dis tinct de crease in pine for est ar eas and an in crease in birch for ests and open com mu ni ties of cold steppe type with a dom i na tion of Ar te mi sia. The pro nounced cli mate and en vi ron ment in sta bil ity dur ing the last phase of the Eemian Inter gla cial may be con sis tent with it be ing a nat u ral phe nom e non, char ac ter is tic of tran si tional stages. Tak ing into con sid er ation the cur rently ob served global warm ing, co in cid ing with a nat u ral cool ing trend, the study of such tran si - tional stages is im por tant for un der stand ing the un der ly ing pro cesses of cli mate change.
Key words: palaeoclimate, intra-inter gla cial cool ing, veg e ta tion his tory, isopolles, pol len anal y sis.
* Corresponding author, e-mail: m.kupryjanowicz@uwb.edu.pl Received: October 20, 2015; accepted: December 7, 2015; first published online: January 22, 2016
INTRODUCTION
The ques tion of en vi ron men tal sta bil ity dur ing the Eemian Inter gla cial has long been dis cussed (Velichko et al., 1982, 2005; Fronval and Jansen, 1996; Litt et al., 1996; Kukla et al., 1997, 2002; Seidenkrantz and Knudsen, 1997; Saarnisto et al., 1999; Guiter et al., 2003; Klotz et al., 2003, 2004; Kühl and Litt, 2003; Seppä et al., 2008, and oth ers). The dis cus sions wid - ened when a re cord of strong tem per a ture fluc tu a tions dur ing the Eemian Inter gla cial was dis cov ered in ice cores in Green - land (Anklin et al., 1993; Dansgaard et al., 1993; Grootes et al., 1993; GRIP, 1993). The re sults of the North Green land Ice Core Pro ject (2004) re vealed, as one of the most im por tant fac - tors, abrupt cli mate warm ing about 115,000 years ago to wards the end of the last inter gla cial, be fore gla cial con di tions were fully de vel oped. The pres ence of strong fluc tu a tions in cli mate in the fi nal phase of the Eemian Inter gla cial was sup ported by pol len data from some Eu ro pean sites. These data come mostly from the long maar se quences in South ern Eu rope (e.g., Tzedakis et al., 1994). In Cen tral and East ern Eu rope such data are scarce (Müller, 1974; Beaulieu and Reille, 1989, 1992a, b;
Mamakowa, 1989; Guiot et al., 1993; Field et al., 1994;
Thouveny et al., 1994; Cheddadi et al., 1998; Granoszewski, 2003; Borisova et al., 2007; Kupryjanowicz, 2008; Boettger et al., 2009). In the Eemian se quence from Gröbern in Ger many, the sta ble iso tope data in di cated a pro nounced warm ing phase to wards the very end of the inter gla cial, just be fore the tran si tion to the gla cial ep och (Boettger et al., 2009). A sim i lar cli ma tic os - cil la tion is re corded in a few other pro files from Ger many (e.g., Neumark-Nord – Eissmann, 2002, and Klinge – Striegler, 1986;
Velichko et al., 2005; Novenko et al., 2008a) and some pro files from Rus sia (e.g., Ples – Borisova et al., 2007). Boettger et al.
(2009) com piled pro files from Cen tral and East ern Eu rope, in - clud ing the pol len re cord of cli mate in sta bil ity at the end of the Eemian Inter gla cial. In this list, how ever, data from Pol ish pro - files are miss ing. Our re view of Pol ish palynological data on the Eemian Inter gla cial dem on strated that at least a dozen such sites ex isted in Po land (Fig. 1). In pol len re cord from these sites the abrupt cool ing was rec og nized by us in the mid dle part of the E7 Pinus zone, which was the fi nal phase of the last inter - gla cial. This event is re corded there as a tem po rary de crease in pine fre quency and a spread ing of birch, and then also of open veg e ta tion (Fig. 2). At the ma jor ity of these sites this cool ing was not dis tin guished by the au thors of the re ports. De spite that, the palynological re cord is, in our opin ion, un am big u ous and fully jus ti fies this dis tinc tion.
Our aim was to look into dis tri bu tion of the pol len re cords of the Late Eemian cli ma tic fluc tu a tions in Po land. The best tech - nique for such a re gional syn the sis of palynological data is the isopollen method. Isopolles are syn chro nous lines lim it ing ar - eas with iden ti cal per cent age shares of pol len from a spe cific plant taxon, link ing points (sites), where the same share of pol - len was re corded at a given time. The isopollen method was in - tro duced by Szafer (1935) in his work on the postglacial his tory of veg e ta tion in Po land. Later, this syn thetic method for the pre - sen ta tion of pol len data was suc cess fully used and de vel oped in dif fer ent coun tries (Sauremo, 1940; Firbas, 1949; Bertsch, 1953; Donner, 1963; Moe, 1970; Birks et al., 1975; Birks and Saarnisto, 1975; Da vis and Webb, 1975; Da vis, 1976; Bernabo and Webb, 1977; Hunt ley and Birks, 1983; Ralska-Jasiewi - czowa, 1983; Webb et al., 1987). Re cently, the role of the isopollen method has be come even more sig nif i cant with the de vel op ment of da ta bases, ded i cated soft ware and nu mer i cal
meth ods (e.g., Ralska-Jasiewiczowa et al., 2004; Obidowicz et al., 2013; Wacnik et al., 2015).
The main ob jec tive of our in ves ti ga tion is to con trib ute to a better un der stand ing of cli ma tic vari abil ity dur ing the ter mi nal phase of the last inter gla cial by study ing pol len pro files from Po - land. It al lows us to in fer that the rec og ni tion of the cli mate changes in the de clin ing part of the Eemian Inter gla cial en ables the pre dic tion of anal o gous nat u ral changes that should have oc curred in the late Ho lo cene, and their sep a ra tion from anthropogenic changes.
MATERIAL AND METHODS
PALYNOSTRATIGRAPHY OF THE LATE EEMIAN IN POLAND
Due to the lack of de ter mi na tion of the ab so lute age of the Eemian lac us trine and mire de pos its, the ho ri zons for which we drew isopollen maps are not time ho ri zons, but they are biostratigraphical ho ri zons, i.e. pol len ho ri zons. The pe riod un - der con sid er ation in cludes the E7 Pinus re gional pol len zone de ter mined for Po land by Mamakowa (1989). It is the last (youn gest, up per most) pol len as sem blage zone of the Eemian Inter gla cial. In this zone we dis tin guished five re gional subzones (Fig. 2 and Ta ble 1) based on fea tures pro posed by Kupryjanowicz (2008), who was the first to draw at ten tion to the cold inter gla cial cli ma tic os cil la tions in these Eemian pol len re - cords. Isopollen maps for se lected pol len taxa of trees (Pinus sylvestris t., Betula and Picea abies t.), shrubs (Juniperus communis), dwarf shrubs (Calluna) and the sum of her ba ceous plants was pre pared for each of these subzones. These plants were the main com po nents of veg e ta tion in the part ana lysed of the Eemian Inter gla cial. To dem on strate the dif fer ence in the pat tern of isopolles (and thus in the re gional di ver sity of veg e ta - tion) be tween a cold event within the pine zone and the first stadial of the Early Vistulian (Weichselian), we also cre ated an isopollen map for the re gional pol len zone rep re sent ing this stadial (EV1).
SITES
Based both on Mamakowa’s stud ies (1989, 2003) and on fur ther orig i nal work from 2003-2014 we cre ated the pol len da - ta base, which in cludes data from 184 sites where a com plete pol len re cord for at least one zone of the Eemian Inter gla cial was reg is tered (Kupryjanowicz et al., in print). The pol len re cord of the pine zone (E7 R PAZ) was rec og nized for as many as 112 of these sites. How ever, sec tions of pro files con tain ing this re - cord were stud ied very var i ously. Most of them are of low res o - lu tion, and there fore they do not con tain a re cord of a cold event within the pine zone. This event was rec og nized at 31 sites (Fig. 3). Its lo ca tion is shown both in Fig ure 1 and on isopollen maps pre sented in this pa per (Fig. 4).
CONSTRUCTION OF ISOPOLLEN MAPS
The data re quired for isopollen maps of the Eemian Inter - gla cial are stored in the POLPAL sys tem (Walanus and Nalepka, 1999; Nalepka and Walanus, 2003; see also web page POLPAL). Most of the pol len ta bles that make up the Eemian da ta base con tain the raw data, i.e. pol len counts. Sev - eral ta bles con tain per cent ages, be cause they were avail able only in this form. More than a dozen ta bles con tain raw data,
Instability of the environment at the end of the Eemian Interglacial as illustrated by isopollen maps of Poland 227
Fig. 1. Location of sites studied (sites with numbers in black squares; the numbers of sites according to the Eemian isopollen data base) on the background of other Polish localities of the Eemian Interglacial from isopollen data base
(sites with numbers in grey)
35 – Horoszki Du¿e (Granoszewski, 2003), 37 – Babiec Piaseczny (Krupiñski, 1995; Kotarbiñski and Krupiñski, 1995), 42 – D¹brówki (Krupiñski, 2005), 54 – Studzieniec (Krupiñski, 2005), 59 – Warszawa-Wawrzyszew (Krupiñski and Morawski, 1993); 67 – Choroszczewo (Kupryjanowicz, 2008), 70 – Dzierniakowo (Kupryjanowicz, 2008), 75 – Hieronimowo (Kupryjanowicz, 2008); 80 – Machnacz (Kupryjanowicz, 1991, 1994), 83 – Micha³owo (Kupryjanowicz and Drzymulska, 2002), 93 – Solniki (Kupryjanowicz, 2008), 102 – Mikorzyn Starszy (Stankowski et al., 1999), 113 – Grudzielec (Malkiewicz, 2002), 124 – Wiœniew II (Pidek and Terpi³owski, 1993, 1995a, b), 142 – Rzecino (Win ter et al., 2008), 148 – Szwajcaria 1 (Borówko-D³u¿akowa and Halicki, 1957), 154 – Konopki Leœne (Borówko-D³u¿akowa and Halicki, 1957), 172 – Otapy I (Bitner, 1956), 173 – Otapy II (Bitner, 1956), 180 – Imbramowice (Mamakowa, 1989), 207 – Dziadowa K³oda (Kuszell, 1998), 229 – Krzepczów 4 (Klatkowa, 1972), 239 – Skaratki (Chmielewski, 1961), 246 – Zgierz-Rudunki I (Jastrzêbska-Mame³ka, 1985), 247 – Zgierz-Rudunki II (Jastrzêbska-Mame³ka, 1985), 262 – Kletnia Stara (Janczyk-Kopikowa, 1986, 1987; ¯arski, 1989), 272 – Warszawa-Wola (Borówko-D³u¿akowa, 1960), 284 – Ruszkówek (Janczyk-Kopikowa, 1997), 300 – Szwajcaria (Ber et al., 1998; Krupiñski, 1998); 307 – Zieluñ ZN.1.00 (Krupiñski, 2005); 308 – Mikorzyn 1 (Stankowski et al., 1999, 2003; Stankowski and Nita, 2004)
Fig. 2. Simplified pollen diagram from the Solniki profile illustrating changes of vegetation during the E7 Pinus regional pollen zone of the Eemian interglacial
(according to Kupryjanowicz, 2008 – modified)
R PAZ – regional pollen assemblage zones, R PASZ – regional pollen assemblage subzones (for characteristics see Table 1), grey rectangle indicates subzones representing the
intra-Eemian cooling
R PAZ Sym bol and name
R PASZ Sym bol and name
De scrip tion of re gional pol len zones or subzones
De scrip tion of pol len ho ri zon used for
draw ing of maps
EV1 Poaceae–Ar te mi - sia–Betula nana
High NAP val ues very great va ri ety of taxa of her ba ceous plants; max i mum of Ar te mi sia, which is the most dis tinc tive pol len taxon of
the zone; in creas ing pol len val ues of Juniperus, Betula nana t. and Salix; pres ence
of sin gle pol len grains of thermophilous trees (redeposition).
NAP max i mum
E7 Pinus
E7e Pinus–Betula Cul mi na tion of Pinus sylvestris t.; de cline of
NAP and Betula alba t. Peak of Pinus
E7d NAP Cul mi na tion of NAP val ues; still high pro por - tion of Betula alba t.; rel a tively low fre quency
of Pinus sylvestris t. Peak of NAP
E7c Betula
Cul mi na tion of Betula alba t. pol len; rise in NAP val ues; de pres sion of Pinus sylvestris t.;
fi nal dis ap pear ance of thermophilous trees and shrubs.
Peak of Betula
E7b Pinus Max i mum of Pinus sylvestris t. pol len; de - crease of Picea abies t. and thermophilous
trees and shrubs.
Pinus max i mum
E7a Pinus–Picea
Sys tem atic rise in Pinus sylvestris t. pol len;
rel a tively high, but grad u ally de creas ing pro - por tion of Picea abies t., reg u lar oc cur rence of Carpinus betulus and Alnus pol len, but
with low val ues.
Mid dle pol len spec trum
Grey shad ing in di cates subzones rep re sent ing the intra-Eemian cool ing
T a b l e 1 Re gional palynostratigraphy of the youn gest, pine phase of the Eemian inter gla cial (E7 R PAZ) and the old est part of the Early
Vistulian (EV1 R PAZ) in Po land and pol len ho ri zons rep re sen ta tive of these units; re gional pol len as sem blage zones (R PAZ) ac cord ing to Mamakowa (1989)
which were cre ated by dig i tiz ing di a grams avail able in the lit er a - ture or in other ar chives. These ta bles con tain the data only for se lected pol len spec tra.
The pol len ho ri zons for which isopollen maps were con - structed are iden ti fied with only in di vid ual pol len spec tra, se - lected within par tic u lar pol len zones or subzones (Ta ble 1).
Hence, these Eemian pol len ho ri zons are not equiv a lents to the pol len as sem blage zones (PAZ), or pol len as sem blage subzones (PASZ) as de fined by Birks (1986).
The no men cla ture of pol len taxa was standardized for the ta bles sub mit ted to the da ta base. The cal cu la tion of per cent - ages is based on the pol len sum of trees, shrubs, dwarf shrubs and ter res trial herbs. The set of isoline val ues for trees and shrubs con sists of 19 per cent age ranges (0.01, 0.1, 0.5, 1, 2, 3, 5, 7, 10, 15, 20, 25, 30, 35, 40, 50, 60, 70 and 80%), and for her - ba ceous taxa 14 ranges (0.01, 0.1, 0.2, 0.5, 0.7, 1, 2, 3, 5, 7, 10, 20, 30 and 40%).
Palynological data were used to con struct isopollen maps of the Eemian Inter gla cial for 18 dif fer ent rank taxa of trees and shrubs, for 6 taxa of dwarf shrubs and herbs, and for the sum of non-arborescent plants (NAP, herbs). This pa per pres ents maps for sev eral se lected taxa which best il lus trate changes in veg e ta tion as so ci ated with the late Eemian cli ma tic os cil la tions.
A com plete se ries of maps for the en tire Eemian Inter gla cial, and a de tailed de scrip tion of the method used, can be found in a mono graph by Kupryjanowicz et al. (in print).
RESULTS
E7A SUBZONE
The pine zone (E7 R PAZ) rep re sents the lat est stage of the Eemian suc ces sion of veg e ta tion. Pro gres sive hu mid ity and cool ing of the cli mate, char ac ter is tic of the telocratic stage of the
inter gla cial cy cle, re sulted in a change of hab i tat con di tions and im posed a trans for ma tion of plant com mu ni ties from the very be gin ning of this zone. Pine was the main for est-form ing tree at that time across Po land. This is re flected in the isopollen maps by very high pine pol len val ues, which in most ar eas reach 60–70%, and only in the south-west are slightly lower, 50–60%
(Fig. 4). Pine for est was per haps like the mod ern Leucobryo-Pinetum (Granoszewski, 2003). This is a tall pine for est within the range of oce anic cli mate in cen tral and east ern Cen tral Eu rope (Matuszkiewicz, 2006), in which birch and spruce lo cally con sti tute an ad mix ture. An isopollen map shows that the share of birch was greater in cen tral, south ern and east ern Po land (gen er ally 5–7%, lo cally to 10%), while in the west and north its im por tance at this time was rel a tively smaller (3–5%). The pol len val ues of Picea, rang ing from 10–15% in the whole of Po land, and in the north-east even to15–20% sug - gest that it was fairly abun dant, and that it played a sig nif i cant role in the com mu ni ties of the whole area un der con sid er ation.
Spruce may have oc curred mainly in the al ready de grad ing wet alderwoods, and in riverine for ests. Also its role in the ex pand - ing pine for ests may have been prom i nent, es pe cially in bog pinewoods. In the low est ar eas per haps even spruce-pine bo - real for ests oc curred, es pe cially in north east ern Po land.
Due to the spread of pine for ests, in which there is lit tle shade, the sig nif i cance of open com mu ni ties in creased fur ther.
This is il lus trated on isopollen maps by a high pro por tion of herbs (to 7–10% in south west ern and north east ern Po land).
The great est di ver sity of her ba ceous plant taxa was as so ci ated with moist and boggy hab i tats, which dom i nated at that time in the land scape of many re gions of Po land (e.g., Mamakowa, 1989; Granoszewski, 2003; Krupiñski, 2005; Kupryjanowicz, 2008). The in creas ing role of these hab i tats is shown by an in - crease in the pol len of Poaceae and Cyperaceae. The grasses were pos si bly as so ci ated mainly with the boggy form of pine for - est at that time (e.g., Granoszewski, 2003). Drier light for est Instability of the environment at the end of the Eemian Interglacial as illustrated by isopollen maps of Poland 229
Fig. 3. Regional pollen assemblage subzones of the pine (E7) zone of the Eemian Interglacial and the first pollen zone (EV1) of the Early Vistulian (Weichselian) represented in particular studied profiles (the numbers
of sites according to the Eemian isopollen data base; for locations of sites see Fig. 1) Grey shading indicates subzones representing the intra-Eemian cooling
Fig. 4. Isopollen maps of selected pollen taxa illustrating the main vegetation changes beginning of the last glaciations Blue frame indicates subzones representing the intra-Eemian cooling;
Instability of the environment at the end of the Eemian Interglacial as illustrated by isopollen maps of Poland 231
dur ing the fi nal phase of the Eemian Inter gla cial (E7 Pinus R PAZ) and at the very (EV1 Poaceae–Ar te mi sia–Betula nana R PAZ) in Po land
for better qual ity of this fig ure see Ap pen dix 1 in the on line ver sion, at doi: 10.7306/gq.1271
hab i tats were per haps oc cu pied mainly by patches of heath, as is in di cated by the in crease in the pol len val ues of Calluna vulgaris, es pe cially in north west ern, east ern and east ern cen - tral Po land.
In view of the changes tak ing place in the plant com po si tion, the con stant oc cur rence at that time of Hedera he lix pol len, and the pres ence of Viscum al bum and Ilex aquifolium pol len is a prob lem that is hard to in ter pret (e.g., Mamakowa, 1989).
These pol len grains are most prob a bly con nected with redeposition. How ever, Hedera and Viscum may have per - sisted in eco log i cally fa vour able hab i tats.
E7B SUBZONE
Pine reached its max i mal inter gla cial spread in this subzone. In the whole of Po land, its pol len val ues in creased above 70%, and in the north-west and north-east even to 90%.
It formed thick, al most pure for ests, prob a bly de vel oped as dif - fer ent com mu ni ties. They oc cu pied nearly all hab i tats. Apart from pine, which was the main for est tree, birch trees also grew there. Their role in north ern and south west ern Po land was slightly greater than in the pre vi ous subzone, as sug gested by an in creased per cent age share of Betula in these re gions (gen - er ally to 7%, and lo cally even to 10%), as shown in the isopollen map; only in the north-west it was still un der 5%.
The ex pan sion of pine and spread of birch were re lated to the de cline of spruce and of all other for est trees. How ever, the rel a tively high val ues of Picea abies t. pol len (gen er ally 3–5%;
2–3% only lo cally in northeast ern and northwest ern Po land) in - di cate that spruce still played a sub stan tial role in for est com - mu ni ties, and prob a bly ac com pa nied pine, form ing pine-spruce for ests. In sparse, well-lit co nif er ous for ests the sig nif i cance of open com mu ni ties con se quently in creased. This is in di cated by a fur ther rise in the per cent age amount of herbs, which for most Pol ish ter ri tory reach up to 7%. Drier for est hab i tats were still oc cu pied by heath, as in di cated by the in crease in the pol len fre quency of Calluna vulgaris to 0.7% in the whole of Po land.
E7C AND E7D SUBZONES – INTRA-INTERGLACIAL COOLING
The E7c and E7d subzones of the pine zone rep re sent a cold cli ma tic os cil la tion. The re cord of that os cil la tion is in cluded in many pro files stud ied from dif fer ent re gions of Po land. The cool ing re cord is the best de vel oped in the pro file from Solniki in north ern Podlasie (Kupryjanowicz, 2008). At this site, thanks to the high res o lu tion of the pol len anal y sis, the two-step course of that os cil la tion was rec og nized for the first time (Fig. 2). The isopollen maps showed that a sim i lar pat tern of this os cil la tion is also re corded in sev eral other pro files (Fig. 4). How ever, zones rep re sent ing this intra-Eemian cool ing are usu ally con tained within a thin in ter val, and they may eas ily be omit ted in stud ies based on ran dom sam ples. This ex plains their pres ence only at some sites, which strongly hin ders the re con struc tion of the re - gional dif fer ences in veg e ta tion changes.
This prob a bly brief cold pe riod was marked by the de cline of pine and a strong in crease in the pro por tion of birch. It is doc u - mented on the isopollen maps of the E7c subzone by a fall in Pinus pol len val ues gen er ally <70%, in the north-east <50%, and in the south-west even <35%, as well as by a rise in Betula pol len pro por tion to gen er ally 20%, and in the north-east and south-west to 50%. The area of the pine for ests was sub stan - tially re duced. They were tem po rary re placed by birch for ests.
Ac cord ing to Granoszewski (2003), at that time birch may have lo cally formed stands of the Betuletum pubescentis type, in boggy for mer hab i tats of al der carr com mu ni ties, acid i fied by
spruce. Pine and spruce also grew there. De spite the tem po rary in crease in the sig nif i cance of birch, it seems that pine-spruce for ests were still pres ent. How ever, the role of spruce di min - ished sig nif i cantly across the whole of Po land, which is re flected in de creased per cent ages of its pol len to 1–2%, in di cated in isopollen maps. Only in north east ern and cen tral Po land the role of spruce was slightly more im por tant (3–5% and 2–3%, re - spec tively).
These changes were ad di tion ally ac com pa nied by an in - crease in the pro por tion of herbs (usu ally to 20%), in clud ing mainly Cyperaceae, Poaceae, Ar te mi sia and nu mer ous oth ers (Mamakowa, 1989; Granoszewski, 2003; Krupiñski, 2005;
Kupryjanowicz, 2008). As a re sult of cli mate wors en ing and the deg ra da tion of soils, the for est prob a bly re treated from the hab i - tats where edaphic and hy dro log i cal con di tions were less fa - vour able. These hab i tats were oc cu pied by com mu ni ties of her - ba ceous plants, prob a bly both on drier soils and on wet ter ones.
In north west ern Po land heathlands dom i nated by Calluna vulgaris (in crease in pol len val ues to 3%) and other mem bers of the Ericaceae fam ily (rise also to 3% – Win ter et al., 2008) were de vel oped at that time. In cen tral Po land Juniperus ap peared in greater amounts (0.5–1%).
The E7d subzone rep re sents the sec ond stage of a cold cli - ma tic os cil la tion within the pine phase. The pol len re cord sug - gests a fur ther cool ing of the cli mate. It led to the re duc tion of birch for ests, which were formed in the pre vi ous subzone (de - cline of Betula pol len val ues to 10%, and lo cally in the north-west even to 7%), and to the spread of the open com mu - ni ties (in crease in the pro por tion of herbs to 30%, in the north-east to 40%, and in the south-west to 50%). Moist hab i - tats were mainly oc cu pied by shrub tun dra with shrub wil low (e.g., Kupryjanowicz, 2008). In drier hab i tats, open steppe grass com mu ni ties spread, with high pro por tions of Ar te mi sia and Chenopodiaceae (e.g., Granoszewski, 2003). A de crease in for est cover may also be sug gested by the in crease of the role of Juniperus, which had al ready ap peared in the pre vi ous subzone, and in this zone its pol len reached val ues of 1%
across al most all of Po land, ex cept the south-west and north-east. Juniperus spread across north west ern and north - east ern Po land (its pol len val ues reach 2% in these re gions).
It seems highly likely that the E7c and E7d subzones re flect some sud den cli mate change to wards both cool ing and wet - ness. The oc cur rence of a short-lived cold os cil la tion is also in - di cated by ev i dence of sur face run off in the de pos its at sev eral sites (e.g., Machnacz – Kupryjanowicz, 1994; Dzierniakowo – Kupryjanowicz, 2008) in the form of sand and gravel lenses.
E7E SUBZONE
Af ter the cold pe riod, the cli mate be came slightly warmer.
Pine re turned to places oc cu pied for a time by birch and open veg e ta tion – this is il lus trated in the isopollen maps by a rise in Pinus sylvestris t. pol len across Po land (usu ally to 80%, and in the east and north-west to 90%) and by a fall in Betula (most marked in north ern and east ern parts of the coun try, <10%; in other re gions to 10–20%), herbs (gen er ally to 10–20%, and in the east and north-west to 7–10%) and Calluna (gen er ally
<0.2%; only in the north-west pol len val ues re mained sim i lar to those in the pre vi ous pe riod, 1–2%). Pine for ests spread anew, but in a struc ture dif fer ent to that of the E7a and E7b subzones.
They were sparser, with a clearly lower pro por tion of spruce, with out fir and thermophilous de cid u ous trees, which were re - placed by birch. The area of open com mu ni ties was sig nif i cantly smaller than in the pre vi ous subzone.
Changes dur ing the E7e subzone can be ex plained by a tem po rary ame lio ra tion in the cli mate at the very end of the Eemian Inter gla cial (Kupryjanowicz, 2008). How ever, the gen - eral ten dency of the veg e ta tion suc ces sion dur ing the E7d subzone would point to con tin u ous de te ri o ra tion of the cli mate start ing di rectly af ter the short warm ing at the be gin ning of this subzone. In the Horoszki pro file this is sup ported by a de crease in or ganic mat ter, a de creas ing trend in the con cen tra tion of sporomorphs, and by an ac com pa ny ing in crease in the pro por - tion of herbs (Granoszewski, 2003).
The low amounts of thermophilous tree pol len prob a bly re - flect long-dis tance trans port. The pine for est was bo real in na - ture, and in the youn gest part of the E7e subzone even had fea - tures of for est tun dra, per haps sim i lar to the mod ern com mu ni - ties of this type (Norin, 1961), and was clearly in re treat.
EV1 ZONE
An in ten si fi ca tion in cli mate change to wards cool ing fi nally brought about the thin ning and re treat of for est com mu ni ties from ar eas of Po land and the spread of open plant com mu ni - ties. This pro cess is re flected on the isopollen maps as a very marked in crease in her ba ceous plant pol len (above 40% in whole Po land, ex cept in its northwest ern part where the share of herbs was rel a tively low, 30–40%, but the pro por tion of Calluna sig nif i cantly in creased, lo cally even to 20%), and the rapid fall of pol len val ues of all trees (de cline of Pinus sylvestris t. pol len val ues to 25–30%, Betula to 15–20%, and Picea abies t. <2% across al most the whole coun try), which marks the bound ary be tween the inter gla cial and gla cial suc - ces sion of veg e ta tion, i.e. be tween the telocratic and cryocratic stages of the gla cial/inter gla cial cy cle.
These changes are ac com pa nied by a suc ces sive in crease in the pro por tion of coarser sand frac tions in the de pos its at nu - mer ous sites, symp tom atic of the re-in ten si fi ca tion of ero sional pro cesses (e.g., Mamakowa, 1989; Krupiñski, 2005;
Kupryjanowicz, 2008). The rise in the pol len val ues of Alnus, Corylus, Tilia, Quercus, and Ulmus in di cates redeposition of inter gla cial sed i ments in the layer in which the pro por tion of coarser sand frac tions of the sed i ments reaches a max i mum.
This zone rep re sents the first post-Eemian cold pe riod of stadial char ac ter. The vegetational land scape at that time was tun dra with clus ters of wil lows and prob a bly shrub birches. A fur ther in crease in the her ba ceous plant pol len value, up to 50%, pro vides ev i dence of the con tin u ously pro gress ing ex pan - sion of var i ous open com mu ni ties. The dis tinct in crease in the pol len val ues of Ar te mi sia and Poaceae sug gests the ris ing im - por tance of grass com mu ni ties, per haps steppe-like in na ture.
Shrub com mu ni ties, the ap pear ance of which is emphasised by the oc cur rence of pol len of Juniperus, gained in sig nif i cance. Among the tree spe cies, only pine and birch might have oc curred, form ing small patches in open ar eas.
DISCUSSION
The re cord of an intra-Eemian cold os cil la tion in the fi nal, pine phase of the last inter gla cial (E7 R PAZ) is noted in 31 pro - files from Po land. In all these pro files the sec tions be long ing to the Pinus zone are made up of ho mo ge neous un dis turbed lac - us trine de pos its (e.g., Mamakowa, 1989; Kupryjanowicz, 1994, 2008; Kupryjanowicz and Drzymulska, 2002; Granoszewski, 2003; Krupiñski, 2005). There fore, the cold os cil la tion de - scribed can not re sult from sed i men tary re work ing. The re cord of the cool ing is the best de vel oped in the pro file from the
Solniki site in north east ern Po land. In the Solniki pol len pro file, thanks to the high res o lu tion of pol len anal y sis, a two-step course of this cool ing is vis i ble (Kupryjanowicz, 2008). The re - sults of Cladocera, mag netic sus cep ti bil ity and sta ble iso tope anal y ses from these same pro files sup port this course of cli - mate change in the re gion (Kupryjanowicz et al., 2005;
Miros³aw-Grabowska et al., 2015). Sim i lar ad di tional anal y ses were car ried also at sev eral other sites ex am ined paly no logi cal - ly, and ev ery where they were con sis tent with a tem po rary drop in tem per a ture within the pine phase of the Eemian Inter gla cial (e.g., Rzecino – Niska and Miros³aw-Grabowska, 2015;
Studzieniec – Miros³aw-Grabowska and Niska, 2007).
The re treat of pine dur ing the E7c and E7d subzones is sig - nalled on the isopollen maps by a sig nif i cant fall in Pinus pol len val ues at all sites stud ied. At the same time there is a rise in the amount of Betula pol len, as can be clearly seen on the isopollen maps of this taxon. Such a change may re flect a cool ing of the cli mate or an in crease in its hu mid ity. Sim i lar os cil la tions, ex - pressed in the palynological re cord by a de crease of per cent - age val ues of Pinus pol len and an in crease in the Betula pol len pro por tion, took place also in the early Ho lo cene dur ing the Preboreal pe riod when pine for est dom i nated, and are rel a tively well doc u mented (e.g., Allen et al., 2002; Feurdean et al., 2008;
PÀlachs et al., 2011; Wan ner et al., 2011).
Subzone E7b rep re sents a pe riod of the max i mal inter gla - cial spread of pine for ests. Spruce grad u ally lost its po si tion in for ests. Thermophilous trees were al most com pletely ab sent.
All these facts sug gest rel a tively cold cli ma tic con di tions with a con ti nen tal char ac ter. How ever, the mean July tem per a ture was not lower than +12°C (Granoszewski, 2003; Kupryja - nowicz, 2008). The mean Jan u ary tem per a ture de creased be - low –5°C. A multi-method ap proach by Brewer at al. (2008) also showed a de crease in tem per a tures and an in crease in dry con - di tions af ter 120 ka BP. In subzone E7c, the es tab lished Pinus for est was in vaded by Betula. A cool ing of the cli mate seems to be the most prob a ble rea son for this change. The mean July tem per a ture prob a bly de creased to about +12°C at that time (Kupryjanowicz, 2008). In subzone E7d, the sig nif i cant re duc - tion in ar eas of bo real pine-birch for ests, and the spread of her - ba ceous com mu ni ties dom i nated by Ar te mi sia, took place.
These changes point to a rel a tively dry con ti nen tal cli mate, with the mean Jan u ary tem per a ture <–10°C, and the mean July tem per a ture only slightly higher than +10°C (Kupryjanowicz, 2008). In subzone E7e, the new ex pan sion of pine for est and a de cline of birch and herbs un doubt edly re flect a slightly more tem per ate cli mate. The mean July tem per a ture once again had to be >+12°C, and the mean Jan u ary tem per a ture >–5°C (Granoszewski, 2003; Kupryjanowicz, 2008).
Beaulieu and Reille (1989) have al ready de scribed a sim i lar cold os cil la tion in the late part of the Eemian Inter gla cial in Les Echets, south ern France. The au thors cor re late it with sim i lar os cil la tions re corded in some pol len pro files from Ger many (Odderade – Averdieck, 1967; Sulzberg-Baden – Welten, 1981; Gondiswill – Wegmüller, 1986), France (La Grande Pile – Woillard, 1978) and The Neth er lands (BrÝrup – Andersen, 1961). This os cil la tion is also dis tinctly vis i ble in the re con struc - tion of mean Jan u ary and July tem per a tures in Bispingen, north ern Ger many (Kühl and Litt, 2003).
The fact that the cold os cil la tion in the Eemian pine zone is not re corded in nu mer ous pro files from al most the whole of Eu - rope may re sult from dif fer ent rea sons. In cases when the thick - ness of sed i ments, in which the fi nal part of the Eemian Inter - gla cial is re corded, is low, the os cil la tion may not be no ticed with the stan dard dis tance be tween ana lysed pol len sam ples (ev ery 5–10 cm). Ac cord ing to Beaulieu and Reille (1989), it is also pos si ble that this short and low-am pli tude os cil la tion in - Instability of the environment at the end of the Eemian Interglacial as illustrated by isopollen maps of Poland 233
duced no ta ble veg e ta tion changes only near ecotones in the north ern cen tral and east ern Eu rope.
A cor re la tion of intra-Eemian cli ma tic os cil la tions con tained in the ter res trial re cords from Eu ro pean (in clud ing Pol ish) sites with a cold event ob served in Green land ice cores or deep ma - rine cores is very dif fi cult. In the great ma jor ity of cases, the lack of ab so lute age de ter mi na tions of those changes is the first, and un doubt edly the most im por tant rea son for the dif fi cul ties.
In pro files that were not sub jected to pol len anal y sis and ab so - lute dat ing, there is no cer tainty that they con tain a re cord of the whole inter gla cial. In con se quence of that, there is an in abil ity to de fine in which part of the inter gla cial the re corded change took place.
At the end of the last inter gla cial, the pol len and geo chem i - cal data for a few se quences in Cen tral and East ern Eu rope show a grad ual cool ing dur ing the inter gla cial/gla cial tran si tion.
This can be cor re lated with the con tin u ous tem per a ture fall at ca. 120 ka BP in the NGRIP ice core from cen tral Green land (North Green land Ice Core Pro ject, 2004). Within this global cli - ma tic tran si tion are clear in di ca tions of in sta bil i ties of cli mate and veg e ta tion dy nam ics, pos si bly caused through brief cool ing and then warm ing to wards the very end of the last inter gla cial in Cen tral Eu ro pean pro files. More over, Menke and Tynni (1984) re ferred to en vi ron men tal os cil la tions in pro files from north west - ern Ger many. Strong cli mate fluc tu a tions ac com pa nied by in - ten sive loess de po si tion dur ing the Late Eemian arid ity pulse, and an abrupt cool ing event ex actly at the time of the Last Gla - cial in cep tion at ca. 118 ka BP, were also noted by Sirocko et al.
(2005), and Seelos and Sirocko (2007) in north ern and west ern Ger many. Pos si bly, these re sults cor re spond to the sea level rise in ferred world wide in the fi nal stage of the last inter gla cial (Hearty et al., 2007). In East ern Eu rope, slight warm ing os cil la - tions oc curred dur ing the very end of the last inter gla cial in the pro file from Ples (Up per Volga re gion), and these also were ear lier noted in sev eral other pro files in the north-west of the Rus sian Plain and Lith u a nia (e.g., Grichuk, 1961; Kondratene, 1996; Novenko et al., 2008b).
The cor re la tion of the cli ma tic fluc tu a tion de tected be tween the con ti nen tal limnic se quences in Po land and also in the whole of Cen tral and East ern Eu rope is well-es tab lished – it def i nitely oc curred dur ing E7 R PAZ, i.e. be fore the be gin ning of the first Early Last Gla cial stadial (Herning). Prob a bly, the cli - mate be fore the inter gla cial/gla cial tran si tion was char ac ter ized by in ter nal in sta bil ity, re sult ing in the se quence of cli ma tic os cil - la tions ob served, within the over all trend to wards cool ing. The pre cise cor re la tion be tween the pro files in ves ti gated and the pat terns ob served in the NGRIP ice core re mains un der dis cus - sion. There are some prob lems with the cor re la tion of DO 25 in NGRIP with ter res trial re cords. Johnsen et al. (2005) re ported that the warm ing event DO 25 lasted some 4 ka. Such a pe riod seems to be too long for cor re la tion with the warm ing event at the very end of the last inter gla cial in ter res trial re cords, al - though the ex act du ra tion of the ob served warm ing events in lake pro files pre sented here re mains un clear.
For the Eemian Inter gla cial, as for the Ho lo cene, af ter sig nif - i cant warm ing dur ing the tran si tion from the gla cial to inter gla - cial and reach ing the cli ma tic op ti mum, sta ble warm cli ma tic con di tions set in with a grad ual tem per a ture de crease with time.
It ap pears that in Cen tral Eu rope a short cool ing and warm ing phase at the very end of the last inter gla cial pre ceded the fi nal tran si tion to gla cial con di tions. This is part of the ob served in - crease in cli ma tic and en vi ron men tal in sta bil ity dur ing this global warm/cold tran si tion, which ap pears to be at least a Eu ro - pean phe nom e non. The ob served cli ma tic and en vi ron men tal in sta bil ity dur ing the fi nal part of the last inter gla cial could be se - ri ously con sid ered as a pos si ble gen eral nat u ral char ac ter is tic of global warm/cold tran si tion pe ri ods. There fore, de tailed stud - ies of these phe nom ena are im por tant for eval u at ing and un der - stand ing the cur rently ob served and very con tro ver sially dis - cussed (e.g., Velichko et al., 2005; Keenlyside et al., 2008) cli - ma tic trends, and for de tect ing anthropogenic in put in cli mate change.
SUMMARY AND CONCLUSIONS
In 31 pol len pro files from dif fer ent re gions of Po land, clearly de fined cli ma tic fluc tu a tions are re corded by pol len data at the end of the Eemian Inter gla cial. They con tain a strong cool ing in the mid dle part of the pine phase of the inter gla cial (subzones E7c and E7d of the E7 Pinus re gional pol len zone), and sub se - quent warm ing at the very end of this phase (E7e subzone).
The pan-re gional char ac ter of the Late Eemian cold event is clearly no tice able on isopollen maps as a sig nif i cant re duc tion in Pinus pol len through out the whole Pol ish ter ri tory. The drop in tem per a ture was at first ac com pa nied by an in creased pro por - tion of Betula pol len (E7c subzone), and then also an in crease in the pro por tion of pol len of non-ar bo real plants (E7d subzone). The warm ing at the very end of the Eemian Inter gla - cial (E7e subzone) is il lus trated on isopollen maps by the re-in - crease of Pinus pol len in the whole ter ri tory of Po land and by a de cline of Betula and her ba ceous pol len. This fluc tu a tion of cli - mate was not just a lo cal or re gional phe nom e non, but seems to have been an event on the scale of the whole of West ern, Cen - tral and East ern Eu rope.
Ac knowl edge ments. We thank Ch. Turner and anon y - mous re viewer for help ful com ments on ear lier drafts. The in - ves ti ga tions have been sup ported by the Pol ish Min is try of Sci - ence and Higher Ed u ca tion (pro ject no. N N304 352238) and by the stat u tory funds of the Uni ver sity of Bia³ystok, Pol ish Geo log - i cal In sti tute – Na tional Re search In sti tute, W. Szafer In sti tute of Bot any PAS, Maria Sk³odowska-Cu rie Uni ver sity, AGH Uni ver - sity of Sci ence and Tech nol ogy, Adam Mickiewicz Uni ver sity, Uni ver sity of War saw, Uni ver sity of Wroc³aw, and Uni ver sity of Silesia.
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