Studia Quaternaria, vol. 31, no. 1 (2014): 51–60. DOI: 10.2478/squa-2014-0005

LATE HOLO CENE CHANGES IN VEGE TA TION OF THE MR¥GOWO LAKELAND (NE ) AS REGIS TERED IN THE POLLEN RECORD FROM LAKE SALÊT

Marta Szal1, Mi ros³awa Ku pry janowicz1, Mariusz Wyczó³kowski2 1 Insti tute of Biol ogy, Uni ver sity of Bia³ystok, Œwierkowa 20b, 15-950 Bia³ystok, Poland; e-mail: [email protected], [email protected] 2 Wojciech Kêtrzyñski Museum, Plac Zamkowy 1, 11-400 Kêtrzyn, Poland; e-mail: [email protected]

Ab stract Pollen anal y sis of sedi ments from the upper part of bot tom depos its from Lake Salêt al lowed re con struc tion of main stages of the Late Ho lo cene veg e ta tion trans for mation in the Mr¹gowo Lake Dis trict (from ca. 3600 cal. years BC) and to cor re late some of these changes with im migra tion and eco nomic ac tiv ity of lo cal hu man groups. Signif i cant spread - ing of sec ond ary semi-nat u ral birch for est, de vel op ment of horn beam for est and in creas ing im por tance of anthro- pogenic open com mu ni ties were the most char ac ter is tic fea tures of veg e ta tion evo lu tion. A def i nite break down of elm took place be tween 2900 and 2500 cal. years BC, slightly af ter in creased con tri bu tion of birch in woodlands. Dis ap - pear ance of ha zel around 1200 cal. years BC, ac compa nied by ex pan sion of horn beam has been ob served and should be linked with ac tiv ity of the Z¹bie-Szestno type cul ture and the Lusatian cul ture tribes during the Bronze Age, but not with a climate change. Consid er able in ten sifi ca tion of set tle ment pro cesses re corded in the younger part of the Subat- lantic chronozone was one of the im por tant rea sons that were re sponsi ble for quick changes in for est struc ture. Strong and con tin ued de for es ta tion started as early as the end of the 10th century AD and was substan tially inten si fied in the first half of the 13th century.

Key words: veg e ta tion trans for ma tions, Late Ho lo cene, pol len anal y sis, Mr¹gowo Lake Dis trict, Masuria, NE Po land.

Manuscript received 28 October 2013, accepted 14 May 2014

IN TRO DUC TION From a paleoecological per spec tive north eastern Poland Pollen analy sis is one of the most widely used re search is an inter est ing region, because it is under influ ence of a tools in paleoecology that helps in detect ing responses of nat- tran sitional cli mate (e.g. Woœ 1999). Oc currences of differ - u ral vege ta tion to human im pacts through history, as well as ent air masses have wide range of in fluence upon natu ral dis- to climate and envi ron men tal change on a vari ety of tem po ral tri bu tions of plants typ i cal for both the mar i time (e.g. Cla- and spa tial scales (e.g. Prentice, 1988; Hunt ley, 1990). Pol- dium mariscus and Juncus subnodulosus) and the boreal cli- len records have been used to study biostratigraphy, vege ta - mate (among others: Polemonium caeruleum, Nymphaea tion patterns and agrarian history of the Masurian Lakeland candida and Nuphar pumila; Ga³ka et al., 2014). Such spe- for many years. Among the sites that have been palyno logi - cific char ac ter of this part of Po land is re flected also in geo- cal ly exam ined within mod ern standards, only a few in - bo tani cal divi sion of the country, in which it was in cluded in cluded pollen sequences cover ing the Subatlantic chrono- the Masuria-Belarus North Divide (Matuszkiewicz 2008). zone. These were pol len se quences from: Lake Miko³ajskie Palaeo eco logi cal re search in this area is a key to un der stand - (Ralska-Jasiewiczowa, 1966), Lakes Kruklin, Mamry and ing of mecha nisms and reasons for plant migra tion (Kupryja- Ta³ty (Stasiak, 1967, 1971), Lake Dga³ Wielki (Filbrandt- nowicz 2008; Lamentowicz et al., 2008). To assess, to what Czaja, 2000), Lakes Mi³kowskie, Wojnowo and £azduny, extent the records from mari time and more conti nen tal re- for mer Lake Staœwiñskie and the Szczepanki site (Wacnik, gions are differ ent from one another, more research and espe - 2009a, b; Wacnik et al., 2012a), Wielkie B³oto mire (Kar- cially multi-proxy in ves ti ga tion is needed. piñska-Ko³aczek et al., 2013) and Skaliski Forest (Ko³aczek Ar chae ol o gists have al ready dem on strated a spe cific and et al., 2013). However, beyond the profiles described by unique char ac ter of eco nomic ac tiv ity in north east ern Po land Wacnik (2009a, b; Wacnik et al., 2012a) most studied se- in rela tion to central Europe (e.g. Okulicz 1973; Karczewski quences were poorly supported by ra diocar bon dating, or the 2011). A partic u lar feature of this area was a prolonged ex is- ages were miss ing. This fact re stricted pos sible re con struc - tence of for ag ing and a very late ac cep tance of farming as the tion of rate of envi ron men tal changes and their corre la tion basis for food procure ment, in contrary to the rest of Poland with ar chae o log i cal, his tor i cal and cli ma tic data. (Wacnik et al., 2012a). 52 M. SZAL et al.

The lake is of glacial ori gin and is located in a depres sion surrounded by morainic hills. Steep escarp ments, over 10 m high, slope down to the water surface on both eastern and western lakesides; whereas northern and southern slopes are more gentle. The lake is divided into two parts – the smaller, northern part is named Lake Salêt Ma³y and the larger south- ern part is Lake Salêt Wielki (Fig. 1). Max i mum wa ter depth is 17.2 m and mean depth is 4.9 m (Jañczak, 1999). The lake is fed by a single stream and in ad dition to surface runoff, an inflow of shallow under ground water from the surround ing mo raine up land is prob a bly also pos si ble. The vege ta tion around Lake Salêt is almost devoid of forest and indi cates substan tial anthropogenic changes. The forest occurs in 10% of the area only whereas most is occu - pied by grasslands: meadows and pastures 30%, ara ble land of 60%. A land strip of 500 m around the lake is overgrown by her ba ceous veg e ta tion, with scat tered trees oc cur ring near buildings, along roads and streams and at the lakeshore only.

Coring and dat ing In winter 2010 a core of sedi ments was taken from the Fig. 1. Lo ca tion of Lake Salêt and cor ing site. frozen surface of Lake Salêt with the use of the Wiêc- kowski’s piston corer and the Kajak gravity corer. Coring was per formed close to the deep est part of the lake (wa ter This study is focused on recon struc tion of the Late Holo - depth 15 m), with bottom depos its 15.70 m thick (Fig. 1). The cene vege ta tion changes in the Mr¹gowo Lake District, present pa per deals with a top part of the log only (depth to based on pollen analy sis and supported by AMS radio car bon 10.80 m). This part of the core is com posed of calcar e ous dating. The pri mary aim was to present veg eta tion devel op - gyttja, rich in organic matter and rela tively poor in sand, clay ment in the surround ings of Lake Salêt from ca. 3600 cal. and silt. years BC and to pro vide data for eval u ation of woodland AMS ra dio car bon dat ing of seven pol len con cen trate trans for ma tion by a man. samples was per formed in the Gliwice Ab so lute Dat ing Methods Centre (Table 1). The isola tion of pollen concen - MA TE RIAL AND METH ODS trates from the sed iment was per formed by re moval of cal - cium carbon ate using 38% HCl and humic acids, using 10% Descrip tion of the study site KOH and 2–4 cm2 of sedi ment was used for this purpose. Lake Salêt (53°56'22.09"N, 21°19'20.19"E) is a eutro- Short-lived radionuclides 210Pb were measured in the up- phic water body with area of 327.7 hect ares (Jañczak, 1999). per most layer, 30 cm thick. The anal ysis was car ried out in It is lo cated be tween the villages Szestno, Ruska Wieœ, the Insti tute of Geolog i cal Sciences of the Polish Academy of Brodzikowo and in the centre of the Mr¹gowo Sci ences in War saw. Lake Dis trict, which is a part of the Masurian Lake Dis trict in north east ern Po land (Fig. 1).

Ta ble 1 AMS radio car bon dating of sedi ments from Lake Salêt

No. Depth (cm) Lab. No. Age 14C (BP) Cal i brated age range 95% (BP) Cal en dar age 1 330 GdA-2979 830±25 785 (95.4%) 689 1116-1261 A.D. 1058 (92.6%) 953 2 370 GdA-2980 1090±25 892-1011 A.D. 947 ( 2.8%) 939 1284 (94.6%) 1171 3 410 GdA-2981 1275±25 666-801 A.D. 1155 ( 0.8%) 1149 4 500 GdA-2982 1610±25 1546 (95.4%) 1415 404-535 A.D. 2713 (35.8%) 2629 2623 (55.2%) 2446 5 640 GdA-2983 2470±25 763-419 B.C. 2410 ( 1.5%) 2397 2392 ( 2.8%) 2369 6 710 GdA-2984 2700±25 2850 (95.4%) 2757 900-807 B.C. 7 800 GdA-2985 3030±25 3343 (95.4%) 3161 1393-1211 B.C. HOLO CENE CHANGES IN VEGE TA TION OF THE MR¥GOWO 53

Pol len anal y sis Samples for palynological analy ses (1 cm3) were col - lected usu ally at 5-10 cm inter vals; only in the bot tom part of the section (900-1080 cm) they were col lected ev ery 20 cm. Sam ples were pre pared us ing the standard proce dure of Erdtman’s acetolysis (Berglund and Ralska-Jasiewiczowa, 1986). Ac cord ing to de gree of con tam i na tion by min eral frac tion, samples were treated with a heavy liquid (CdI2+KI). Lycopodium tab lets were added to each sample to en able quan ti ta tive anal y sis of microfossil con cen tra tions (Stock- marr, 1971). Pol len anal ysis was car ried out with the Olympus BX43 light micro scope with magni fi ca tion of 600×; a larger magni - fica tion was used to identify problem atic and small palyno- Fig. 2. Age-depth model of the Lake Salêt pro file based on AMS morphs. For tax o nom i cal iden ti fi ca tion pol len keys (e.g. 14C and 210Pb dates. Beug, 2004) and a refer ence collec tion of mod ern pol len slides were used. vour of the increas ing role of birch. Oak was a compo nent of More than 1000 terres trial pol len grains were counted sev eral commu ni ties, es pe cially with ha zel and pine. Due to and identified in each sample. its high eco log i cal tol er ance, oak de vel oped in di verse hab i- Cal cula tions and pre senta tion of palynological data in a tats, from poor (Quercus petraea) to very rich (Quercus ro- sim pli fied per cent age pol len di a gram were per formed with bur) (Szymañski, 2006). From 3567 to 3399 cal. years BC, POLPAL for Windows soft ware (Nalepka and Walanus, oak was the most im portant com po nent of mixed and multi- 2003). AP+NAP sum was used for percent age calcu la tions. species decid u ous forests in the surround ings of Lake Salêt. The dia gram was divided into local pollen assem blage zones Ac cord ing to Ralska-Jasiewiczowa et al. (2003), there are no (L PAZ) with Constrained Cluster Analy sis (CONISS). To close an a logues to con tem po rary com mu ni ties dom i nated by dis cuss in ter ac tion of anthropogenic and nat u ral in flu ences, oak and ha zel. They were formed in re sponse to cy clic occu- the dia gram was constructed with groups of taxa estab lished pation and exploi tation of the land by the Neolithic people. by Behre (1981), with mod ifi ca tion suggested by Berglund A rising pollen curve of Corylus avellana was proba bly due and Ralska-Jasiewiczowa (1986) and Veski (1998). Pollen to slight opening of woodlands, which favoured inten sive de- curves of herbs were grouped into three cate go ries: (1) culti - velop ment of their understory. However, this process could vated crops, (2) weeds and ruderals, and (3) grass land and have been also induced by drier climate condi tions (low wa- eco log i cally un de fined plants. ter level in lakes and dry phase in peat bogs) between 4050 and 3650 cal. years BP (¯urek and Pazdur, 1999), and its RE SULTS great po tential for vege ta tion survival (Tallantire, 2002). In the pol len spec tra from the Lake Salêt profile, Viscum al bum Chro nol ogy has been recorded in that time. In the neigh bourhood of the 14C and 210Pb dates were used for construc tion of the stud ied area pres ence of this con tinen tal spe cies has been age-depth model for the analysed section (Fig. 2). This model also noted (Wacnik, 2009a) and it strongly suggested warm was built by means of a polyno mial 2nd degree curve fitting sum mers, with av er age tem per a ture of the warm est month in the POLPAL program (Nalepka and Walanus, 2003; above 15.5°C (Iversen, 1944). Walanus and Nalepka, 2004) and resulted in drawing the op- A distinct breakdown in the elm curve was noted in the timal depth-age curve on the basis of cali brated radio car bon Lake Salêt region between 2887 and 2563 cal. years BC dates. (Fig. 3). A sim ilar process occurred also in the Gostynin Lake District, where it was dated to 2850–2750 cal. years BC Palynological data (Wacnik et al., 2012b) and the increas ing role of hazel was observed in the follow ing centu ries. The same pattern was Palynological analy sis was performed for 128 samples noted in the surround ings of Lake Salêt, where beyond ha zel, from the upper part of the sec tion. Four lo cal pollen assem - in creased signif icance of pine and birch was re corded. Si - blage zones (L PAZ) were distin guished (Fig. 3, Table 2). mul ta neously with changes in for est com po si tion, dif fer ent open anthropogenic com mu nities with ruderal, weed and IN TER PRE TA TION AND DIS CUS SION grass land taxa de veloped, and in the youn gest part of this pe- riod cere als have also appeared in pollen spectrum (palynolo- Early and middle Subboreal (3600–1850 cal. gical human phase 1). years BC) The problem of decline of Ulmus in the Holo cene has been in tensively discussed during the last de cades. Differ ent Quercus-Corylus L PAZ, depth: 1080–870 cm hypoth e ses were proposed, e.g. climate change, human im - A vicin ity of Lake Salêt was densely for ested by multi- pact and finally, spreading of the so-called “Dutch” elm dis- species decid u ous woodland. Oak was the dom inant tree in ease caused by an ascomycete fun gus (Ceratocystis ulmi), the older part of this period, but it gradu ally declined in fa- and carried over by bark bee tles Scotylus (e.g. Rackham, 54 M. SZAL et al. n w o r b - y e r g

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) P A N ( axat ne axat llop breh fo spuorg dna )PA( sburhs dna seert fo axat detce axat fo seert dna sburhs )PA( dna spuorg fo breh llop les – mar g aid eg atne crep ne llop dei fil pmiS .têlaS ekaL ekaL .têlaS pmiS fil dei llop ne crep atne eg aid g mar – les .ajttyg su tir ted-su o era clac era o ted-su tir su .ajttyg . 3

. g i F HOLO CENE CHANGES IN VEGE TA TION OF THE MR¥GOWO 55

Ta ble 2 Lake Salêt – description of local pollen assem blage zones (L PAZ)

Name of L PAZ Depth [cm] and age [cal. De scrip tion years] of pollen sam ples Dom i na tion of Pinus sylvestris t. (21–47%); rela tively high frequency of Picea abies (to 7%); grad ual de crease of Alnus (to 7–8%), Quercus (to ca. 2–3%) and Carpinus betulus (to ca. 1%); rather low val ues of Betula alba t. (below 23%); low Pinus-Picea-NAP fre quency of Corylus avellana (to 3,5%), Ulmus (to 1,8%), Tilia cordata (to 2,4%), Fraxinus excel sior (to 0,8%), Salix (to 0–365 cm 1,3%) and Fagus sylvatica (to 0,9%); rela tively fre quent Juniperus communis; sharp increase of NAP from 7% to 16%, 2005–981 AD and then to 48% – highest amount of human indi ca tors in the whole dia gram (cul tivated plants to 16%; ruderals and weeds to 8%). Upper boundary: rapid fall of Betula alba t. be low 18%; rise of Pinus sylvestris t. above 20% and Picea abies to ca. 5%. Ab so lute dom i na tion of Betula alba t. (20–44%); high tri ple cul mina tion of Carpinus betulus (18%, 12% and 12%); still Carpinus-Betula-Quercus high values of Quercus (8–14%); strong fluctu a tions of Pinus sylvestris t. (8–30%); low propor tion of Ulmus (ca. 2%), 370–770 cm Tilia cordata (ca. 3%), Fraxinus ex cel sior (ca. 1%), Salix (to 1.5%) and Fagus sylvatica (be low 1%); grad ual de crease of 960 AD–1183 BC Alnus from 20–24% to 5–11%; increase of herba ceous plants to 12% (culti vated plants up to 3%; ruderals and weeds up to 2%). Carpinus-Corylus-Quercus- Upper boundary: rapid rise of Betula alba t. above 35% and fall of Corylus avellana. Betula Rapid in crease of Alnus (from 10% to 23%); decrease of Quercus to 8% and Corylus avellana to 6%; low percent ages of 780–860 cm Ulmus (1–2% and Tilia cordata (2–4%); the first culmi na tion of Carpinus (10%); rel atively low val ues of Pinus sylvestris 1246–1821 BC t. (15–28%) and Betula alba t. (18–32%); rise of her baceous plants to ca. 10% with clear increase of human indicators. Upper boundary: rise of Carpinus betulus to ca. 7%. Rel a tively high fre quency of Alnus to ca. 22–29%; cul mina tions of Quercus (26%) and Pinus sylvestris t. (38%) in the bot- Quercus-Corylus tom part of the zone, and then their grad ual decline; low frequency of Ulmus to ca. 1% at the depth 960 cm (ca. 2563 BC); 870–1080 cm al most con stant Tilia cordata curve (ca. 5%); fluctu a tions of Corylus avellana around 20%; gradual rise of Carpinus 1894–3567 BC betulus in the youn ger part of the zone; system atic increase of herba ceous plants (NAP) to ca. 5% corre spond ing with ris- ing values of Betula alba t. up to 19%.

1980). Elm reduc tion corre sponds quite logi cally with such man phase 2). Moreover, human activ ity result ing in clearing pathogenic hypoth e sis and it is not very concor dant with mi- of some woodland areas, promoted spreading of hornbeam. gration routes of suc ces sive Neolithic cul tures. However, it Hornbeam has been well docu mented as a moder ate pio neer seems convinc ing (Girling and Greig, 1985; Birks, 1986, spe cies in re cent years. It has rap idly col o nized ar eas af ter Ralska-Jasiewiczowa and van Geel, 1998; Peglar, 1993; fall of oak-hornbeam woods in the Bia³owie¿a Forest (Ber- Innes et al., 2010) that Neo lithic people took advan tage of nadzki et al., 1998). elm disease to clear decid u ous woods growing on more fer- In the younger part of this period a hornbeam area has de- tile soils. Pat tern of dis tri bution of Ulmus on Polish isopollen creased, and with high proba bil ity it could result from inten - maps in di cates that its de cline has been more dif fer en tiated in si fi ca tion of set tle ment at the tran si tion of the Neo lithic and time and space than pre vi ously ac cepted (Ralska-Jasiewi- Bronze Ages (Manasterski 2009). Cleared ground was used czowa et al., 2003). For in stance, in the Bieszczady Moun- as pas tures, mead ows, fields and lo cali ties for set tlements. tains elm occurs around 4400–4200 14C years BP (2450– Large ar eas of grass lands at that time indi cated pre domi - 2250 cal. years BC) only (Ralska-Jasiewiczowa, 1980) and nance of ani mal husbandry rather than culti va tion. in some lowland sites as early as 6000–5700 14C years BP Start ing from ca. 1850 cal. years BC a gradual de crease (4050–3750 cal. years BC) (Lata³owa, 1992). Decline of in impor tance of hazel was noted. Very distinct decline of Ulmus is not a synchro nous event in Esto nia, as the dates this species at that time is also recorded in numer ous other range from 3800 to 3200 cal. years BC, likely due to human pollen di agrams from Poland (e.g. Ralska-Jasiewiczowa et ac tiv ity, con nected with in tro duc tion of ar a ble farm ing rather al., 2003), as well as in several sec tions from adja cent coun- than by natu ral factors (Saarse and Veski, 2001). In the pol- tries, mainly from Germany (e.g. Pott, 1986; Brauer et al., len di agram from Lake Salêt a very dis tinct elm de cline is 2000; Kubitz, 2000). The onset of this process was deter - dated to the inter val 2887–2563 cal. years BC (Fig. 3). Simul - mined to ca. 3400 14C years BP (ca. 1750 cal. years BC) in the taneously with the decline of Ulmus, Picea abies de vel oped Lake Goœci¹¿ (Ralska-Jasiewiczowa et al., 2003), and at ca. in the vi cin ity of the studied site. 3490 14C years BP (ca. 1860 cal. years BC) in lakes from the Eifel region in Germany (Brauer et al., 2000). At about 1700 Late Subboreal (1850–1200 cal. years BC) cal. years BC forest transfor ma tions occurred around the Carpinus-Corylus-Quercus-Betula L PAZ, depth: Puck Bay in northern Poland (Miotk-Szpiganowicz, 1997; Uœcinowicz and Miotk-Szpiganowicz, 2003), and somewhat 860–770 cm sim i larly, Carpinus betulus started to develop in forest at the The most signif i cant feature of this period was a very dis- ex pense of Corylus avellana in the ¯u³awy region in the tinct spreading of hornbeam in stands of the Lake Salêt re- Vistula River mouth (Zachowicz et al., 1982, Zachowicz and gion. This process clearly coin cided with decreased impor- Kêpiñska, 1987). There are some hints of both a ther mal os- tance of hazel as well as lime, elm and oak. Changes in for est cilla tion towards a cooling (Zolitschka, 1992) and an in- com po si tion were most ev i dently a con se quence of lo cal crease (al though os cil lat ing) in cli mate hu mid ity (Ralska- anthropogenic pressure (a begin ning of the palynological hu- Jasiewiczowa and Starkel, 1988), which might be a proba ble 56 M. SZAL et al. rea son for de cline of ha zel. In creas ing cli mate hu mid ity ca. 500 cal. years BC (Fig. 3), just af ter the Lusatian culture set - 1700 cal. years BC has been re constructed for northern Brit - tlement, which was the most widespread prehis toric cul ture ain (Charman et al., 2006) and north ern Norway (Vorren et in Poland, charac ter ized by very high density of popu la tion al., 2007). From ca. 1500 cal. years BC a groundwa ter table (God³owski and Koz³owski, 1979). A decline of this culture rose in mires of North Ireland (Swin dles et al., 2010) and in is re flected in a pollen di a gram from Lake Salêt as the tem po - south ern Fin land (Väliranta et al., 2007). rary re gen er a tion of for est com mu ni ties, first of pi o neer A broad spectrum of data from Polish flu vial, lake and birch woods and then of decid u ous forest with dom inant peatland en vi ron ments in di cated also an in crease in hu mid ity hornbeam. The follow ing period of woodland reduc tion, between 4400–4100 and 3500–2900 cal. years BP (2450– with areas defor ested or thinned by a man became overgrown 2150 and 1550–950 cal. years BC, respec tively) (Ralska- by vari ous anthropogenic open commu ni ties and by a helio- Jasiewiczowa and Starkel, 1988; ¯urek and Pazdur, 1999; philous birch forest. The second, less inten sive hornbeam ex- Starkel et al., 2006; Paw³owski et al., 2012). This part of the pansion started from ca. 200 AD. A history of hornbeam in Subboreal chronozone was consid ered by Starkel (1990) and the Ho lo cene in Cen tral Eu rope was strictly con nected with Starkel et al. (2006) to have been a wet pe riod, char ac ter ized hu man ac tiv ity (see Ralska-Jasiewiczowa, 1964; Ralska- by more frequent rainy sea sons. In the Pol ish isopollen map Jasiewiczowa et al., 2004). This species was partic u larly for 4000±100 14C years BP (ca. 2050 cal. years BC) Miotk- dev as tated in pe ri ods with in ten sive set tle ment due to fer til - Szpiganowicz et al. (2004) indi cated that nearly the whole ity of the occu pied hab itats. On the other hand, pe riods with area of Poland was occu pied by Corylus avellana pollen con - re duced anthropogenic pres sure al lowed par tial re gen er a tion tents of 10–15% (western, central and northern Poland, of forest commu ni ties with hornbeam, because of its great re- Carpathians Moun tains and Roztocze). The values over 15% gen er a tive abil i ties. occurred in the peri-Baltic area only and below 10% in the Large areas in the vi cin ity of Lake Salêt were still occu - eastern part of the country up to the Podlasie border and the pied by oak. Birch forest played a signif i cant role in the land- Carpathian Foredeep in the Ma³opolska Upland. The later scape, be ing prob a bly sec ond ary semi-nat u ral com mu ni ties de creased to be low 5% pro ceeded from SE-E across the which were formed in habi tats transformed by exploi ta tion whole country and the mentioned authors assumed that there during earlier settle ment stages. An expan sion of birch was were other rea sons than a con tinen tal climate. It can be con - also the most char ac ter is tic fea ture of this pe riod in other cluded that devel op ment of the Early Bronze Age cultures, parts of the Masurian Lake District. For exam ple, this pro- prac tic ing rather prim i tive ar a ble ag ri cul ture but broadly de- cess was evi dent in pollen spectra from Lake Miko³ajskie pendent on cattle grazing (Manasterski 2009) may have con- (Ralska-Jasiewiczowa, 1966), Woryty (Pawlikowski et al., trib uted to shrink age of ha zel dis tri bu tion. 1982), Lake Dga³ Wielki (Filbrandt-Czaja, 2000) and Lakes In the middle of this period (ca. 1500 cal. years BC) areas Mi³kowskie, £azduny and Wojnowo (Wacnik et al., 2012a). with lime and tempo rarily, also the ar eas with elm have de - Sim i lar de vel op ment of com mu ni ties with dom i nant birch clined. The elm de cline ca. 1700 years BC was also re corded was re corded in more or less the same time in di a grams from in sections from the Lakes Mi³kowskie and Wojnowo (Wac- other parts of Poland, for instance from the Biebrza River nik et al., 2012a). If these changes are ac cepted to have re - Valley (Balwierz and ¯urek, 1987), Suwa³ki Lake District sulted from hu man ac tiv ity, then increased pol len pro duction (Lake Kluczysko – Wacnik, pers. comments), cen tral Po land of Corylus avellana can be explained by opening of the pre- (Biñka et al., 1991), Gostynin Lake District (Wacnik et al., viously densely shaded decid u ous forest by cutting of elm 2011; Wacnik et al., 2012b) and from Wielkopolska (To- and lime branches for cattle fodder. Patches of open hab itats bolski and Okuniewska-Nowaczyk, 1989). In the vicin ity of began to spread in the area. Lake Salêt there were also pine and dif fer ent mixed for ests with pine-oak and pine-birch. Late Subboreal and early Subatlantic (1200 cal. Alder grew on wetland habi tats in the surround ings of years BC –1000 AD) the lake and in out skirts of rivers val leys. A rel a tively small Carpinus-Betula-Quercus L PAZ, depth: area was oc cu pied by elm, lime, ash and spruce. Other trees were rare. 760–370 cm The rise in frequency of pollen in dica tors of grasslands, Deep changes in structure of a mixed de cidu ous forest i.e. meadows and pastures (the palynological hu man phase 3) took place at the very begin ning of this period, indi cated by indi cates that surround ings of Lake Salêt have been to some rapid spread ing of birch. Veg e ta tion trans for ma tions were extent influ enced by a man. Inten sive agri cul ture and a broad caused both by nat u ral inter gla cial suc ces sion pro voked by extent of disturbed soil with high NO3 con tent were confir- climate change and by human economic activ ity. The domi - med by widespread presence of weeds and ruderals. Open ar- nating role of birch and hornbeam made forest structure dif- eas were used for culti va tion as denoted by contin u ous pres - ferent from the previ ous one. Hornbeam formed presum ably ence of Cerealia t. pol len, reg u lar oc cur rence of Secale the high est canopy layer that casted shadows on all lower cereale pollen, and sporadic appear ance of Triticum t., Avena storeys of the forest. This elimi nated hazel from phytoce- t., and Hordeum t. pol len (this is not shown in the pol len di a- nosis and lim ited its occur rence, proba bly to forest periph er - gram). Rye, as a wind-polli nated crop, has higher pollen pro- ies only. Hornbeam, due to its high ability to form suckers duc tion and better dis persal pos si bil i ties than other ce re als, from trunk or stumps, could be the main ele ment of regen er - and thus its frequen cies must repre sent not only local culti va - ated forest, quickly occu py ing all larger clearings (Wacnik et tion but also a re gional background (cf. Koff and Punning, al., 2012b). Max i mum horn beam ex pan sion oc curred about 2002). It is also possi ble that fields at the edge of Lake Salêt HOLO CENE CHANGES IN VEGE TA TION OF THE MR¥GOWO 57 and ap pear ance of for est graz ing in di ca tors (e.g. Pteridium to regen er ate in a compar a tively short time and to spread over aquilinum) point to somewhat more cat tle-rear ing ac tiv ity in certain ter rito ries as one of the main com po nents of an the area. In thinned and grazed forests Juniperus communis oak-horn beam for est. and Calluna vulgaris occurred. Some open areas may have become overgrown by wood, in which Picea abies was an impor tant ele ment. Spruce ex- Late Subatlantic (1000–2005 AD) pansion in the Lake Salêt re gion was dated to ca. 1000 AD, as has been observed in some other dia grams from the Masurian Pinus-Picea-NAP L PAZ, depth: 365–0 cm Lake District (e.g. Filbrandt-Czaja, 2000; Wacnik et al., The be ginning of this pe riod was charac ter ized by rapid 2012a). An in creasing presence of Calluna vulgaris and de for es ta tion and by marked in crease in hu man ac tiv ity. Pteridium aquilinum, both of which favour poorer sandy Withdrawal of tree pollen must have been connected with soils, suggests openings in a dry pine forest and subse quent forest clearing under taken in order to enlarge the areas of cul- appli ca tion of these plots for ani mal husbandry. Pinus sylves- tivated fields or pastures and with timber exploi ta tion, which tris spread most proba bly on less fertile, sandy habi tats, is doc u mented in ar chae o log i cal and his tor i cal sources (e.g. which were covered previ ously by a mixed conif er ous forest. Wróblewski et al., 2003; Nowakiewicz, 2006, 2010). From Current distri bu tion of pine was also connected with human around 1000 AD exten sion of human economic activ ity star- ac tiv i ties. Cul mi na tions of Pinus sylvestris t. pollen curve ted to transform the natu ral envi ron ment in a process persis tent im me di ately af ter each de for es ta tion were al most cer tainly and contin ued up to the present time. Accord ing to Ralska- due to transport of pine pollen from distant areas and were Jasiewiczowa (2006) this gener ated an osten si ble picture re- also an evi dence for spread ing of pine to hab itats occu pied sem bling the ap proaching end of the Ho locene intergla cial cy- formerly by an oak-hornbeam forest. cle: dis ap pear ance of a de cid u ous for est and first mass sprea- At the same time her ba ceous veg eta tion at tained greater ding of pine, fol lowed by to tal de for es ta tion and oc cu pa tion of sig nif i cance. Pol len grains of cul ti vated plants oc curred fre- its terri tory by open vege ta tion. Low values of tree pollen and quently. Hith erto un re corded Fagopyrum pollen ap peared pre dom i nance of long-dis tance trans ported pine pol len dem - also in pollen spectra from the Lake Salêt profile (1116 AD), onstrated exis tence of large open areas, which in turn sug- in di cat ing that buck wheat was prob a bly cul ti vated. This is in gested pro gres sive de vel op ment of set tle ments. agree ment with other Euro pean data showing that buck- At the begin ning of this zone there was a period of in - wheat, which was native to Central Asia and was intro duced creased Carpinus betulus pollen percent ages, which was to Europe by the Mongols, was culti vated from 12th– 14th con tem po ra ne ous with slightly grow ing hu man eco nomic cen tu ries on wards (Speranza et al., 2000; Lata³owa et al., ac tiv ity. A great re gen er a tion abil ity of horn beam al lowed 2007; StanèikaitÅ et al., 2008), although the earli est Polish for its fast ex pan sion in for est un til the early Mid dle Ages. record, known from the town of Wolin, dated back to 9th–10th Soon after wards, ca. 1250 AD, the areas domi nated by horn- cen tu ries (Alsleben, 1995). Fagopyrum is an in sect-pol li- beam shrank grad u ally. Al most to tal elim i na tion of Carpinus nated plant which produces a small quantity of pollen that is betulus from for est stands was proba bly the effect of farm ing usually underrepresented in pollen spectra. Results of mod- in ten si fi ca tion and grow ing de mand for fer tile soils. Di min- ern pol len mon i tor ing in di cated that buck wheat, wheat and ished sig nif i cance of Carpinus betulus in a forest could have barley pollen ap peared in low percent ages, even when fields been also caused by many years’ cat tle graz ing, a con se - were lo cated at a small dis tance from the sampling place quence of which was a forma tion of well-lit oak-pine forest (Pidek, 2009). Field and ruderal weeds appeared in higher (Faliñski and Pawlaczyk, 1993). per cent ages. An impor tance of oak was still sig nif i cant ini tially (981– 1033 AD). This could have been caused by selec tive protec - CONCLU SIONS tion of this tree by a man in or der to provide acorns, which could be stored for up to three years (Weckerly et al., 1989) Pollen analy sis of the up per part of the sedi men tary se- and used as fodder for pigs, partic u larly in winter time. quence from Lake Salêt allowed recon struc tion of vege ta tion Acorns are a source of a high-energy food and nowa days, in changes in the Mr¹gowo Lake District during Sub-boreal and some seasons of a year, they may con sti tute up to 50% of the Subatlantic chronozones of the Holo cene (from ca. 3600 cal. diet of some wild an imals. Oak leaves are also rich in pro teins years BC). (Pekins and Mautz, 1988; Elowe and Dodge, 1989). It is pos- In the early Subboreal chronozone (3700–2900/2600 sible that acorns were also consumed by people, just as was cal. years BC) oak-pine forest devel oped, but mixed decid u - the case up to re cently among the ab origi nes of North Amer- ous woodland with elm, lime and hazel still occurred. ica. Accord ing to Pekins and Mautz (1988) acorn produc tion Very dis tinct trans for ma tion of a for est co in cided with could have in creased sig nif icantly by se lective fall of trees in tran sition between early and middle Subboreal chronozone a forest and by protec tion of mature speci mens. A sudden (2900–2600 cal. years BC). This is indi cated by defi nite breakdown in a role of Quercus ca. 940 AD, syn chronous breakdown of elm, preceded by gradual drop of oak and with Carpinus betulus de cline and in crease of Pinus sylves- spread of birch, which started from ca. 3300 cal. years BC. tris t., should be strictly connected with human activ ity. Then, from ca. 2300 cal. years BC, devel op ment of horn- Ralska-Jasiewiczowa (1964) ex plained that oak and horn- beam and tem po rary exten sion of spruce oc curred. At that beam were par tic u larly dev as tated spe cies in pe ri ods of in - time the first clearing by human communities occurred. tensive set tlement on account of fertil ity of the occu pied In the late Subboreal chronozone grad ual elim ina tion of hab itats. She concluded also that these trees had great ability hazel (1800–1200 cal. years BC) was accom pa nied by ex- 58 M. SZAL et al. pan sion of horn beam and pro gres sive de vel op ment of birch. cene proxy cli mate records: stacked Holo cene peatland pa- This process should be proba bly linked with human activ ity laeo-wa ter ta ble re con struc tions from north ern Brit ain. Qua - rather (the Lusatian culture) and not with a climate change – ter nary Sci ence Re views 25, 336–350. crop culti va tion started from 1800–1700 cal. years BC. 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