Late Holocene palaeoclimate variability: The significance of bog pine dendrochronology related to peat stratigraphy. The Puścizna Wielka raised bog case study (Orawa – Nowy Targ Basin, Polish Inner Carpathians)

Late Holocene palaeoclimate variability: The significance of bog pine dendrochronology related to peat stratigraphy. The Puścizna Wielka raised bog case study (Orawa – Nowy Targ Basin, Polish Inner Carpathians)

Quaternary Science Reviews 148 (2016) 192e208 Contents lists available at ScienceDirect Quaternary Science Reviews journal homepage: www.elsevier.co...

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Quaternary Science Reviews 148 (2016) 192e208

Contents lists available at ScienceDirect

Quaternary Science Reviews journal homepage: www.elsevier.com/locate/quascirev

Late Holocene palaeoclimate variability: The significance of bog pine dendrochronology related to peat stratigraphy. The Puscizna Wielka raised bog case study (Orawa e Nowy Targ Basin, Polish Inner Carpathians)  c, d, Marek Kra˛piec a, *, Włodzimierz Margielewski b, Katarzyna Korzen a d e _ Elzbieta Szychowska-Kra˛piec , Dorota Nalepka , Adam Łajczak w, Poland AGH University of Science and Technology, A. Mickiewicza Ave. 30, 30-059, Krako w, Poland Institute of Nature Conservation, Polish Academy of Sciences, A. Mickiewicza Ave. 33, 31-120, Krako c w, Poland Kazimierza Wielkiego Str. 110/2-3, 30-074, Krako d w, Poland W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz Str. 46, 31-512, Krako e w, Poland Pedagogical University, Institute of Geography, Podchora˛ z_ ych Str. 2, 30-084, Krako a

b

a r t i c l e i n f o

a b s t r a c t

Article history: Received 11 March 2016 Received in revised form 17 June 2016 Accepted 19 July 2016

The results of dendrochronological and palynological analyses of subfossil pine trees occurring in the peat deposits of the Puscizna Wielka raised bog (Polish Carpathians, Southern Poland) e the only site with numerous subfossil pine trees in the mountainous regions of Central Europe presently known e indicate that the majority of the tree populations grew in the peat bog during the periods ca 5415 e3940 cal BP and 3050e2560 cal BP. Several forestless episodes, dated to 5245e5155 cal BP, 4525 e4395 cal BP and 3940e3050 cal BP, were preceded by tree dying-off phases caused by an extreme periodical increase in humidity and general climate cooling trends. These events are documented based on analyses of pollen and non-pollen palynomorph assemblages, dendrochronological analyses of the trees, as well as numerous radiocarbon datings of the sediment horizons occurring within the peat bog profile. The phases of germinations, and, in turn, of tree and shrub invasions of the peat bog areas have been closely connected to drying and occasional warming of the regional climate. The last of the forestless periods began about 2600 years ago and continued up to the very recent times. Currently, as a result of desiccation of the peat bog and the lowering of the groundwater level (due to improved water drainage system), pine trees have returned the peat bog again. These results demonstrate that studies of subfossil bog-pine trees are quite effective in documenting and reconstructing periods of humidity fluctuation that occurred within the Carpathian region over the last several millennia. © 2016 Elsevier Ltd. All rights reserved.

Keywords: Climate change Raised bog Pinus sylvestris Subfossil wood Tree-rings analyses Multi-proxy analysis Holocene Polish Inner Carpathians

1. Introduction Subfossil pine wood (Pinus sylvestris L.) occurring in the European peat bogs, has been commonly used in dendrochronological studies to reconstruct and establish chronology of the Holocene palaeoclimatic events. Numerous studies have been carried out in Northern and Western Europe, including in Germany (Leuschner

* Corresponding author. E-mail addresses: [email protected] (M. Kra˛ piec), [email protected].  ), d.nalepka@botany. pl (W. Margielewski), [email protected] (K. Korzen pl (D. Nalepka), [email protected] (A. Łajczak). http://dx.doi.org/10.1016/j.quascirev.2016.07.022 0277-3791/© 2016 Elsevier Ltd. All rights reserved.

et al., 2007; Eckstein et al., 2008, 2010, 2011), Sweden (Gunnarson, 1999, 2008; Gunnarson et al., 2003; Edvardsson, 2010; Edvardsson et al., 2012), Great Britain (Lageard et al., 1995, 2000; Moir et al., 2010, 2012), Scotland (Bridge et al., 1990; Moir et al., , 2001; 2010); Ireland (Pilcher et al., 1995), Lithuania (Pukiene Edvardsson et al., 2016) (Fig. 1a- sites: 1e7) and Poland (Fig. 1a e site 7) (Barniak et al., 2014). A considerable portion of the compiled chronologies of subfossil pine trees are floating chronologies, dated using the radiocarbon method (among others Lageard et al., 1999; , 1997; Eckstein et al., 2009; Edvardsson et al., 2016), with Pukiene notable exceptions of chronologies from Northwestern Germany (see Fig. 1a - site 1) and Southern Sweden (Fig. 1a - site 2) where absolute dating was possible due to the heteroconnection with the

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Fig. 1. Study area: a - Location of the Puscizna Wielka raised bog; locations of other European peat bogs deposits with Scots pine (Pinus sylvestris) occurrences are also shown: 1 e Germany (Eckstein et al., 2011); 2 e Southern Sweden (Edvardsson et al., 2012); 3 e England (Lageard et al., 2000); 4 e Scotland (Moir et al., 2010); 5 e Northern Ireland (Pilcher , 2001; Edvardsson et al., 2016); 7 e Northern Poland (Barniak et al., 2014); 8 e Puscizna Wielka raised bog; b e SRTM topography model of the et al., 1995); 6 e Lithuania (Pukiene Puscizna Wielka bog within the Orawa - Nowy Targ Basin.

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Lower Saxony bog oak chronology (Eckstein et al., 2009; Edvardsson et al., 2012). In East-Central Europe, no systematic studies using subfossil wood from peat deposits to analyze Holocene paleoclimate are known to have been conducted or published. The Puscizna Wielka raised bog located in the Orawa-Nowy Targ Basin (Inner Carpathians, Southern Poland) provides an excellent opportunity for such study (Fig. 1a and b). Numerous pine tree trunks are buried (Fig. 2) in this peat bog which formed during the Middle Holocene (sensu Walker et al., 2012). Based on data collected during dendrochronological analyses of tree-trunks excavated from this peat bog, the compilation of pine chronologies that span over a period of 1700 years, covering time eranges 5415e3940 cal BP and 3050e2560 cal BP has been completed (Kra˛ piec and SzychowskaKra˛ piec, 2016). The main purpose of this study is the use of the dendrochronological analysis to reconstruct bog pine population dynamics, affected by the paleoecological changes that occurred during the woodland stages of the peat bog. These changes, which were directly related to more widespread palaeoclimatic fluctuations, such as periods of cooling and moistening, have been recorded in peat deposits for the last 5500 years, and are reconstructed using multi-proxy analysis.

mountains (such as Pinus sylvestris) (Staszkiewicz and Tyszkiewicz, 1969; Koczur, 2008) are present. An exceptional feature of the Orawa-Nowy Targ peat bogs is the occurrence of mountain pine (Pinus mugo Turra), typically found at higher altitudes, and the  gger (Staszkiewicz presence of rare swamp pine Pinus x rhaetica Bru and Tyszkiewicz, 1969). The Puscizna Wielka peat bog occupies a special place among the peatlands of the Orawa - Nowy Targ Basin due to its large size, the considerable thickness of the peat, and preservation of the large part of the dome in its natural condition until today. This peat bog is located in the western part of the Orawa-Nowy Targ Basin, on the western side of the European Watershed and within the catchment area of the Czarna Orawa (Fig. 1b). The bog occupies the altitude of 639.0e681.0 m a.s.l. whereas its dome is situated at an altitude of 650.0e679.0 m a.s.l (N49 26.321'; E19 46.561'). The present dimensions of the peat bog are 4  2 km, and of the dome 3  1.5 km (Fig. 2a). While the peat deposits are on average 3 m thick, thicker peat sequences reaching up to 10 m have been identified (Baumgart-Kotarba, 1991-1992). The peat deposit have been periodically excavated here for at least 200 years. Due to the extensive peat excavation since 1967 by the Zakład Torfowy factory (the mining field area is about 12 ha), numerous tree trunks have been exposed at the current surface of the peat bog (Fig. 2b).

2. Study area

3. Materials and methods

The Puscizna Wielka raised bog is located in the Orawa - Nowy Targ Basin, a relatively large intermontane basin that has a shape of an irregular triangle covering an area of 262 km2. The basin is situated at the altitude of ca 490e680 m a.s.l., and borders the Western Beskidy Mountains (Western Outer Carpathians) to the north and the Pieniny Klippen Belt, Spisko-Gubałowskie Foothills and the Tatra Mountains (Inner Carpathians) to the south (Fig. 1b). The main European Watershed dividing the catchment areas of the Baltic Sea (the Czarny Dunajec e a tributary of the Dunajec in the Vistula Basin) and the Black Sea (the Czarna Orawa/Black Orava e a tributary of the V ah in the Danube Basin) runs accross the basin (Baumgart-Kotarba, 1991-1992). Geologically, the Orawa - Nowy Targ Basin is a tectonic intermontane depression filled with Neogene and Quaternary deposits (Watycha, 1976). The altitude of the Orawa-Nowy Targ Basin is at some places 1000 m less than the altitude of the neighbouring mountainous surrounding the basin (Fig. 1b). Such morphology causes frequent occurrences of temperature inversion (on average, close to 260 days annually), which in turn causes the occurrence of cold air pockets and contributes to the relatively long retention of snow cover. The Basin is situated in a moderately cool climatic region; the mean annual temperature is only þ5.5  C here (Hess, 1965; Obre˛ bskaStarklowa, 1977). On the other hand, during the summer months (MayeOctober) precipitation greatly exceeds evaporation with average total annual amounts of 740e815 mm and 330 mm for precipitation and evaporation, respectively (Obre˛ bska-Starklowa, 1977; Kowanetz, 1998). Prevailing heavy cloudiness and frequent fog further contribute to the condensation of water within the basin (Staszkiewicz and Szela˛ g, 2003). Numerous dome-shaped raised bogs, developed in some cases since the Late Glacial are located within the Orawa-Nowy Targ Basin (Fig. 1b) (Koperowa, 1962; Obidowicz, 1990; BaumgartKotarba, 1991e1992; Łajczak, 2006; Kołaczek et al., 2010). The unique combination of habitat and climatic conditions within the Orawa-Nowy Targ Basin, as well as its proximity to high mountain ranges, have lead to the formation of distinctive vegetation within the local peatlands (Koczur, 2006, 2008). Apart from numerous glacial relics found in this area (e.g. Eriophorum gracile; Rubus chamemorus), lowland species that are not found higher in the

3.1. Field sampling Subfossil pine trees occurring in the peat deposit are most commonly found as horizontaly-oriented almost complete trunks, sometimes even with the remnants of roots. A total of 614 wood samples were collected for the dendrochronological analyses during field work conducted over a three year period (2011e2014). Majority of samples were obtained from almost entirely preserved trunks of fallen pine trees, which often had their roots still attached. These samples were collected as round disks (slices) by cross-cutting trunks and stumps using a gas-engine chainsaw. A considerably smaller number of samples (33) were collected just from stumps with well-preserved root system. The original position of the sampled subfossil pine tree in the bog profile was usually not known because their trunks were recovered from the peat prior to sampling, during the course of the current commercial exploitation (Fig. 2b). However, with respect to the above-mentioned stumps with completely preserved root systems, it is safe to assumed that they have remained until the present day almost in situ. This conclusion is partly based on the observation that even mineral soil from the initial phase of the tree growth on the peat bog could still be seen within the root system attached to two of the sampled stumps. Prospecting drillings on the peat bog deposits were made using a spiral bit. Drilling and sampling (see Fig. 2a) was done at four locations (in the vicinity of the occurrence of subfossil wood) within profiles of varying thickness of the peat deposits. Cores were taken with an INSTORF sampler (a Russian peat sampler), 8 cm in diameter. In total, 8 cores [logs] ranging in lenght from 2.4 to of 4.5 m, were taken using the INSTORF drill; subsequently three of the cores were subjected to detailed analyses (see Fig. 2c e logs: PW6; PW2; PW7). 3.2. Sediment analyses Identification of the organic deposits was performed in thin sections using a light microscope and was based on a tissue analysis of peat samples. The peat types were identified according to peat classification proposed by Tołpa et al., 1971. Mineral sediments occurring at the bottom of peat bog were identified based on the

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Fig. 2. Plan of the Puscizna Wielka bog showing locations of boreholes (a) (modified after Łajczak, 2006) and view of the exploitation field (b); c e lithostratigraphical profiles of the Puscizna Wielka peat bog, with loss on ignition curves (for each 2.5 cm deposit sequences), pollen LPAZ (with modelled age of their boundaries based on age-depth curve) and radiocarbon 14C datings (both conventional and calibrated). The position of the artefact (on the photo) is marked on the peat profile. The position of subfossil trees in the deposits reconstructed on the basis of time-depth curve analysis (see Fig. 3).

Bouyoucos-Casagrande areometric analysis (Mycielska-Dowgiałło and Rutkowski, 1995). Minerogenic sediments were classified according to Pettijohn's classification (Pettijohn, 1975; Battaglia et al., 2002), using the Wentworth grain size scale (Wentworth, 1922). For the mineral sediments, granulometric indexes such as mean

grain size (Mz), standard deviation (s1) and skeweness (Sk), were calculated according to the procedure proposed by Folk and Ward (1957) (Fig. 2c). Loss on ignition analyses of sediments were made for each 2.5 cm thick layer of the analysed logs. Pre-prepared sections of the

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log were dried and subjected to heating for 4 h in a muffle furnace at 550  C, according to the procedure proposed by Heiri et al. (2001). In total, 442 loss on ignition analyses were performed providing data for constructing loss-on-ignition curves for all three analysed profiles (Fig. 2c). 3.3. Radiocarbon dating and age-depth model Conventional radiocarbon dating of organic material using the liquid scintillation counting method (LSC) was performed in the Laboratory of Absolute Dating in Krakow (Poland). Samples were chemically pre-treated using the AAA (acid-alkali-acid) method. The procedure included standard synthesis of benzene from organic samples (Skripkin and Kovalyukh, 1994). 14C measurements were carried out with a 3-photomultiplier spectrometer, the HIDEX 300SL (Kra˛ piec and Walanus, 2011) and Quantulus 1220. Calibrated radiocarbon ages (cal a BP) were determined using the IntCal13 radiocarbon calibration dataset (Reimer et al., 2013) and the OxCal 4.2 calibration software (Bronk Ramsey, 2009). All presented margins of error are the uncertainties at a 95.4% probability. A total of 25 14C datings were made on samples of various compositions/ origin taken from the sediments of the Puscizna Wielka bog (Tables 1 and 2). The chronology (age-depth curve) of the Puscizna Wielka bog (PW-2 log in Fig. 2c), is based on the OxCal P_Sequence model (Bronk Ramsey, 2008). In total 17 radiocarbon dates were used for the construction of the age-depth model (Table 2). The changes in the pattern of peat accumulation (e.g. from minerogenic to ombrogenic) were factored in during the creation of the age-depth model. Therefore, single-boundary constraints were introduced in our model, and assigned to the depths of 328 cm (Fig. 3). The overall agreement index of this model amounts to 66%. The k parameter of the P_Sequence function (describes a magnitude of fluctuations from a constant deposition rate) is assumed to be 1 cm1 because such a value can guarantee that an overall agreement index of the model amounts to >60% (critical value). Based on of the age-depth model, the probability distributions of the modelled calendar ages for selected events related to the local palaeoenvironmental changes were calculated. 3.4. Dendrochronological methods Dendrochronological analyses were performed using standard

research procedures (Schweingruber, 1988). Prior to the measurements, 2 to 4 measurement grooves, typically 2e3 cm wide, were made with a dissecting knife on each of the samples along the radii. Samples prepared in this manner were then subjected to very precise measurements of the annual growth rings with the accuracy of 0.01 mm, using the Dendrolab 1.0 apparatus for dendrochronological measurements (Zielski and Kra˛ piec, 2004). The TREERINGS computer software (Krawczyk and Kra˛ piec, 1995), as well as the TSAP software (Rinn, 2005) were used to process the measured tree-ring sequences. The coefficient of the parallel run (Gl) and the t-coefficient were used (Eckstein and Bauch, 1969; Baillie and Pilcher, 1973) during the evaluation of the similarity of the dendrochronological sequences of ring-widths. A visual evaluation of the similarity between the dendrograms was used as the ultimate test for the indicated position adjustment. The visual evaluation of the parallel run between the dendrograms involved the application of logarithmically transformed raw curves. In the next stage, the accuracy of the measurements and the quality of the cross-dating were checked by means of the COFECHA program (Holmes, 1983). This program enables the efficient identification of missing rings and measurement errors. Chronologies were compiled based on the best correlated individual sequences using the TSAP software. Finally, the data were subjected to the standardisation procedures in order to minimize non-climatic ringwidth variations related to the age and geometry (Fritts, 1976). Taking into account highly probable extraordinarily narrow increments in the first and last decades of the tree growth, the dendrochronological sequences were standardized with a 67% flexible spline. The standardized chronologies were produced using the ARSTAN_4.1d program (Cook and Krusic, 2005).

3.5. Pollen, and non-pollen palynomorph analyses For pollen analyses, the cores were sampled at 5 cm intervals. Every sample of 1 cm3, was subjected to a standard chemical preparation: treatment in a 40% HF solution and acetolysis as described by Erdtman (1960), with a pre-treatment to remove calcium carbonate using a 10% HCl solution, and the removal of humus compounds by boiling in a10% KOH solution (Faegri et al., 1989). Pollen grains were identified using a light microscope (400 and 1000 magnification); up to 600 grains of trees and shrubs were counted for each sample. The percentage values of sporomorphs were calculated in proportion to the total sum, including

Table 1 Radiocarbon datings (14C) of samples taken from various profiles (logs) of the Puscizna Wielka raised bog. 14

No.

Samples' name/depth

Dated material

Lab. code

Age

1

Puscizna Wlk., PW1A; 385e387.5 cm

Peat

MKL - 581

6100 ± 90

2

Puscizna Wlk., PW1A; 387.5e390 cm

Peat

MKL - 582

6180 ± 90

3

Puscizna Wielka, PW 6; 365e369 cm

Peat

MKL - 1715

5070 ± 110

4

Puscizna Wielka, PW3, sample 3

Peat

MKL - 1716

4350 ± 70

5 6

Puscizna Wielka, PW3; sample 2 Puscizna Wielka, PW 7; 307.5e312.5 cm

Charcoal Peat

MKL - 1717 MKL - 1714

6570 ± 90 4740 ± 50

7

Puscizna Wielka, PW7; 310e312.5 cm

Charcoal

D-AMS 2239

5129 ± 33

8

Puscizna Wielka, artefact 368e370 cm

Wood fragment

Poze39936

4450 ± 40

C (BP)

Age cal. BP (2s) (95,4%) 7179e6745; 7240e7202 6815e6800; 7275e6846 6020e5591; 6111e6080; 6173e6156 5078e4824; 5136e5105; 5280e5163 7610e7309 5410e5325; 5588e5445 5829e5751; 5941e5857; 5982e5976 4935e4882; 5145e4956; 5288e5155

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Table 2 Radiocarbon datings (14C) of samples taken from various layers of the log PW - 2 of the Puscizna Wielka raised bog (position of log PW-2 on Fig. 2a). 14

C (BP)

No

Depth [cm]

Dated material

Lab. code

Age

1

25e27.5

peat

MKL - 1718

260 ± 70

2

72.5e75

Peat

MKL - 1720

450 ± 80

3 4 5

120e125 125e128 135e140

Peat Peat Peat

MKL - 1721 MKL - 1722 MKL - 1725

860 ± 70 950 ± 70 1120 ± 70

6

177.5e182

Peat

MKL - 1727

1620 ± 60

7 8 9

182e186.5 187.5e190 190e192.5

Peat Peat Peat

MKL - 1728 MKL - 1729 MKL - 1730

1760 ± 60 1880 ± 80 1900 ± 80

10

231.5e234

Peat

MKL - 1732

2190 ± 90

11

257.5e262.5

Peat

MKL - 1734

2350 ± 75

12 13 14

282e284.5 325.5e328 328e330.5

Peat Peat peat

MKL - 1736 MKL - 1737 MKL - 1738

2510 ± 70 3060 ± 100 3370 ± 60

15 16 17

362e364.5 365.5e368 368e370.5

Peat Peat peat

MKL - 1740 MKL - 1741 MKL - 1742

3990 ± 140 4430 ± 100 4450 ± 110

all tree and shrub taxa (AP) and herbaceous plants (NAP) and excluding the local plants, such as Cyperaceae, aquatic plants and spores (Berglund and Ralska-Jasiewiczowa, 1986). The results are presented as percentage pollen diagrams divided into local pollen assemblage zones (L PAZ) (according to Janczyk-Kopikowa, 1987), using the dendrogram constructed based on the ConsLink (Constrained Single Link of samples) analysis (Nalepka and Walanus, 2003). The data were stored and processed using the POLPAL software (Nalepka and Walanus, 2003). Apart from the quantity of pollen, four major types of nonpollen palynomorphs (NPP) were recognized and identified in the analysed sample material: stomata of plants, algae, fungi (hyphae and spores of fungal parasites, which live in association with host plants, commonly roots) and Rhizopoda (amoebae fauna) (Van  and Ha jek, 2006). Geel, 1978; Van Geel et al., 2003; Opravilova The NPP assemblages were analysed and results reported following the same procedure used for pollen analyses. 4. Results and interpretations 4.1. Sequence of deposits Within the area of the peat bog adjacent to the site of the occurrence of subfossil pine trunks, the thickness of the peat deposits does not exceed 4.0 m (logs: PW-6 and PW-2 in Fig. 2c). Organic sediments overlie clayey silt here (Mz-6.82 e 6.95 4), a very poorly sorted sediment (s1 ¼ 3.2e3.22 4) (Fig. 2c). Due to the granulometric composition, these sediments are typical for overbank deposits, characteristic of river marshes (of a crevasse-type sediments) (see: Kalicki, 1996). The mineral sediments are overlain by a thin layer of minerogenic peat reaching a fairly constant thickness of approx. 0.5 m within the peat bog (Fig. 2c). This is mainly woody peat, strongly decomposed. The minerogenic peat is

Age cal. BP (2s) (95,4% probability)

Modeled age cal BP (95,4% probability)

34 e … (7%); 115 e 73 (2,4%); 225 e 136 (19,4%); 498 e 254 (66,5%) 562 e 309 (88,8%); 635 e 595 (6,6%) 919e682 980e705 1186 e 921 (92,6%) 1238 e 1206 (2,8%) 1629 e1378 (92,4%); 1660e1655 (0,7%); 1693e1666 (3,0%); 1823e1549 1995e1615 1676 e 1620 (4,4%); 2006e1686 (90,3%); 2038e2024 (0,7%) 1961e1950 (0,6%); 2355e 1969 (94,8%) 2260e 2158 (11,5%); 2620e 2299 (72,2%); 2705e 2628 (11,7%) 2748e2379 3470e2965 3729e 3456 (90,5%); 3765 e 3748 (1,5%); 3823 e 3792 (3,4%) 4839e4089 5315e4840 5326e 4835 (93,3%); 5446 e 5404 (2,1%)

23 e (3)

517e433 914e864 952e899 1080e1002 1609e1523

1738e1622 1739e1623 1762e1640

2218e2110 2493e2400

2750e2678 3379e3221 3532e3409

4846e4733 4992e4855 5192e4951

overlain by a homogenous complex of ombrogenic peat, predominantly sphagnum peat (Eusphagneti type according to Tołpa et al., 1971 classification), with a tihickness of 3.5 to 2.5 m (Fig. 2c). Locally, irregular inserts of sphagnum-cottongrass peat (EriophoroSphagneti type) of various thickness occur (Fig. 2c). A similar irregularity characterizes the horizons of peat decay observed in the profiles as layers of decomposed sphagnum peat (Fig. 2c). The charcoals occurring in the mineral sediments underlying the peat bog (clayey silt) were dated by the 14C method at 7610e7309 cal BP (PW-3 core in Fig. 2a; Table 1) and at 5982e5751 cal BP (Fig. 2c, Table 1). The beginning of the organic accumulation in the peat bog area, dated by the radiocarbon method was quite synchronous with the Early Subboreal (ca 5.6e4.8 ka cal BP) (Fig. 2c). However, single dates of the bottom sediment of the peat bog collected at various parts are older (ca 7275e6800 and 6173e5591 cal BP - Fig. 2c; Table 1). In a profile collected in the central parts of the exploitation field (and thus representative of the tree trunks excavated during the exploitation), a minerogenic peat sample collected from the bottom of the peat bog (368e370.5 cm) was dated at 4450 ± 110 BP (5446e5404; 5326e4835 cal BP) (Fig. 2c e log PW2). From the same horizon of peat (368e370 cm), during the drilling with the INSTORF drill, a sharpened fragment of a wooden rod with a length of approx. 8 cm and a diameter of 1.5e2 cm was extracted (Fig. 2c e see photo below). The analysis of the object performed by prof. Paweł Valde-Nowak (Jagiellonian University in Krakow) revealed that it was an artefact made by a prehistoric man. The dendrological analysis showed that the artefact had been made from the wood of a Scots pine (P.sylvestris L.). It was dated using the AMS technique at 4450 ± 40 BP (5288e5155; 5145e4956; 4935e4882 cal BP) (Table 2). The age of the artefact corresponds to the period of the Funnel Beaker Culture or the Baden Culture settlements, which were widespread at that time in

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Fig. 3. Age-depth curve of the Puscizna Wielka peat bog, with time-range of subfossil trees (dated by 14C and dendrochronologically). Part of the curve concerning the minerogenic stage of the mire is marked in blue, the ombrogenic peat bog e in red. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

the Orawa-Nowy Targ Basin (Rydlewski and Valde-Nowak, 1979; Valde-Nowak, 1988). A pollen analysis performed for one of the profiles indicates that the bottom sections of this part of the peat bog (still minerogenic, fen-type at that time) began to grow in the Subboreal, which is confirmed by the radiocarbon datings from this profile (Figs. 2 and 6c,a). The change of the feeding of the peat bog from soligenic and fluviogenic into ombrogenic and the beginning of the accumulation

of ombrogenic peat, were dated in the main profile (at the depth of ca 328e330 cm) to 3370 ± 60 BP (3729e3456; 3765e3748; 3823e3792 cal BP) (Fig. 2c e log PW-2). The loss on ignition analysis of the sediments of three profiles indicates that the peat (both minerogenic and ombrogenic) of the Puscizna Wielka peat bog are characterised by a significant organic content amounting on average to 95% of loss on ignition (Fig. 2c). With some exceptions, fluctuations within the loss on ignition

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curves are insignificant and fall within the range of the standard error (4e5% e see Heiri et al., 2001). Only in the upper sections of all three of the profiles (uppermost, 20e25 cm, section) a drop in the loss on ignition value amounting to approx. 12e15% is observed. It probably related to the exploitation of the surface layer of the peat and the natural regeneration of the upper parts of the peat bog (Fig. 2c). The beginning of regression on the loss on ignition curve in the upper sections of the profile was dated at 260 ± 70 BP (the last 498 calendar years). However, regeneration of the upper layer of the peat deposit might have occurred already under conditions of human influence caused by the deforestation of considerable parts of the alluvial plains of the rivers in the Orawa Basin, aeolisation of the silty deposits in the exposed areas, and the delivery of fine mineral particles to the upper layers of the recovering peat dome (see Gurnell et al., 2008). 4.2. Dendrochronological analysis The age distribution of the analysed trees is presented on Fig. 4. Over 70% of the investigated trees from the Puscizna Wielka peat bog were at least 100 years old. Of these, 5% were pines more than 250 years old, the oldest having reached the age of 291 years. Out of the 614 individual sequences containing over 60 annual growth rings 440 ones were synchronised (correlated) dendrochronologically. Based on same-aged, correlated dendrochronological sequences, 5 chronologies were produced: 2NTT_AA2, 2NTT_BC2, 2NTT_D, 2NTT_E and 2NTT_E, determined by at least 10 trees each (Table 3). The longest chronology compiled, spanning 627 years, was developed from 262 samples, while the next in order, 2NTT_AA2, comprising 491 years e from 125 samples (Fig. 5). Altogether the chronologies covered 1751 calendar years. However, it could be a few dozen years less, taking into account possible slight overlap of some of the floating patterns, but too short to merge them. Due to the lack of sufficiently long pine chronologies from this region (SzychowskaeKra˛ piec, 2010) and also lack of convergence with peatland sequences from more distant areas of Germany and Sweden, the dating of the chronologies compiled was carried out using the wiggle-matching method. The chronologies and the results of the wigglematching dating were presented in detail in the paper by Kra˛ piec and Szychowska-Kra˛ piec (2016). The pine chronologies of

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the Puscizna Wielka and the results of the wiggle-matching dating are presented in Table 3. Older generations of pine trees (2NTT-D, E and F) are characterised by the occurrence of germination phases (G) lasting for 20e50 years as well as tree dying-off phases (DOE) lasting for 120e150 years (Fig. 5). The pine trees defining the chronologies represented by lower numbers of trees (2NTT_F and 2NTT_E) are characterised by the occurrence of numerous zones of reduced annual increments, which seems to indicate significantly worse growth conditions for trees in the periods 5415e5245 cal BP (2NTT_F) and 4395e4150 cal BP (chronology 2NTT_E). This concerns also, to a lesser extent, trees from the period 4160e3940 cal BP (Table 3). Among the older generations of pine trees, an exception are those originating from the maximum afforestation phase of the peat bog, determining the pattern 2NTT_BC2 (5155e4525 cal BP). Here, unlike in the case of the other generations, the dying off of trees has continued simultaneously since 5005e4525 cal BP, but three germination phases can be distinguished, dated to circa 5150 cal BP; 5020e4950 cal BP; 4920e4820 cal BP and 4790e4740 cal BP) (Fig. 5). The lack of distinct germination and dying-off phases (GDO) may attest to relatively stable growth conditions continuing for several hundred years, especially that the feature distinguishing the trees from this generation is a considerable proportion of very old pine trees, over 150 years old. Pine trees from the youngest generations, determining the sequence 2NTT_AA2, show a clear division into two generations separated by a clear GDO (Fig. 5). The older pine trees began to encroach on the peat bog around 3050 cal BP. It was a process spread over a long period of time. Beginning from about 2870 cal BP, the pine tree dying-off phase from this generation began and continued until about 2790 cal BP. Because several pine trees had survived and were still growing when new trees encroached upon the then almost forestless part of the peat bog, the relative dating of both groups was possible. The time of germination by the new generation of trees was relatively short. The majority of the trees appeared within about 30 years. The tree dying-off phase is also clearly marked, lasting almost 100 years, from about 2660 to 2560 cal BP. In both of the periods of dying off of pine trees, a clear reduction of tree-ring width can be observed, suggesting adverse conditions for tree growth.

Fig. 4. Histogram of ages of the trees analysed dendrochronologically.

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Table 3 Sequences of subfossil pine trees from the Puscizna Wielka peat bog. Sequence dating was made with the WM technique (2s range calibration). No

Chronology

Years

No of sequences

Dating (cal BC)

Dating (cal BP)

1 2 3 4 5

2NTT F 2NTTBC2 2NTT_E 2NTT D 2NTT_AA2

168 627 245 220 491

10 262 10 33 125

3465e3295 (±25) 3205e2575 (±25) 2445e2200 (±50) 2210e1990 (±40) 1100e610 (±10)

5415e5245 5155e4525 4395e4150 4160e3940 3050e2560

(±25) (±25) (±50) (±40) (±10)

Fig. 5. The subfossil pine chronologies from the Puscizna Wielka bog. (a) The horizontal lines represent individual trees sorted by ending years. (b) Tree-germination and dying-off phases, shown in green and red respectively. (c) Sample replication. (d) Mean age of samples. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

4.3. Vegetation and palaeoecological changes in the Puscizna Wielka raised bog: pollen and non-pollen palynomorph analyses The history of the flora of the Puscizna Wielka peat bog (and its surroundings) is presented chronologically, appropriately for each of the L PAZs distinguished on the pollen profile (Fig. 6a). Palaeoecological changes taking place in the peat bog during the duration of each local pollen zones were additionally characterised

based on supplemental analyses of non-pollen palynomorphs (NPP) (Fig. 6b). Picea L PAZ (382.0e349.5 cm). This zone, dated with radiocarbon to between 5404 and 5326 cal BP and 4485e4009 mod. cal. BP, is characterised by the dominance of Picea abies. A considerable proportion of Alnus and Corylus avellana pollen, as well as a relatively large percentage of Quercus pollen also occur within this level (Fig. 6a). Small share of Tilia as well as some Ulmus. Carpinus, Fagus,

M. Kra˛ piec et al. / Quaternary Science Reviews 148 (2016) 192e208 Fig. 6. The results of pollen analyses (a) and non pollen palynomorph (NPP) analyses (b) of the Puscizna Wielka peat bog profile, with modeled radiocarbon datings of the chronozone boundaries (see Fig. 3). Location of the profile PW-2 on Fig. 2a).

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Abies, Betula and Pinus pollen have also been noted. Herbaceous vegetation was represented by Artemisia, Rumex, Plantago, Poaceae, Apiaceae and local reed bed plants e Typha latifolia and Sparganium t., as well as Cyperaceae (Fig. 6a). A number of NPPs of various single types appear throughout all levels of this section of the profile. Such variable composition of the NPP assemblages may have resulted from the variability and instability of the paleoecological conditions within the peat bog that persisted during the initial phase of its formation (Fig. 6b). In the bottom layer, levels with only Gelasinospora (type 1) exist e a sporophyte considered to be indicative of fires and desiccations due to increased sporulation in burnt areas (Pirozynski, 1989; Kuhry, 1997; Muller et al., 2008). This sporophyte is typically found on dead wood and in the faeces of herbivores (Lundqvist, 1972; Krug et al., 2004). Another saprophyte e Helicoon pluriseptatum (type 30) occurs within two levels (362 cm and 367 cm) together with Endophlyctis lobata (type 13) e a parasite of sphagnum mosses, and Arcella flavum (type 31). Tilletia sphagni (type 27) e another parasite of sphagnum mosses, is also found within these two levels. Abies L PAZ (349.5e279.5 cm). This zone was dated to between 4485 and 4009 mod. cal. BP and 2745e2560 mod. cal. BP (see Fig. 3) and was established as a zone with dominant Abies alba and Fagus sylvatica within the pollen spectrum (Fig. 6a). A decrease in the percentage of Picea abies pollen is clearly visible in this zone. Some other trees are present here, including pine and birch. Herbaceous plants are represented by Artemisia, Poaceae, Asteraceae, Brassicaceae, Rubiaceae ¼ Galium t. and Rosaceae (Fig. 6a). A predominance of the Arcella flavum (type 31) palynomorphs is observed within NPP assemblages analysed within this section of the profile (Fig. 6b). A large proportion of the Habrotrocha angusticollis and Arcella flavum (type 31) suggests presence of a high water level in the peat bog (see Lamentowicz and Mitchell, 2005; Payne et al., 2012). Sections of the profile where the hydrological conditions might have been less stable are documented based on decreases in the Arcella flavum curve and an increase of the Assulina seminulum (type 32B), a taxon known to have a higher tolerance for unstable aquatic conditions (Fig. 6b) (see Payne et al., 2012). Abies-Fagus-Alnus L PAZ (279.5e99.5 cm). In this zone, which formed between 2745 and 2560 mod. cal. BP and 764e586 mod. cal. BP, a predominance of Abies alba and Fagus sylvatica pollen is notable, with a large proportion of Alnus and small percentage of the Picea abies pollen and some other tree species, such as maple, elm, oak, hornbeam, ash, willow, pine and birch. Herbaceous plants were still poorly represented, mostly by Artemisia and Poaceae. The dominance of the Arcella flavum (type 31), with a very large share of Habrotrocha angusticollis indicates presence of a high level of water table in the peat bog (Lamentowicz and Mitchell, 2005; Payne et al., 2012). Moreover, the intermittent occurrence of Microthyrium sp. (type 8), Tilletia sphagni (type 27) and the type 18/ type 19 palynomorphs, as well as a large share of Endophlyctis lobata (type 13) all indicate that the paleoecological conditions within the peat bog were quite variable. Gelasinospora sp. (type 1) may suggest periodic desiccations or occurrence of fires (Pirozynski, 1989; Kuhry, 1997; Muller et al., 2008). In the levels showing almost complete absence of testate amoebae, species otherwise common for raised peat bogs, a very high percentage of Endophlyctis lobata (type 13) was noted. This suggests that periodic desiccations of the peat bog and changes in the conditions for the development of sphagnum mosses might have occurred (Fig. 6b). Abies-Fagus L PAZ (99.5e28.75 cm; 764e586 mod. cal. BP) is quite similar in pollen composition to the previous zone. As the percentage of Alnus pollen is decreased here, the Ericaceae, mainly Calluna vulgaris are more common and there is great increase in the number of Sphagnum spores in pollen spectrum. The predominance of the Arcella flavum (type 31) within this

level points again to a high water table level in the peat bog (see Payne et al., 2012). The diagram shows several single episodes associated with hydrological disturbances when Arcella flavum decreased in favour of Assulina seminulum (type 32B) (Fig. 6b). Pinus-NAP L PAZ (28.7e6.75 cm). Pinus and NAP (mainly heather) pollen clearly predominate within this level, with a large share of Drosera intermedia and Bryales spores. As shown in the diagram (see Fig. 6a), the occurrence of relatively larger shares of Poaceae and Cyperaceae are easily noticeable. A predominance amongst the NPP assemblages within this levelof Endophlyctis lobata (type 13) e a parasite typical in sphagnum mosses (Van Geel, 1978) is also noted (Fig. 6b). Its large share of the NPP spectrum may signify the occurrence of some specific conditions favourable for the development of sphagnum mosses. The almost total absence of testate amoebae, otherwise common in raised peat bog environments, points to a desiccation of the peat bog, probably caused by improvement of the water drainage and on-going peat exploitation. The presence of Gelasinospora sp. (type 1) may indicate systematicdesiccations or periodic occurrences of fire (Pirozynski, 1989; Kuhry, 1997; Muller et al., 2008; Feeser and O'Connel, 2009). Pinus L PAZ (6.75e0 cm). The pollen spectrum within this level is absolutely dominated by Pinus. Due to this extremely large share of pine pollen no conclusions regarding the state of regional vegetation or possible presence of other species can be made. There is also great increase of Ericaceae pollen (Fig. 6a). A predominance of Assulina seminulum (type 32B) e taxon typically highly tolerant and adaptive to less stable aquatic conditions (Payne et al., 2012) is easily noticeable within the NPP assemblage. The absence of other testate amoebae, including Arcella flavum (type 31), water table level dependent and usually common in raised peat bogs (Lamentowicz and Mitchell, 2005; Payne et al., 2012), may indicate a desiccation of the peat bog and/or instable hydrological conditions such as fluctuations in the water level in this part of the profile. 5. Discussion The results of the dendrochronological analyses and wigglematch dating of pines from the Puscizna Wielka peat bog indicate that the overwhelming majority of the trees studied grew in the peat bog from ca 5415 to ca 3940 cal BP with two breaks dated to 5245e5155 cal BP, and 4525e4395 cal BP (Figs. 5 and 7). The invasion of the trees on the peat bog took place relatively early, during the initial phase of the peat bog development. Such conclusion is supported by radiocarbon dating of charcoal occurring in the mineral deposits (silty clay) directly underlying the peat (ca 5976e5751cal BP), as well as the data concerning main stage of minerogenic peat accumulation (ca 5400e4800 cal BP) (Figs. 2 and 7). The first stage of the growth of trees in the vicinity of the peat bog thus coincided with the beginning of the development of numerous, small minerogenic mires, that gradually began to fill isolated abandoned palaeochannels of the Czarny Dunajec and the Orava Rivers (Baumgart-Kotarba, 1991-1992). Undoubtedly, just prior to the formation of the peat bog, some critical hydrological events, typical of strong climate moistening, must have occurred, a major change in the channel system of the rivers of the Orawa-Nowy Targ Basin from multiple-channel to single-channel and frequent flooding of the rivers resulting in depositions of crevasse sediments typical of marshes (silty clay) on the gravel-filled river banks. These sediments, which contain charcoal dated at 7610e7309 cal BP and 5982e5751 cal BP (Table 1, see also Fig. 7), as well as the oldest peat (dated at 7275e6800 and 6173e5591 cal BP) deposited within the isolated abandoned palaeochannels, were deposited during periods of severe

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Fig. 7. Correlation of the palaeoenvironmental changes recorded in the Puscizna Wielka raised peat deposits (on the basis of pollen and non-pollen analyses) and pine palaeoecology (on the basis of dendrochronological analysis of the Puscizna Wielka subfossil trees). Time intervals of pine tree growth on European peatlands after various authors (peatland locations and their numbers as well as related authors quoted in Fig. 1). Palaeoclimatic changes are presented as a concentration of radiocarbon datings (Cumulative Probability Density Function) of mass movements and related processes, as well as fluvial processes in Poland (after Margielewski, 2006; Starkel et al., 2013). The time series of the frequencies of the Holocene cold and wet events in the Northern Hemispheree after Wanner et al. (2015). A positive or negative anomaly of temperature or humidity presented in ±standard deviation of the Holocene mean value-see also Wanner et al. (2011). Cold events culmination after Bond et al., 1997, 2001; Wanner et al., 2011. Holocene chronostratigraphy after Walker et al., 2012, and traditional (but not formal) Holocene subdivision after Mangerud et al. (1974), chronozones boundaries calibrated by Walanus and Nalepka (2010).

moistening and cooling of the climate that occurred close to the end of the Atlantic and in the Early Subboreal. Palaeoclimatic analyses have shown that in ca 7700 cal BP, as well as in 6400e5600 cal BP, episodes of particularly severe climate humidification took place in the whole of the Northern Hemisphere (Fig. 7) (Wanner et al., 2011, 2015), including Central Europe (Starkel et al., 2013). The increase in climate humidity, especially in the more recent periods of humidification, caused surge in both the fluvial activity of the Upper Vistula River (Starkel et al., 1996, 2006) and of mass movement processes in the Western Carpathians (Fig. 7) (Margielewski, 2006; nek et al., 2013). At around the same time Starkel et al., 2013; Pa (6500e5900 cal BP), a widespread climate cooling occurred (Wanner et al., 2011, 2015), causing the advance of Alpine Glaciers (Rotmoos/Piora Stage, Bortenschlager, 1982). The beginning of the main stage of the peat accumulation is generally synchronous in all of the studied peat bog profiles (ca 5446e4835; ca 5588e5325 cal BP; 5280e4824 cal BP) (Fig. 2c, Table 1). Just before the main phase of covering of all palaeochannels with peat began, the bog area was briefly populated by a

relatively small population of pine trees during 5435e5245 cal BP (Fig. 5, sequence 2NTT_F). This short-lived invasion of the trees over the peat bog occurred during the time of lower water table, incision of the rivers in the Orawa - Nowy Targ Basin into their alluvia and gradual warming and drying of the climate. The climatic changes caused general attenuation of fluvial activity of the rivers of southern Poland, significantly increasing in the rate of mineral deposition in peatlands after ca 5.6 ka cal BP (Starkel et al., 2013). The trees growing at that time in the Puscizna Wielka raised bog show numerous zones of reduced annual tree ring width, most likely indicating that the growth conditions for trees deteriorated significantly during the period 5420e5230 cal BP (sequence 2NTT_F e see Fig. 5) leading to the pine trees dying-off phase (5330e5245 cal BP) (Figs. 5 and 7). The results of the analyses of non-pollen palynomorphs from the main profile of the Puscizna peat bog demonstrated that a high and stable water level in the peat bog persisted at that time, and might have been the direct cause for this pine trees dying-off episode (Figs. 6 and 7b). The simultaneous drop in the number of pollen of the thermophilous

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Corylus, Quercus and Ulmus in the Puscizna bog profile (see Fig. 6a) also suggests that deterioration of the ecological conditions occurred in the vicinity of the peat bog. These local changes in climatic conditions might have been related to the more widespread climate cooling that occurred in Central Europe at ca 5500e5250 cal BP (Fig. 7) (Magny, 2004; Pe˛ dziszewska and Latałowa, 2016; Wanner et al., 2015). The second phase of germination of the peat bog by pine trees was a long-lasting (it continued for 630 years) and was characterised by the mass occurrence of pine in the peat bog (Fig. 5 e sequence 2NTT_BC2). The maximum afforestation phase of the peat bog took place in the period 5160e4530 cal BP (Figs. 5 and 7). Our data indicate the occurrence of 3 distinct germination phases during this stage: circa 5150 cal BP, 5000e4950 cal BP and 4790e4740 cal BP (Figs. 5 and 7). However, the trees were dying off within the Puscizna Wielka peat bog simultaneously during almost this entire period: from 5000 to 4530 cal BP (Figs. 5 and 7). The occurrences of several other pine tree bogs in the same period, have been documented throughout Europe. These include Campemoor and Dreiecksmoor peat bogs in Northern Germany (Eckstein et al., 2009, 2010), with clearly marked GDOs and age ranges shorter than that of the Puscizna Wielka (see Fig. 7), bogs in the British Isles (Lageard et al., 1999; Moir et al., 2010) and Ireland. According to Pilcher et al. (1995), although the growth of the pine trees on the Irish peat bogs began at ca 5400 cal BP (so it was simultaneous with the invasion of trees on the Puscizna Wielka bog), their chronologies are longer than those of the trees covering the Puscizna Wielka bog (see Fig. 7). As recorded in non-pollen palynomorph assemblages, the first germination phases of trees of this generation in the Puscizna Wielka peat bog (first germination phases of 2NTT_BC2 sequence) corresponded to the lowering of the water table level and the decline of the high water level in the peat bog (Figs. 6 and 7b). The successive dying-off of trees over quite a long period was likely a natural process related to the advanced age of the trees, perhaps slightly combined with the harsh climatic conditions typical for the Orawa - Nowy Targ Basin at that time. The termination of bog pine growth however, might have strongly influenced by the climate cooling as well as the increase in humidity that occurred in the Northern Hemisphere around 4800e4500 cal BP and 5100e4500 cal BP, respectively (Fig. 7) (Wanner et al., 2011, 2015). The general climate cooling at that time caused an expansion of the Alpine (Rotmoos 2 phase) (Bortenschlager, 1982) and Scandinavian glaciers (Karlen and Kuylenstierna, 1996). Ca 4850 cal BP, an increase in both humidity and the cooling of the climate was recorded in Poland, intensifying fluvial activity and mass movements in the Western Carpathians (Margielewski, 2006; Starkel et al., 2013; nek et al., 2013). Considerable increase in occurrences of the Pa minerogenic horizons (sensitive indicators of an increase in humidity and cooling) in the landslide peat bogs of the Polish Carpathians, was recorded in the period 4750e4550 cal BP (see Fig. 7)  ski et al., (Margielewski, 2006; Margielewski et al., 2011; Michczyn 2013). Accordingly, the germination phase of trees of this generation might have taken place in the course of a significant increase in climate humidity (Fig. 7). The distinctive feature of trees from this generation is a significant proportion of aged pine trees, over 150 years old (Fig. 5). The second phase of peat bog afforestation ended probably as a result of a specific ecological disaster, possibly caused by Neolithic man. A distinct fire horizon, which likely ended this phase of mass germination of the peat bog by pine trees, was recognized in the sediments of the minerogenic mire within the peat dated at ca 4.4e4.5 ka cal BP. Tree germination continued soon after, with two more phases of tree growth during the periods 4350e4210 cal BP (sequence 2NTT_E) and 4180e3960 cal BP (sequence 2NTT_D)

(Figs. 5 and 7). Timings of these sequences correspond well to germinations time intervals reported from peat bogs in Northern Germany (4350e3950 cal BP) (Eckstein et al., 2009, 2010) and Ireland (4450e3950 cal BP) (Pilcher et al., 1995) (Fig. 7). Both of these phases were typical germination dying-off phases (GDOs). They are characterised by 20e50 years of germination at the onset of each phase followed by dying-off periods lasting 120e150 years (Figs. 5 and 7). The number of trees occupying the Puscizna Wielka raised bog at that time was however much smaller than in the previous phase (Fig. 5). Furthermore, these trees are characterised by the occurrence of numerous zones of reduced annual tree rings. This characteristic indicate that the tree growing conditions, most notably during 4350e4210 cal BP (sequence 2NTT_E) and to a lesser extent during 4180e3960 cal BP (sequence 2NTT_D) (Fig. 5), were noticeably worse compared to those of the previous phase of germination of the peat bog. In general, climatic conditions were quite stable during both these phases of tree growth, with reduced humidity (period: 4850e3700 cal BP e see Fig. 7). Extreme geomorphological processes or hydrological phenomena were considerably less frequent in that period (Fig. 7) (Starkel et al., 2013). The cold Bond event reportedly occurred ca 4.2 ka cal BP (Bond et al., 1997; Birks, 2008), although more recent studies place the events of climate cooling at ca 4.7 and 3.8 ka cal BP (Fig. 7) (Wanner et al., 2011, 2015). Admittedly, in this part of the pollen profile of the Puscizna Wielka bog, a gradual decrease in the number of Picea pollen and a quite rapid expansion of Abies alba and (to a lesser degree) Ulmus are observed (Fig. 6a) The curves for the thermophilous Corylus, Quercus, Tilia do not display any declines, indicating fairly stable thermal conditions. At the same time, a change in the feeding of the peat bog, from soligenic (fluviogenic) to ombrogenic, and the beginning of ombrogenic peat accumulation occurred (sensu Kaule € ttlich, 1976). These events were likely associated with hyand Go drological changes in the vicinity of the peat bog, probably caused by a lowering of the water table level due to incision of the rivers flowing throughout the Basin into the bedrock. The desiccations of the peat bogs is shown by the level of peat humification related to the commencement of the development of the ombrogenic stage of the peat bog (Fig. 2c). Sundew (Drosera) pollen, a species typical of ombrogenic peat bogs, co-appears with sphagnum in the peat bog and there is a notable increase in the number of Pinus sylvestris t. pollen in the vicinity of the peat bog (Fig. 6a). A sustained decrease in the quantity of the pollen of the hygrophilous Alnus is observed while a significant permanent decrease in the number of the thermophilous Corylus avellana indicates the tendency towards climate cooling at that time (Fig. 6a). The termination of this phase of germination was followed by the second forestless period of the peat bog development which lasted for almost 900 years. It began after 3960 cal BP and continued until 3050 cal BP. This event can be also recognized in other European peat bogs. After 3950 cal BP, the number of pine trees in the peat bogs in Northwestern Europe is clearly on the decrease: only few pine sequences were still in existence including one only longer pine sequence described from that time from the peat bogs of southern Sweden (Edvardsson, 2010; Edvardsson et al., 2012), and two shorter from Lithuania (Edvardsson et al., 2016) (Fig. 7). This decrease probably occurred in response to the climate change towards cooler climatic conditions (Nesje et al., 1991; Holzhauser et al., 2005), likely associated with the more global change of atmospheric circulation in the region of North Atlantic occurred that led to a general decrease of summer temperature and an increase of precipitation (Nesje et al., 1991; Hammarlund et al., 2004; Mayewski et al., 2004; Holzhauser et al., 2005; De Jong et al., 2009). Data from the Northern Hemisphere show two separate cold events clustering around 3.8 and 3.1 ka cal BP associated

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with the climate humidity increase (Fig. 7) (Wanner et al., 2008, 2015). This cooling trend caused advancement of the Scandinavian (Karlen and Kuylenstierna, 1996) and Alpine glaciers (phase Loeben) (Bortenschlager, 1982) at that time. During the next and most recent phase of germination, which began at circa 3 ka cal BP, pine trees returned to the Puscizna Wielka bog (wich was just of ombrogenic type at that time) in much greater number as compared with the previous phase. This germination phase continued for another 500 years (Figs. 5 and 7) and was preceded by a fire in the peat bog which, based on data collected form a fire horizon found in the sediments (that also contain Gelasinospora type 1 sporophyte), occurred ca 3.3e3.4 ka cal BP, hence predating the appearance of trees by about 100e200 years. The pine trees appearing during this most recent 2NTT_AA2 phase can be easily subdivided into two generations separated by clear germination and dying-off phases (GDOs) (Figs. 5 and 7). The older pine trees began to encroach on the peat bog at ca 3080 cal BP. It was a process spread over a long period of time. The stages of germination and dying off of trees during the course of this stage of germination are well correlated with the hydrological changes recorded in the peat bog (Figs. 6 and 7). The first two stages of germination (3025e2965 cal BP and 2925e2890 cal BP) took place between periods of a high level of water table in the peat bog. The increase of the water level in the peat bog prior to the appearance of pine trees at ca 3100e3150 cal BP (Fig. 7) was indicated by composition of the non-pollen palynomorph assemblages, as well as the appearance of pollen of Lemna and Typha latifolia in the palynological profile (Fig. 6a and b). Another long-term flooding of the peat bog occurred at ca 2900e2600 cal. BP triggering deterioration of ecological conditions for trees overgrowing the mire. Dying-off phase of the pine trees from this generation started at ca 2920 cal BP and lasted until ca 2830 cal BP (Figs. 5 and 7). That event, definitely associated with high water level in the peat bog, caused by a severe and rapid increase in climate humidity in the Northern Hemisphere ca 3.0e2.8 ka cal BP (Fig. 7) (Wanner et al., 2011, 2015), has also been recognized elsewhere in Poland (e.g. Margielewski, 2006; Starkel et al., 2013; Pe˛ dziszewska and Latałowa, 2016). Remarkably, some pine trees had survived this stage of dying off and were still growing within the Puscizna Wielka bog when new trees encroached upon then almost forestless part of the peat bog. Hence, the relative dating of both groups was possible. The majority of the later-generation trees appeared within about 30 years (2775e2720 cal BP) when a short-term improvement of the ecological conditions for tree growth in the peat bog probably took place (local drops of the water table?). Another dying-off phase, lasting for almost 100 years, ca 2690e2590 cal BP, was related to the re-intensification of the adverse climatic conditions at that time. The severe cooling of the so-called “Cold Iron Age Epoch” was recorded at ca 2850e2250 cal BP (Bond et al., 1997, 2001; Birks, 2008) or, according to Wanner et al. (2011, 2015), around 3300e2500 cal BP with culmination ca 2.7 ka cal BP (Fig. 7). In the Alps, an expansion of the mountain glaciers of the Goeschener 1 phase was taking place at that time (Bortenschlager, 1982). Climate cooling and moistening caused intensification of slope processes and fluvial activity in Poland (with culmination at ca 2850 cal BP) (RalskaJasiewiczowa and Starkel, 1988; Margielewski, 2006; Margielewski et al., 2010; Starkel et al., 2013). It is symptomatic that during the last dying off phase, larch (Larix) pollen occur in of the Puscizna Wielka peat bog, as evident from the palynological diagram (Fig. 6a). This is the pioneering tree typical of the alpine tree-line, showing a high tolerance to thermal and humid conditions that began to spread in Poland during in the More Recent Dryas Phase and continued until the Preboreal Phase (Ralska-

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Jasiewiczowa et al., 2004). The adverse climate prevailing in both of the pine trees dyingoff phases is also indicated by notable reduction of the tree ring widths, a phenomenon typically resulting from adverse conditions for the growth of pine trees (Fig. 5). These two dying-off episodes are well known from other regions of Central Europe although within the bog pine populations only the more recent phase was documented in pines occurring in Scotland (ca 2760 cal BP e Moir et al., 2010) and in the peat bogs of Lithuania (ca 2800 cal BP) (Edvardsson et al., 2016) (Fig. 7). According to available data, no bog pines developed in Sweden and Northern Germany after 3050 cal BP (Fig. 7). Long-term reductions of tree rings width and dying off of oak trees were noted during 2805e2785 cal BP, 2685e2665 cal BP and 2655e2635 cal BP in the peat bogs of Lower Saxony (Fig. 7; site location e see Fig 1a, site-1). Also, generation changes of trees growing in peatlands ca 2800 cal BP and 2650 cal BP (Leuschner, 1992) were reported from the North Sea coast. Similar changes were recognized in the Rucianka peat bog (NE Poland) (site location: Fig. 1aesite 7) (Barniak et al., 2014). The more recent dying-off episode can be associated with the climatic changes that occurred in Northwestern Germany as noted by Schmidt (1992), who discovered well-preserved track-ways built on wetlands dated to the period 2667e2663 cal BP. It appears that these track-ways, used only for short period of time, were buried in a water-saturated layer and quickly covered by peat resulting in almost perfect preservation of wood. A similar event took place in the case of the settlement of the people of the Lusatian culture settlement in Biskupin (central Poland) abandoned after 2670 cal BP by the population living there because the water surface in the nearby Lake Biskupin rose to a level that made further settlement impossible (Niewiarowski et al., 1992). The last dying-off phase lasted for about 100 years (2690e2590 cal BP) DOE and was followed by about 2600 years of forestless period in the Puscizna Wielka peat bog (Figs. 5 and 7). Although relatively long in duration, this forestless period in the Puscizna Wielka, is much shorter than those established in Northern Germany and Sweden where bog pines had already disappeared around 3050 cal BP, and slightly shorter than in Scotland where the decline of the pine trees took place after 2760 cal BP (Fig. 7) (Moir et al., 2010). Elswhere in Northeastern Europe, in places such as in Lithuania, trees are known to have grown in peat bogs over the period of the last 2500 years with the exceptions of 9the10th centuries AD and the Little Ice Age (Fig. 1a e site 6, and , 2001; Edvardsson et al., 2016). Fig. 7) (Pukiene Most recently, pine trees have returned to the Puscizna Wielka peat bog (Fig. 2b). This has been caused by the lowering of the water table in the peat bog associated with its drainage and the exploitation activity. The desiccation of the peat bog is well recorded by the change in the composition of pollen and non-pollen palynomorph assemblages (Fig. 6). Pinus sylvestris is accompanied  gger by Pinus mugo and a rare species Pinus x rhaetica Bru (Staszkiewicz and Tyszkiewicz, 1969). 6. Conclusions Our study carried out on site at the Puscizna Wielka raised bog indicate that pine tree (Pinus sylvestris) which grew on this peat bog during the period ca 5415e3940 cal BP (with several germination and dying-off phases) and ca 3050e2560 cal BP, is a species very sensitive to fluctuations of climatic conditions closely related to changes in the hydrology and thermal regime occurring in the peatlands. Within the peat bogs of the Orawa-Nowy Targ Basin, which occur in the area that has repeatedly experienced particularly adverse climatic conditions (so-called “cold pole” of Poland, temperature inversions, prevalence of precipitation over

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evaporation), the reaction of pine trees to the changes in ecological conditions (climate cooling, increase in humidification resulting in higher water level) is particularly noticable. Our analyses have demonstrated that the pine trees dying-off phases were undoubtedly caused by rising water levels in the peat bog closely associated with the increase in climate humidity. These events are perfectly reflected in the composition of pollen and non-pollen palynomorph assemblages occurring throughout the peat bog profile. Unfavourable climatic and, consequently, hydrological conditions resulted in significant reduction of the tree rings width, clearly marking periods of deterioration of the paleoecological conditions within the peat bog environment. Moreover, it is likely that the generally adverse climatic conditions of the Orawa - Nowy Targ Basin accelerated the pine trees dying-off and the reduction of their tree rings width. In contrast, the phases of germination and expansion of the trees and shrubs over the area of the peat bogs have been associated with the drying and occasional warming of the climate. These conclusions are consistent with the results of similar dendrochronological and palynological studies carried out elswhere in Europe (Edvardsson et al., 2012, 2016; Eckstein et al., 2010; Leuschner et al., 2007). Acknowledgements The studies presented above have been funded by the Ministry of Science and High Education (NCN grant) No: N N307 774 340, conducted in period: 2011e2014. The Authors would like to thank two anonymous Reviewers for their valuable remarks and comments concerning the manuscript. Appendix A. Supplementary data Supplementary data related to this article can be found at http:// dx.doi.org/10.1016/j.quascirev.2016.07.022. References Baillie, M.G.L., Pilcher, J.R., 1973. A simple cross-dating program for tree-ring research. Tree-Ring Bull. 33, 7e14. Barniak, J., Kra˛ piec, M., Jurys, L., 2014. Subfossil wood from the Rucianka raised bog (NE Poland) as an indicator of climatic changes in the first millennium BC. Geochronometria 41 (1), 104e110. Battaglia, S., Leoni, L., Sartori, F., 2002. Mineralogical and grain size composition of clays developing calanchi and biancane erosional landforms. Geomorphology 49, 153e170. Baumgart-Kotarba, M., 1991-1992. Geomorphological development of the Kotlina Orawsko-Nowotarska Basin in condition of neotectonic movements. Stud. Geomorphol. Carpatho-Balcanica 25e26, 3e28. Berglund, B.E., Ralska-Jasiewiczowa, M., 1986. Pollen analyses and pollen diagrams. In: Berglund, B.E. (Ed.), Handbook of Holocene Palaeoecocology and Palaeoecology and Palaeohydrology. Wiley, Chichester, pp. 455e484. Birks, H.J.B., 2008. Holocene climate research e progress, paradigms, and problems. In: Battarbee, R.W., Binney, H.A. (Eds.), Natural Climate Variability and Global Warming: a Holocene Perspective. Wiley-Blackwell, Oxford, pp. 7e57. Bond, G., Showers, W., Cheseby, M., Lotti, R., Almasi, P., Domenocal, P., Priore, P., Cullen, M., Hajdas, I., Bonani, G., 1997. A pervasive millennial e scale cycle in North Atlantic Holocene and global climates. Science 278, 1257e1266. Bond, G., Kromer, B., Beer, J., Muscheler, R., Evans, M.N., Showers, W., Hoffman, S., Lotti-Bond, R., Hajdas, I., Bonani, G., 2001. Persistent solar influence on North Atlantic climate during the Holocene. Science 294, 2130e2136. Bortenschlager, S., 1982. Chronostratigraphic subdivision of the Holocene in the Alps. Striae 16, 75e79. Bridge, M.C., Haggart, B.A., Lowe, J.J., 1990. The history and palaeoclimatic significance of subfossil remains of Pinus sylvestris in Blanket Peats from Scotland. J. Ecol. 78, 77e99. Bronk Ramsey, C., 2008. Deposition models for chronological records. Quat. Sci. Rev. 27, 42e60. Bronk Ramsey, C., 2009. Bayesian analysis of radiocarbon dates. Radiocarbon 51 (1), 337e360. Cook, E.R., Krusic, P.J., 2005. ARSTAN V. 41d: a Tree-ring Standardization Program Based on Detrending and Autoregressive Time Series Modeling, with Interactive Graphics. Tree-Ring Laboratory, Lamont-Doherty Earth Observatory of Columbia University, Palisades, New York USA.

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