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Holocene treeline and timberline changes in the South Carpathians

(Romania): Climatic and anthropogenic drivers on the southern slopes of the
Retezat Mountains

Article in The Holocene · April 2017

DOI: 10.1177/0959683617702227

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HOL0010.1177/0959683617702227The HoloceneVincze et al.

Research paper

The Holocene

Holocene treeline and timberline

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DOI: 10.1177/0959683617702227
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(Romania): Climatic and anthropogenic
drivers on the southern slopes of the
Retezat Mountains

Ildikó Vincze,1,2 Ildikó Orbán,3,4 Hilary H Birks,3 Ilona Pál,1,2

Walter Finsinger,5 Katalin Hubay,6 Elena Marinova,7
Gusztáv Jakab,8,9 Mihály Braun,6 Tamás Biró,10 Mónika Tóth,11,12
Claudia Dănău,13 Iosif V Ferencz14 and Enikő K Magyari1,15

Abstract
Two high-altitude lake-sediment sequences (Lake Lia, 1910 m a.s.l. and Lake Bucura, 2040 m a.s.l.) from the Retezat Mountains (South Carpathians,
Romania) were analysed using multi-proxy methods to study responses of treeline, timberline and alpine/subalpine vegetation to climate change and
human impact during the past 16,000 years. Woody species (Pinus mugo, Pinus cembra, Picea abies and Juniperus communis) reached Lake Lia between 12,000
and 11,800 cal. yr BP, whereas P. mugo colonised the shores of Lake Bucura at 9600 cal. yr BP. Lake Lia was in the timberline ecotone between 8000
and 3200 cal. yr BP, in semi-open P. cembra and Picea abies woodland, probably mixed with P. mugo on the steeper slopes. Lake Bucura was surrounded
by the upper part of the krummholz zone during the mid-Holocene. The increase in P. cembra after c. 6000 cal. yr BP around Lake Lia suggests that the
composition of the timberline forest changed. The disappearance of P. cembra and Picea abies around Lake Lia at ~3000 cal. yr BP reflects descent of the
timberline. A large mean July temperature decline between 3300 and 2800 cal. yr BP may have driven or at least contributed to the descent of the Picea
abies–P. cembra forests. An increase in human indicator pollen types in Lake Bucura around 4200 cal. yr BP may reflect human impact in the naturally open
alpine zone in the Late Bronze Age. In contrast, human impact likely appeared considerably later, around 2650 cal. yr BP (Early Iron Age) around Lake
Lia in the upper subalpine zone. Human impact likely intensified after 2200 cal. yr BP at both sites that resulted in the lowering of the krummholz zone.
We conclude that climate change and human impact both played an important role in the lowering of the treeline and timberline in the late-Holocene.

Keywords
Holocene climate, human impact, plant macrofossils, Retezat Mountains, timberline, treeline

Received 6 July 2016; revised manuscript accepted 9 February 2017

Introduction
The treeline ecotone is a transitional belt between the upper limit of Tausz, 2007). In the treeline ecotone, the density of trees decreases
closed forest (altitudinal timberline) and treeless alpine vegetation upwards, and the highest elevation reached by single tree individu-
in high mountains (Däniker, 1923; Lotter et al., 2006; Wieser and als taller than 2–3 m defines the treeline (Arno and Hammerly,

1MTA-MTM-ELTE Research Group for Paleontology, Hungary 10Department of Physical Geography, Eötvös Loránd University, Hungary
2Department of Physical and Applied Geology, Eötvös Loránd 11Balaton Limnological Institute, Centre for Ecological Research,
University, Hungary Hungarian Academy of Sciences, Hungary
3Department of Biology and Bjerknes Centre for Climate Research, 12Centre for Ecological Research, Hungarian Academy of Sciences,

University of Bergen, Norway GINOP Sustainable Ecosystems Group, Hungary

4Department of Plant Systematics, Ecology and Theoretical Biology, 13Administratia Parcului National Retezat (APNR), Romania

Eötvös Loránd University, Hungary 14Department of Archaeology, Muzeul Civilizatiei Dacice si Romane
5Palaeoecology, ISE-M (UMR 5554 CNRS/UM/EPHE), France Deva, Romania
6Hertelendi Laboratory of Environmental Studies, Institute for Nuclear 15Department of Environmental and Landscape Geography, Eötvös

Research, Hungarian Academy of Sciences, Hungary Loránd University, Hungary

7Center for Archaeological Sciences, GEO-Instituut, University of

Leuven, Belgium Corresponding author:

8Faculty of Economics, Agricultural and Health Studies, Szent István Ildikó Vincze, Department of Physical and Applied Geology, Eötvös
University, Hungary Loránd University, Pázmány Péter sétány 1/C, Budapest, 1117 Pest,
9Institute of Archaeology, Research Center for Humanities, Hungarian Hungary.
Academy of Sciences, Hungary Email: [email protected]
2 The Holocene 00(0)

1984; Schwörer et al., 2013; Tinner and Theurillat, 2003). Dwarfed Coldea and Cristea, 1998). Geanana (1991) concluded that logging
trees and shrubs growing above the treeline form the krummholz and grazing widely affected the landscape during the 20th century
zone (Holtmeier, 1981; Körner, 2012a; Tinner, 2007). lowering the elevation of the treeline about 100 m below the poten-
Since the elevation of the potential timberline and treeline is tial tree limit. These inferences contrast to records from the Eastern
primarily determined by climate (Holtmeier, 1981; Körner, 2003), Carpathians and the Alps. The elevation of the treeline in the Cen-
researchers generally agree that the treeline ecotone will be tral Alps was probably 150–200 m higher than today’s tree limit
strongly affected by present-day warming (Grace et al., 2002; between 11,000 and 4500 cal. yr BP (Tinner and Theurillat, 2003);
Solomon et al., 2007). Signs of upward treeline and timberline and the onset of the exploitation of the alpine grassland zone started
shifts have already been detected in several decadal-scale studies much earlier (late Neolithic; Oeggl, 1994; Tinner et al., 2007). In
(Harsch et al., 2009; Heiri et al., 2014; Körner, 2012b; Theurillat the Alps, timberline descended around 6500 cal. yr (Tinner, 2007)
and Guisan, 2001; Tinner and Ammann, 2005), supporting pre- and in the Eastern Carpathians around 6000–5000 cal. yr BP (Feur-
dictions that ongoing global warming leads to rises in treeline and dean et al., 2016; Geanta et al., 2014).
timberline with corresponding reductions in alpine grassland The aims of this study are to investigate the vegetation changes
(Schwörer et al., 2013). at the current treeline ecotone over time and to understand
Treeline and timberline dynamics can be understood using a whether the driving factors of these changes were climate change
combination of short-term ecological observations with longer- or human impact. To address these aims, we use plant macrofos-
term reconstructions based on palaeoecological proxies such as sil, pollen, stomata and charcoal records from two lakes situated
fossil pollen, plant macrofossils, stomata and charcoal, and in at different altitudes in the same mountain valley. The study sites
some cases megafossils and tree rings (Birks and Birks, 2000; are located in the Retezat Mountains (Figure 1), about 300 km
Reasoner and Tinner, 2009; Tinner, 2007). Several long-term south of the sites in the Eastern Carpathians investigated by Feur-
multi-proxy records from the Alps illustrate the effects of past dean et al. (2016) and Geanta et al. (2014). Neolithic and Bronze
climate and land-use changes on treeline ecology (Berthel et al., Age archaeological sites have been reported from the lowlands of
2012; Finsinger and Tinner, 2007; Gobet et al., 2003, 2005; Lang the nearby Haţeg Basin and from south-east of our study area
and Tobolski, 1985; Rey et al., 2013; Schwörer et al., 2013, 2014; (Institutul National al Patrimoniului, 2013; Luca et al., 2005),
Tinner et al., 1996; Tinner and Kaltenrieder, 2005; Tinner and indicating human presence in the region. Our key questions are as
Theurillat, 2003; Wick et al., 2003; Wick and Tinner, 1997). The follows: (1) how did Holocene treeline ecotone vegetation in the
use of plant macrofossil and stomata analysis in many of these Retezat Mountains respond to climate change? (2) did human
studies provides a detailed picture of local vegetation dynamics activities influence high mountain ecosystems? and (3) how do
around the treeline ecotone (Tinner, 2007), because of the lower Holocene treeline fluctuations in the Retezat Mountains compare
dispersal distance of plant remains compared with pollen with the treeline variations in the Eastern Carpathians and other
(Ammann et al., 2014; Birks and Bjune, 2010). In contrast, plant European mountains?
macrofossil analysis has only recently been adopted to investigate
long-term treeline dynamics in the Carpathians (Feurdean et al.,
2016; Geanta et al., 2014), and previous reconstructions were Study area
mainly based on pollen records (Fărcaş et al., 1999, 2013; Feur- The Retezat Mountains are situated on the western side of the
dean and Willis, 2008). Because of the low number of long-term South Carpathians in Romania (Figure 1). The mountain range
records and the inherent limitations of pollen records as a tool to stretches from east to west between the Jiu river valley (about 45
reconstruct treeline dynamics, knowledge of the long-term km north from the valley) and the Haţeg Basin (about 25 km from
dynamics of upper treeline and timberline in the Carpathians is the basin; Figure 1b), and most of it belongs to the Retezat
scarce and fragmentary. National Park, which hosts a rich flora with 90 endemic and 130
Currently, evidence for past human activity and their effects on rare and protected species (Borza, 1934; Csűrös et al., 1956;
mountain vegetation in the South Carpathians and transhumance is Nyárády, 1958). The bedrock is predominantly granite and grano-
based on documentary archives, modern vegetation studies and diorite with crystalline schist intercalated between the northern
archaeological data. According to historical evidences, the first Retezat and the southern Buta massifs (Jancsik, 2001). There are
shepherds moved up to the high mountain pastures from the plains more than 50 mountain peaks reaching altitudes >2000 m a.s.l.,
only 800 years ago and started clearing forests to create pastures and the highest is the Peleaga peak (2509 m a.s.l.).
and meadows in the Romanian section of the Carpathians, later The climate is temperate continental in the lowland area (Far-
spreading northwards along the Carpathian chain (Mertens and cas and Sorocovschi, 1992), and alpine climate characterises the
Huband, 2004). Written sources document that grazing contributed high mountains. Because of the combined oceanic and Mediterra-
to the lowering of the timberline in the Iezer Mountains (Mihai nean climatic influence, it is one of the wettest regions in the
et al., 2007), while the enlargement of grazing fields by slash and Romanian part of the Carpathians (Magyari et al., 2013). The pres-
burn activities is documented in the Fagaras Mountains (Nedelea ent treeline is located at 45°35′N, 22°87′E at 1900 m a.s.l., while
et al., 2009). Prehistoric settlements are found in the lowlands timberline runs at 45°34′N, 22°88′E at 1850 m a.s.l. on the south-
surrounding the Retezat Mountains (Supplementary Figure 6, ern slope. According to the study of Farcas and Sorocovschi
available online), and evidence for prehistoric pastoral farming is (1992), January mean temperature is −8°C at the treeline (1900 m
also available in the nearby areas (e.g. Foeni-Salaș, W-Banat, a.s.l.) and between −7°C and −8°C at the timberline on the south-
Romania; Arnold and Greenfield, 2006). A recent study by Juler ern slope of the Retezat Mountains. July mean temperatures are
(2014) focused on Medieval long-distance transhumance in 10°C at the treeline and between 10°C and 11°C at the timberline.
Romania and its fast disappearance after communist times. In that The annual mean precipitation is c. 1200 mm at the treeline, 1300
study, an important transhumance route runs through the Retezat mm at the timberline and <1200 mm above 2000 m a.s.l. These
Mountains. Some vegetation studies conclude that intensive sum- environmental conditions determine the altitudinal zonation of the
mer alpine pasturing had a direct influence on the treeline ecotone vegetation (Borza, 1934; Csűrös et al., 1956; Nyárády, 1958) that
vegetation (Coldea and Cristea, 1998) and suggested that land use is summarised in the Supplementary Material (available online).
led to the appearance of new plant communities (e.g. Potentillo–
Nardion and Calamagrostion) and facilitated the spread of heath-
lands (Loiseleurio–Vaccinion, Rhododendro–Vaccinion) in the Study sites
low-alpine zone following clearance of the native woody vegeta- We analysed two lake-sediment sequences from the Bucura valley
tion (Rhododendro–Pinetum mugi and Bruckenthalio–Piceetum; on the southern flank of the Retezat Range (Figure 1d). Lake Lia
Vincze et al. 3

Figure 1. (a) The location of the Retezat Mountains (RM) in Europe and in the Southern Carpathians (H.b. = Haţeg basin). (b) Selected
sites with important palaeorecords mentioned in the text (b and c): 1. Lago Basso (2250 m a.s.l.; Wick and Tinner, 1997), 2. Gouille Rion
(2343 m a.s.l.; Wick and Tinner, 1997), 3. Sägistalsee (1935 m a.s.l.; Wick et al., 2003), 4. Hinterburgsee (1515 m a.s.l.; Heiri et al., 2003), 5.
Bachalpsee (2265 m a.s.l.; Lotter et al., 2006), 6. Feld mire (2130 m a.s.l.; Lotter et al., 2006), 7. Emines (2288 m a.s.l.; Berthel et al., 2012), 8.
Lauenensee (1381 m a.s.l.; Rey et al., 2013), 9. Iffigsee (2065 m a.s.l.; Schwörer et al., 2013), 10. Lac de Fully (2135 m a.s.l.; Finsinger and Tinner,
2007), 11. Lej da Champfèr (1791 m a.s.l.; Gobet et al., 2003), 12. Lej da San Murezzan (1768 m a.s.l.; Gobet et al., 2003), 13. Lake St Anne
(Eastern Carpathians; 950 m a.s.l., Magyari et al., 2014). (d) Location of Lake Lia (e) (1910 m a.s.l.) and Lake Bucura (f) (2040 m a.s.l.) in the
Bucura valley on the southern slope of Retezat Mountains. (e) Bathymetry of Lake Lia: 45°21′7.3″N, 22°52′39.3″E; coring point marked by a
cross. (f) Bathymetry of Lake Bucura: 45°21′42.9″N, 22°52′27″E coring point marked by a cross. The bathymetric maps were modified after
Vespremeanu-Stroe et al. (2008).

at 1910 m a.s.l. is a small shallow lake (1.26 ha, average water Lake Bucura at 2040 m a.s.l. lies above the treeline and is the
depth: 0.8 m; Figure 1e) located at the present treeline. It has two largest lake in the Retezat (8.92 ha, maximum water depth: 17.5
inflow streams and one outflow. Dense Pinus mugo thickets m (Vespremeanu-Stroe et al., 2008) and Figure 1f). It receives
occupy its steep western slope and only one dwarfed Picea abies water from two inflow streams and has an outflow on its southern
individual is well established on the lakeshore. Picea abies and side towards Lake Lia. It is surrounded by open alpine meadow
Pinus cembra are present near the southern shore c. 150 m away, vegetation and screes. At present, P. mugo is scarce on the lake-
and a few specimens grow on the ridge above the lake. The lake shore (10 individuals growing <20 m from its shores), but it also
and its inflowing streams are fringed by herbs of mesic or wet soils grows above the lake. On rocky terrain Juniperus communis is
(e.g. Caltha palustris, Rumex alpinus, Aconitum tauricum, Adeno- also present. Cattle grazing is frequent during the summer around
styles alliariae, Epilobium angustifolium, Heracleum palmatum). both lakes. The treeline is formed by Picea abies and P. cembra
4 The Holocene 00(0)

individuals around 1900 m a.sl., while the timberline is around smallest taxon level by comparison with the reference collections
1850 m a.sl. on the southern slope of Bucura valley. of the MTA-MTM-ELTE Research Group for Paleontology, the
Hungarian Natural History Museum and the Palaeoecology Labo-
ratory of Bergen University. For conifer needles, the numbers of
Materials and methods complete needles (WN), needle fragments (NF), needle tips (NT)
Sediment sampling and needle bases (NB) and the highest value among the needle
tips and needle bases (NH) were noted. The minimum number of
Sediment cores from Lake Lia (LIA-1) and Lake Bucura (BUK-
needles (MN) in a sample was estimated using the following for-
5) were obtained in 2008 using Livingstone and Kullenberg pis-
mula: MN = (WN + (NF/2) + NH). Sediment components (e.g.
ton corers. The 880-cm-long sediment core from Lake Lia was
Cladocera, chironomids, bryophyte leaves and Cenococcum scle-
taken close to the western shore (45°21′7.3″N, 22°52′39.3″E)
rotia) of Lake Bucura were counted and calculated according to a
under 1.1 m water (Figure 1e). The 550-cm-long sediment core
modified version of the QLCMA method of Barber et al. (1994),
from Lake Bucura was taken in the northern sub-basin
as detailed in Jakab et al. (2004). Plant macrofossil concentrations
(45°21′42.9″N, 22°52′27.0″E) under 13.9 m water (Figure 1f).
were standardised to a reference sediment volume of 20 cm3 for
The Bucura sediment was contaminated below 495 cm depth
Lake Lia (average samples volume was 21 cm3) and 5 cm3 for
because of technical problems, so only the top 495 cm was anal-
Lake Bucura (average sample volume was 5.4 cm3). Macrofossil
ysed. Because of the unavailability of a specialised surface sedi-
concentration diagrams were drawn using TILIA software
ment sample, the core tops do not cover the most recent times. In
(Grimm, 1992) and divided into macrofossil assemblage zones
the laboratory, the sediment cores were photographed and
(LM 1–5, BM 1–5) visually, taking into account major composi-
lithostratigraphically described following Troels-Smith (1955;
tional and concentration changes in the macrofossil assemblages.
Figure 2), sliced into plastic bags and refrigerated.

Radiocarbon dating and chronology Pollen and stomata analyses

Table 1 lists the 28 accelerator-mass spectrometry (AMS) 14C Subsamples of 1 cm3 were taken at 8-cm (Lake Lia) and at 4- to
measurements obtained. Plant macrofossils were 14C dated plus 8-cm (Lake Bucura) intervals and processed for pollen analysis
three dates on the <250 µm organic-rich sediment fraction abun- using standard methods (Bennett and Willis, 2001), but excluding
dant in fine organic debris, Cladocera and Chironomidae the acetolysis step. Pollen, spores and stomata were identified and
remains. Previous attempts to compare the 14C-dating results of counted under an Olympus CX41 light microscope at ×400 and
various sediment components of the lake sediments in the ×1000 magnification. At least 500 terrestrial pollen grains were
Retezat Mountains suggested that the lakes show no reservoir counted at each level. For identification, the pollen atlases of
effect (Hubay et al., 2016; Magyari et al., 2009a, 2012); the bed- Reille (1992, 1995, 1998) and the pollen identification keys of
rock is poor in calcium, and generally terrestrial plant macrofos- Moore et al. (1991) and Beug (2004) were used. For the identifi-
sil components provide similar 14C ages to the aquatic animal cation of the Sordariales-type spores, descriptions in Van Geel
components (Magyari et al., 2009a, 2012). No macrofossils et al. (1989) were used. Pollen diagrams were drawn using Psim-
were found in the highly inorganic late-glacial sediment of Lake poll v.27 (Bennett, 2005). Local pollen-assemblage zones were
Lia, so we dated bulk sediment. 14C dates were calibrated into determined using the method of optimal splitting by information
calendar years before present (AD 1950) using the IntCal13 content on the terrestrial pollen taxa that reached at least 5% in
calibration curve (Reimer, 2013). Bayesian age–depth model- one sample as implemented in Psimpoll. Seven pollen zones were
ling was used to detect outlier radiocarbon dates (Blaauw and identified in each sequence. Pollen percentages are based on the
Christen, 2013) with the R-BACON package (Supplementary terrestrial pollen sum (M). Relative frequencies (F) of aquatic
Figure 1, available online). The programming algorithm of pollen types + fungal spores + algae (A) were calculated accord-
BACON uses the deposition time of the previous depth interval ing to the formula F = ΣA/(M + A).
to predict age–depth relations in the subsequent interval. Even Stomata of Picea abies, Abies alba, J. communis, Larix
though this so-called ‘memory time’ can be modified, when decidua, P. cembra and species of the subgenus Pinus (diploxylon
there are rapid shifts in deposition time the model still provides pines) were identified and counted using the keys of Sweeney
biased age–depth curves. Therefore, we experimented with (2004) and Magyari et al. (2012). Stomata of the subgenus Pinus
more traditional curve fitting methods in CLAM v2.1 using the (diploxylon pines) are probably produced by P. mugo, because it
outlier information from BACON (Figure 2). Finally, we is the only diploxylon pine at mid and high altitudes in the Retezat
decided to use the smooth-spline curve fitting function to model Mountains today.
the age–depth relationship in the two lake-sediment sequences Pollen types indicative of human impact are presented in Fig-
(Blaauw, 2010), as shown in Figure 2 together with the sediment ures 5 and 7 and Supplementary Figure 5 (available online). They
deposition time. Further details on the radiocarbon dating and include herbs associated with pastures, meadows and trampled
age–depth modelling of the Retezat lake sediments are provided ground at both high and low altitudes: Plantago lanceolata,
in Hubay et al. (2016). Plantago major/media, Artemisia and Poaceae. Because of their
wide distribution, good pollen productivity and dispersal, we
interpreted the relative frequency increases in these pollen types
Plant macrofossil analysis as indicative of either local or regional (lower elevation) human
Two-cm-thick subsamples (average volumes 21 and 5.4 cm3 in impact. Alpine grasslands and mats respond to pasturing and
Lake Lia and Lake Bucura, respectively) were taken at 8- to mowing by the expansion of nutrient-demanding herbs (Oeggl,
16-cm intervals from the LIA-1 core and at 8-cm intervals from 1994), many of which are insect-pollinated, producing low
the BUK-5 core. Bucura subsamples were briefly soaked in 10% amounts of pollen that do not travel far from their origin. They are
NaOH and wet-sieved through 250 and 180 µm meshes. Clay-rich therefore underrepresented in the pollen records. Supplementary
samples from Lake Lia were treated with Na4P2O7 and all were Table 2 (available online) lists some of these taxa in the Retezat
sieved through a 125-µm mesh. Plant remains were picked out Mountains frequently encountered on our pollen slides. We inter-
systematically under a stereomicroscope (Olympus SZ 51 and preted the relative frequency increase in these herb pollen types as
LEITZ, ×10 magnification), identified using keys (Bojnanský and indicative of local and extra-local human impact, mainly pastur-
Fargašová, 2007; Katz et al., 1965; Schermann, 1967; Tomlinson, ing in the vicinity of the studied lakes. These taxa include Vac-
1985; Velichkevich and Zastawniak, 2008) and determined to the cinium-type, Adenostyles, Achillea, Sedum and Phyteuma-type.
Vincze et al. 5

Figure 2. Age–depth models and deposition times for the (a) Lake Lia and (b) Lake Bucura sediment profiles. The age–depth models are based
on 16 14C dates from Lake Lia and 12 14C dates from Lake Bucura sequence calibrated using CALIB Rev. 7.2.0. and the IntCal13 data-set (Table
1). The age–depth models were created with the smooth-spline curve fitting function in CLAM v2.1 (Blaauw, 2010). The lithostratigraphy is
represented using the symbols of Troels-Smith (1955) on the left of each model and detailed in Supplementary Table 1 (available online). Troels-
Smith’s symbols: Gyttja (Ld), substantia humosa (Sh), herbaceous organic mud (Ld), fine sand (Ga), gravel (Gg), clay (As), silt (Ag), herbaceous
remains of bryophytes (Th), coarse sand (Gs).

Pollen studies from high altitudes in the Alps show that the pollen (microCHAR). These records are used to reconstruct regional
of local pasturing indicator herbs can be used successfully to (20–50 km extent) fire activity around the study sites following
identify high-altitude pasturing (Bortenschlager, 1991; Court- Tinner et al. (1998). Experimental and empirical evidence indi-
Picon et al., 2006; Vorren et al., 1993). cates that most macrocharcoal particles (>160 µm in diameter)
originate from fires nearer to their source area (max. 10–20 km)
because of the more limited dispersal of larger charcoal particles
Loss-on-ignition
(Lynch et al., 2004; Oris et al., 2014; Peters and Higuera, 2007).
Subsamples of 1 cm3 sediment were taken at 1-cm intervals from Long-term variations in total macroCHAR may reflect biomass
core LIA-1 and at 2-cm intervals from core BUK-5. Samples burning, that is, an integrated signal of charcoal production and
were dried at 105°C overnight and then burned at 550°C for 4 h. fire frequency. To determine long-term variations in regional fire
Organic matter content was estimated as the percentage loss in activity and local biomass burning, microCHAR and macro-
dry weight after ignition (Heiri et al., 2001). CHAR records were smoothed with a 900-year-bandwidth lowess
(Cleveland and Devlin, 1988). Because the sample deposition
Charcoal analyses times (years sample−1) for Lake Bucura were too high below
100 cm depth, the record was not analysed to infer local fire
For macrocharcoal analysis, subsamples were taken at every cen-
episodes using the decomposition approach (Finsinger et al.,
timetre from the top 100 cm of the BUK-5 core, followed by 2 cm
2014; Higuera et al., 2009; Kelly et al., 2011).
contiguous (merged) sub-sampling below 100 cm. For core LIA-
1, samples were taken at 1-cm contiguous intervals. Methods
were as described in Finsinger et al. (2016), and they resulted in Results and interpretation
counts of macrocharcoal particles and their surface area (macro-
charcoal accumulation rates (macroCHAR)). Chronology and sediment stratigraphy
Microcharcoal particles (>10 µm) were counted on the pollen For a detailed lithostratigraphic description of the profiles, see
slides following Tinner and Hu (2003) and Finsinger and Tinner Supplementary Table 1 (available online). The lithostratigraphy is
(2005) and presented as microcharcoal accumulation rates shown in Figure 2 using the Troels-Smith’s (1955) notation.
6 The Holocene 00(0)

Table 1. AMS 14C dates from Lake Bucura (BUK-5) and Lake Lia (LIA-1). 14C measurements were done in the Hertelendi Laboratory of
Environmental Studies, Institute of Nuclear Research of the Hungarian Academy of Sciences, Debrecen. Outliers were detected with the
BACON R package (Supplementary Figure 1, available online). For the LIA-1 core the age of the lake-sediment surface was estimated by
extrapolation.

Core Lab. code Dated material Depth (cm)a 14C age yr BP ± 1 σ Calibrated range yr Remarks
BP (±2 σ)

BUK-5 DeA-1636 <250 µm sediment fraction 50 1817 ± 34 1691–1827

BUK-5 DeA-3244 P. mugo needle, plant macrofossil 101 2826 ± 46 2841–3067
BUK-5 DeA-1635 <250 µm sediment fraction 150 2640 ± 36 2727–2800 Outlier
BUK-5 DeA-3245 Plant macrofossil 186 3569 ± 51 3702–3984
BUK-5 DeA-1634 <250 µm sediment fraction 223 3868 ± 39 4222–4415
BUK-5 DeA-3271 Plant macrofossil 271 4028 ± 38 4417–4584
BUK-5 DeA-1633 <250 µm sediment fraction 330 5199 ± 44 5892–6027
BUK-5 DeA-3303 Plant macrofossil 381 6962 ± 139 7565–8040
BUK-5 DeA-1632 <150 µm sediment fraction 430 7531 ± 52 8280–8416
BUK-5 DeA-3273 Bulk (LT)b 494 9168 ± 40 10,236–10,428
BUK-5 DeA-3274 Bulk (HT)b 494 9153 ± 39 10,230–10,419
BUK-5 DeA-3246 Picea abies needles, plant macrofossil 495 9209 ± 77 10,235–10,564
LIA-1 DeA-1647 <250 µm sediment fraction 80 1018 ± 35 900–985
LIA-1 DeA-3134 P. mugo needle 96 950 ± 36 789–930 Outlier
LIA-1 DeA-3135 Plant macrofossil 133 2782 ± 37 2784–2961
LIA-1 DeA-1646 <250 µm sediment fraction 162 3715 ± 40 3961–4157
LIA-1 DeA-3136 Picea abies needle 218 4057 ± 40 4424–4644
LIA-1 DeA-1645 <250 µm sediment fraction 278 5576 ± 43 6293–6436
LIA-1 DeA-3137 Picea abies needle 337 6147 ± 43 6929–7166
LIA-1 DeA-1644 <250 µm sediment fraction 383 7419 ± 48 8165–8363
LIA-1 DeA-3138 P. mugo needle 433 7987 ± 45 8699–9005
LIA-1 DeA-1643 <250 µm sediment fraction 494 8846 ± 53 9732–10,165
LIA-1 DeA-1641 >250 µm sediment fraction 560 9386 ± 54 10,491–10,747
LIA-1 DeA-3139 P. mugo needle 590 9521 ± 43 10,672–10,884
LIA-1 DeA-3270 Rumex-type fruit 624 9560 ± 42 10,727–11,095 Outlier
LIA-1 DeA-1639 >150 µm sediment fraction 683 11462 ± 63 13,160–13,440
LIA-1 DeA-1640 <150 µm sediment fraction 683 11324 ± 65 13,070–13,291
LIA-1 DeA-3247 Bulk 703 9884 ± 82 11,173–11,625 Outlier

aDepths (cm) are the bottom depths of samples; all dated sample width is 1 cm long.
bLow temperature (LT): 400°C; high temperature (HT): 800°C.

The BACON age–depth model of Lake Lia suggested three Lia. The single charred Picea abies needle fragment could have
outlier 14C dates (Table 1, Supplementary Figure 1, available been transported for hundreds of metres in the convective col-
online) that were excluded from the smooth-spline age–depth umns of fires at lower altitudes (Pisaric, 2002).
model. The smooth-spline age–depth model (Figure 2a) was con- The treeline reached the catchment around 12,010 cal. yr BP
structed between the top and bottom of the core (between 31 and (start of zone LM-2; 12,010–9700 cal BP; 636–483 cm) as shown
830 cm). The estimated ages of the top and bottom samples (~250 by the first appearance of P. mugo and low concentrations of P.
and ~17,100 cal. yr BP, respectively) were obtained by extrapola- cembra needles that were closely followed by the first occurrences
tion of the age–depth model and therefore have high uncertainty. of macrofossil remains of Picea abies and J. communis. Stomata
The average sediment deposition time is 26 yr cm−1 before 12,000 of P. mugo–type were first detected at 11,400 cal. yr BP, closely
cal. yr BP, 16 yr cm−1 between 12,000 and 6900 cal. yr BP, followed by the appearance of P. cembra and Picea abies stomata
increasing to 22 yr cm−1 between 6900 and 2000 cal. yr BP, before at 11,000 cal. yr BP. One L. decidua stoma was found at 10,700
decreasing towards the present. cal. yr BP (568 cm). During the equivalent pollen zone, L-4 (Fig-
One outlier 14C date was detected at Lake Bucura (Table 1, ure 5) percentages of Picea abies, Corylus and tree pollen of Quer-
Supplementary Figure 1, available online) and was excluded from cetum mixtum (Quercus, Ulmus, Tilia and Fraxinus) increased
the smooth-spline age–depth model (Figure 2b). The estimated and herb pollen of Artemisia, Chenopodiaceae and Poaceae dis-
age of the top of the core is ~650 cal. yr BP, obtained by extrapo- tinctly decreased reflecting the vegetation response to the
lation of the age–depth model and therefore it has a high uncer- Younger Dryas/Holocene transition that is also evident in other
tainty. The average sediment deposition time is 26 yr cm−1 records from the region (Magyari et al., 2012). Macrofossils of
between 10,300 and 6300 cal. yr BP, 18 yr cm−1 between 6300 and dwarf shrubs, such as Rhododendron kotchii and Vaccinium sp.,
4200 cal. yr BP and 17 yr cm−1 between 4200 and 700 cal. yr BP. were also present suggesting their rapid local establishment at
1910 m. R. alpinus–type and Heracleum sphondylium/H. palma-
tum became abundant around the lakeshore together with Rumex
Vegetation histories cf. acetosa, C. palustris, Epilobium sp. and Sedum sp. (Figure 3).
Lake Lia. The macrofossil record from Lake Lia is presented in This herb assemblage indicates damp ground and open dry rocks
Figure 3 and the pollen record in Figure 5a. The basal plant mac- and soil. Rumex species may have favoured base-rich disturbed
rofossil zone, LM-1 (between 15,900 and 12,000 cal. yr BP; 782– surfaces after ice retreat, while the presence of Sedum sp. remains
636 cm), contained one charred Picea abies needle fragment at points to dry habitats. The most common Sedum species is Sedum
14,600 cal. yr BP and some seeds of Silene sp., R. alpinus-type atratum in the mountain today that usually occupies scree sur-
and Carex sp. indicating that the treeline was located below Lake faces. The high number of herbaceous remains may indicate strong
 Vincze et al.

Figure 3. Plant macrofossil concentration diagram of Lake Lia plotted on a calibrated age scale showing selected taxa together with the stomata accumulation rates. All macrofossil samples are standardised to
numbers in 20 cm3 sediment. Stomata were counted along with pollen. LM zones are local macrofossil assemblage zones in Lake Lia, delimited visually.
MN: min number of needles; A: anthers; BSC: bract scales; T: twigs; B: buds; S: seeds; F: fruits; L: leaves.
7
8 The Holocene 00(0)

discharge of the inflowing streams or a great extent of permanently Lake Bucura. Lake Bucura is located 130 m above Lake Lia. The
saturated soils near the lakeshore because of high precipitation and macrofossil record is presented in Figure 4 and the pollen record
water levels resulting from melting of large amounts of snow and in Figure 5b.
of the discontinuous permafrost in rock glaciers next to the lake J. communis, R. alpinus, Vaccinium myrtillus and Galium mac-
(Kern et al., 2004; Urdea, 2004; Vespremeanu-Stroe et al., 2012). rofossils are present in the oldest zone (BM-1: 10,300–9530 cal. yr
Around 9700 cal. yr BP (onset of zone LM-3, 483–371 cm), BP, 492–468 cm). The absence of P. mugo and other tree macrofos-
the timberline was probably very close to the elevation of Lake sils clearly indicates that this lake was located above the treeline at
Lia as shown by increased macrofossil concentrations of Picea a time when Lake Lia was between the treeline and the timberline.
abies and by the presence of P. cembra remains. Increasing abun- In zone BM-2, from 9800 to 9500 cal. yr BP, P. mugo needles and
dance of Picea abies macrofossils from 9700 to 8000 cal. yr BP stomata together with P. cembra stomata appeared in Lake Bucura
indicates that the timberline was rising throughout this period. regularly, indicating that the treeline was ascending and that small
However, P. mugo, which competes well in conditions of thick numbers of P. cembra trees reached Lake Bucura by 9500 cal. yr BP.
snow cover and can also colonise avalanche tracks because of its Therefore, Lake Bucura was now in the krummholz zone and the
high flexibility and resistance to breakage (Holtmeier, 2009), treeline was not far below it. Hence, the krummholz zone reached
became more abundant in the local vegetation as shown by high Lake Bucura about 2000 years later than Lake Lia, which was at
needle concentrations and continuous presence of stomata. This this time probably located below the timberline.
suggests that the woodland was relatively open. Betula cf. pubes- In zone BM-3 (7500 and 4900 cal. yr BP, 388–284 cm,) no P.
cens fruits occurred around 8300 cal. yr BP. On the basis of its mugo macrofossils were found between c. 7500 and 6300 cal. yr
modern ecology, B. pubescens is a good coloniser because of its BP and P. cembra stomata were absent. However, unidentifiable
resistance to breakage by snow pack (Holtmeier, 2009), but it was conifer needles were present throughout the zone and P. mugo
rare in the Bucura valley. reappeared after 6300 cal. yr BP suggesting that P. mugo persisted
Between 7915 and 5030 cal. yr BP (zone LM-4, 371–225 cm), on the lakeshore and therefore the lake was still in the krummholz
Picea abies macrofossils were dominant. Conspicuously lower zone. Rumex alpinus and R. acetosa also disappeared from the
concentrations of P. mugo and herbs indicate that the timberline record at the beginning of the zone, but later they reappeared in
was located above the lake, which was now surrounded by closed high concentrations.
Picea-dominated forest. Around 6000 cal. yr BP, the composition Zone BM-4 (4900 and 2200 cal. yr BP; 284–68 cm) was char-
of the timberline forest changed with the increase in P. cembra. acterised by high but decreasing concentrations of P. mugo nee-
Between 5030 and 3300 cal. yr BP (zone LM-5; 225–147 dles and stomata, the presence of P. cembra and some Picea abies
cm), Picea abies still dominated the woodland around Lake Lia, stomata, probably reflecting the highest elevation of the treeline
with a low abundance of P. mugo. Pinus cembra needle concen- at that time. Concentrations of Cladocera ephippia and Chiron-
trations were higher than in the previous zone, suggesting that P. omidae head capsules increased during the past 4000 years (Sup-
cembra became part of the Picea abies–dominated forest. The plementary Figure 3, available online). The increase in Cladocera
increased abundance of P. cembra suggests that the timberline may be because of increasing nutrient availability and hence
was shifting downwards, or that the P. cembra population increasing phytoplankton production stimulated by disturbance in
expanded at the timberline. It is possible that P. mugo was not the catchment. Soil inwash is also indicated by increasing amounts
only part of the Picea abies–P. cembra forests, but may have of the soil mycorrhizal fungus Cenococcum geophilum (Figure 4)
dominated the steep scree slope to west, which is covered by P. and gradually decreasing LOI values (Figure 7) after 4200 cal. yr
mugo thickets today (Figure 1d). BP. Pinus mugo needle concentrations gradually decreased
The uppermost zone between 3300 and 245 cal. yr BP (zone although unidentified needle fragments were still detected until
LM-6, 147 to 31 cm) was characterised by a slight increase in P. 2200 cal. yr BP. At 4200 cal. yr BP, percentages of Sordariales
mugo needles and the near absence of Picea abies and P. cembra: fungal spores (Supplementary Figure 2, available online) peaked
one P. cembra needle at 2700 cal. yr BP and one Picea twig at 79 at Lake Bucura (Figure 5b). The start of increasing percentages of
cm (1090 cal. yr BP) and one Picea anther at 95 cm (1550 cal. yr Poaceae and herb pollen types, including Plantago lanceolata,
BP). Overall, this suggests that the timberline may have shifted Plantago major/media, Artemisia, Adenostyles, Phyteuma-type
below the elevation of lake at 3300 cal. yr BP. However, the con- and Vaccinium-type, and the decreasing percentages of Picea
tinuous presence of Picea abies, P. cembra and P. mugo stomata pollen likely indicate the start of human impact both in the
and of P. mugo needles shows that the treeline was still located mountain valleys and at higher altitudes.
above the lake, as is the case today. Rumex alpinus macrofossils disappeared at 3000 cal. yr BP
The first nearly continuous appearance of pollen of Plantago when Lake Bucura was located above the treeline, while they per-
lanceolata and increased Plantago major/media and Artemisia sisted at Lake Lia (Figures 3 and 4) coinciding with decreasing
pollen in Lake Lia were recorded from c. 2200 cal BP (Figure 5a), Rumex pollen percentages at both sites (Figure 5a and b). Rumex
while Adenostyles, Achillea and Phyteuma-type pollen frequen- species (mainly R. alpinus and Rumex thyrsifolius) are still abun-
cies started to increase from 3000 cal. yr BP (Supplementary Fig- dant in the area today, and the absence of flowering and the
ure 5, available online). In combination with higher Poaceae increasingly vegetative reproduction at increasing altitude
pollen percentages, these probably reflect the expansion of human (Šťastná et al., 2012) may explain the absence of Rumex seeds in
activities in the mountain valleys and also at higher altitudes. Lake Bucura. At Lake Lia, R. alpinus probably found ideal grow-
However, the marked increase in Poaceae pollen (at c. 2700 cal ing conditions associated with soil nitrification near animal enclo-
BP; LPAZ L-7a; Figure 6a) preceded the first appearance of Plan- sures (Šťastná et al., 2012).
tago lanceolata and coincided with a decrease in loss-on-ignition After 2200 cal. yr BP (BM-5; 68 cm), the absence of P. mugo
(LOI) values suggesting that soil erosion increased after the tim- needles, of unidentified conifer needles, and the lack of tree sto-
berline descent. The sediment accumulation rate also increased at mata (apart from a single P. cembra stoma at 1700 cal. yr BP) are
this time (inverse of deposition time, Figure 2). A single Plantago taken to indicate that the treeline continued to descend. Only
sp. seed was found at 31 cm (~245 cal. yr BP), which could reflect Heracleum and Silene sp. seeds, fruits and a few stomata of P.
grazing and/or trampling near the lake. Intensified human activity mugo were still found in Bucura until 1000 cal. yr BP. The corre-
around the lake was also confirmed by the re-appearance of J. sponding increases in Juniperus, Poaceae, Plantago lanceolata,
communis and Potentilla cf. P. argentea macroremains during the Plantago major/media and Sedum pollen suggest more intense
late-Holocene (from 1100 and 300 cal. yr BP; Figure 3). land use during the past 2000 years.
 Vincze et al.

Figure 4. Plant macrofossil concentration diagram of Lake Bucura plotted on a calibrated age scale, together with the stomata accumulation rates. All macrofossil samples are standardised to numbers in 5 cm3
sediment. Stomata were counted along with pollen. BM zones are local macrofossil assemblage zones in Lake Bucura delimited visually.
MN: min number of needles; BS: bract scales; A: anthers; S: seeds; F: fruits; L: leaves; P: perianth.
9
10 The Holocene 00(0)

Figure 5. Pollen percentages of selected arboreal and herbaceous pollen types and fungal spores in (a) Lake Lia and (b) Lake Bucura. The
pollen sum for calculation was at least 500 terrestrial pollens from each sample. Quercetum mixtum includes Quercus, Ulmus,Tilia and Fraxinus
pollen percentages. Empty curves show ×10 exaggerations. LPAZ zones are local pollen-assemblage zones in Lake Lia and Lake Bucura. Pollen
analyst: Ilona Pál.
Vincze et al. 11

Figure 6. Microcharcoal (microCHAR) and macrocharcoal (macroCHAR) accumulation rates in (a) Lake Lia and (b) Lake Bucura. The black
lines are raw data. Grey lines are the data smoothed by lowess (see text).

Figure 7. Summary diagram showing selected proxy records from Lake Lia and Lake Bucura and the chironomid-inferred mean July air
temperatures ± 1 standard errors from Lake Brazi (north slope of Retezat Mountains, 1740 m a.s.l.; Tóth et al., 2015). The vertical red line
indicates the present-day mean July air temperature (~11.2°C; Magyari et al., 2013). LOI 550 = loss-on-ignition at 550°C. Macrofossil records
are summed fragments for the selected taxa. MacroCHAR = macrocharcoal accumulation rate. Stomata AR = stomata accumulation rate. The
pollen of Plantago lanceolata is shown in red as a human indicator pollen type. Macrocharcoal analysis of Lake Lia focused on the Holocene part
of the sediment sequence.

From 2200 cal. yr BP, recurring human impacts are detected, shown that the increase in CHAR values around 500 cal. yr BP is
which likely affected the montane and subalpine forests and the probably not significant. Similarly, the high microCHAR values
alpine grassland zone. These are indicated by the relative pollen in Lake Bucura between 5000 and 3500 cal. yr BP may be because
frequency changes in trees and herbs (Figure 5 and Supplemen- of a sharp change in the sediment accumulation rates (Figure 2)
tary Figure 5, available online, red arrows), especially by the and sediment focusing.
gradual decrease in Picea abies pollen. The fluctuation of Carpi- MacroCHAR (Figure 6) show higher peaks at Lake Lia than at
nus betulus, Fagus sylvatica and Corylus percentages at both Lake Bucura, reflecting high early-Holocene burning of the forest
sites probably reflects human impact at lower elevations (Figure before c. 7000 cal. yr BP and another high biomass burning phase
5 and Supplementary Figure 5, available online, green arrows) centred on 5000 cal. yr BP (Finsinger et al., 2016). Local fires do
together with increases in Poaceae, Alnus viridis, Artemisia, R. not seem to have been intense at Lake Bucura, probably because
acetosa/acetosella and other herb percentages. of the lower biomass availability or the greater distance to burnt
areas. Periods of increased biomass burning as inferred from the
smoothed macroCHAR record from Lake Lia occurred between
Fire histories 9000 and 7000 cal. yr BP, 5500 and 4500 cal. yr BP and around
MicroCHAR (Figure 6) in both sequences indicates higher fire 500 cal. yr BP. A slight increase in macroCHAR starting from
activities between about 10,000 and c. 8000–7000 cal. yr BP in 2000 cal. yr BP could reflect a moderate increase in biomass
the early-Holocene. Fire activity increased again at Lake Lia burning. At Lake Bucura, highest macroCHAR values were
around 500 cal. yr BP. However, Finsinger et al. (2016) have recorded between 10,000 and 7000 cal. yr BP and between 6000
12 The Holocene 00(0)

2015). Concomitantly, treeline ascended and conifers first

appeared around Lake Lia. Warmer-than-present conditions
lasted until around 6300 cal. yr BP (Tóth et al., 2015), when July
temperatures decreased towards present-day levels. On the basis
of the Lake Brazi diatom record, this summer cooling was
accompanied by rising lake levels after 6000 cal. yr BP (Buczkó
et al., 2013). These temperature and hydrology changes coin-
cided with the start of an increase in P. mugo at Lake Bucura. At
Lake Lia, P. cembra became more abundant in the Picea forest
after 6000 cal. yr BP. The increase in P. cembra and the continu-
ing abundance of Picea abies suggest closed forest conditions.
Pinus cembra is more frost resistant than Picea abies (Holtmeier,
2009) and is most abundant near the altitudinal timberline and in
the treeline ecotone. This may suggest that the timberline was
moving downwards.
Figure 8. Schematic illustration of the altitudinal changes in the
The chironomid-inferred temperatures show a marked
treeline (red line) and timberline (blue line) on the southern slope
of the Retezat Mountains during the past 14,000 years. The first decrease to 1.8–1.9°C below modern values between 3000 and
woody macrofossils appeared at Lake Lia around 12,000 cal. yr 1000 cal. yr BP (Tóth et al., 2015). The onset of this decrease
BP. The limits of treeline and timberline are placed between the broadly coincides with the timberline descent below Lake Lia and
sites recording the presence of the respective vegetation type as with the treeline descent further below Lake Bucura. In addition,
inferred by macrofossil and stomata analyses, following Tinner and the diatom record between 3300 and 2800 cal. yr BP characterises
Theurillat (2003). Dashed lines denote stronger uncertainties in the highest lake level in Lake Brazi during the Holocene (Buczkó
the past altitudinal positions. The chronology relies on AMS dating et al., 2013). Overall, summer temperatures played an important
of sediment components from Lake Lia and Lake Bucura; calibrated role in the macrofossil-inferred timberline upward and downward
radiocarbon dates are shown with black dots at the elevations of
shifts during the early and mid-Holocene.
the lakes.
The low plant macrofossil concentrations in Lake Bucura may
imply more open vegetation cover than at Lake Lia. However,
and 3500 cal. yr BP. However, given the likely influence of sedi- taphonomic factors may have been responsible. Lake Lia is small
ment focusing on microCHAR values, the latter high biomass and shallow, whereas Lake Bucura is large and deep. In addition,
burning period is considered an artefact. A slight increase in the bottom of Lake Bucura is covered by aquatic mosses (Supple-
smoothed macroCHAR between 2700 and 1700 cal. yr BP may mentary Table 1, available online), which might have hindered
reflect moderately increased biomass burning at that time. the dispersal of terrestrial macrofossils to the coring site (see, for
MacroCHAR records partially reflect variations in micro- example, Birks et al. (2012)), which was located in the centre of
CHAR (notably at Lake Bucura between 10,000 and 7000 cal. yr the northern sub-basin. In addition, lower sediment-sample vol-
BP, at Lake Lia around 9000–8000 and 500 cal. yr BP) reflecting umes in Lake Bucura (mean 5 cm3) reduce the likelihood of find-
either a synchronicity between local and regional fire occurrences ing plant macrofossils in the sediments compared with Lake Lia
or the deposition of microcharcoal particles because of local fires. (mean volume 21 cm3). However, the reconstructions of the tim-
berline and treeline changes are not based on absolute differences
in plant macrofossil concentrations among the sites, but on the
Discussion variations in plant macrofossil concentrations within each record
The position of the timberline is sensitive to climate changes and the composition of the macrofossil and pollen assemblages.
(Tinner and Theurillat, 2003), because it is mainly influenced by
the accumulated heat during the growing season that is closely
What was the role of humans in the Holocene
connected to the mean temperature of the warmest month (Körner,
treeline and timberline changes?
2012a) and land-use change (Holtmeier, 2009). Our results sug-
gest that between 8000 and 3200 cal. yr BP, the timberline reached Pollens from cultivated plants (e.g. Cerealia-type) were not found
its highest elevation and was located above Lake Lia (1910 m in the sediment of the two lakes. At Lake Bucura, Plantago lan-
a.s.l.), but below Lake Bucura (2040 m a.s.l.; Figures 7 and 8). ceolata pollen first appeared at 4200 cal. yr BP and percentages
They also suggest that Picea abies reached its highest elevation increased together with increases in Plantago major/media,
between 3500 and 3000 cal. yr BP, probably because of more Poaceae, Artemisia, Adenostyles, Vaccinium-type and A. viridis
favourable soil forming conditions (Henne et al., 2011). To inves- pollen. The coincident increase in C. geophilum sclerotia (Figure
tigate the role of summer temperature changes and human activi- 4), the decrease in LOI values (Figure 7, Table 2) and decreasing
ties on the timberline and treeline, we compare the reconstructed P. mugo needle concentrations all indicate the opening of the
treeline and timberline changes (Figure 8) with the charcoal and P. mugo thickets. This could reflect a natural, macroclimate-
human indicator pollen proxies (Figure 7) and with the chirono- induced expansion of alpine meadows with the gradual recession
mid-inferred July temperature changes in Lake Brazi on the of the treeline and timberline already by 4200 cal. yr BP. The
northern slope of the Retezat Mountains (Tóth et al., 2015). We coincident appearance and increase in secondary anthropogenic
ask the following questions: Was the elevated timberline position indicator pollen types, on the other hand, suggest that human
a response to higher growing-season temperatures in the mid- impact in the form of grazing likely appeared around Lake Bucura
Holocene? Was the timberline descent at ~3000 cal. yr BP because coincidentally or shortly after the expansion of the alpine meadow
of Holocene cooling, or did it reflect human impact? areas. Soil erosion may have resulted from both natural and
anthropogenic processes at this time. Regarding the geographical
position of the human impact (whether lowland piedmont or high
Did summer temperature change at the time of mountain origin), it is important that no comparable changes were
treeline ecotone changes in the Bucura Valley? found at Lake Lia around 4200 cal. yr BP. Here, the montane
Summer temperatures increased rapidly in the early-Holocene Picea forest persisted until at least 3200 cal. yr BP and increases
(Figure 7) between 11,500 and 10,200 cal. yr BP (Tóth et al., Plantago major/media, Plantago lanceolata and Poaceae pollen
Vincze et al. 13

Table 2. Indications of human impact (age, cal. yr BP) in Lake Lia and Lake Bucura. The different proxy records are considered to be indicators
of human impact, when accompanied by major changes in the pollen record. Secondary anthropogenic indicator pollen types (Plantago
lanceolata, Plantago major/media, Poaceae, Achillea, Artemisia), tree species from lower altitudes (Fagus, Carpinus betulus), Picea abies and Pinus
pollen percentage changes were considered as indicators of human impact (see Supplementary Figure 5, available online). Note: +: the proxy
likely supports human impact; −: the proxy does not support human impact; 0: no data. The synthetised consequence of human impact is based
on any combination of the previously mentioned comparable proxies from both lakes.

Study sites/proxies Age (cal. yr BP) Sordariales Pollen LOI 550% Microcharcoal Macrocharcoal Cenococcum

Lake Lia (1910 m 400 − + + + + 0

a.s.l.) 600–900 − + − + + 0
1700 − + + − − 0
2200 − + + − − 0
2200–2600 − + + − − 0
2500 − + + − − 0
2600–3000 − + − − + 0
3500 + − + − − 0
3900 − + + + − 0
Lake Bucura (2040 1000 + + + + + +
m a.s.l.) 1500 + + + − − +
1700 + − − + + +
2100 + + + − + +
2500 − − + + + +
2700 + + − − + +
3000 + − + + − +
3300 − − + − − −
3500 − + + + − +
3800 + + + − + +
4000–4200 + + + + + +

percentages occurred later (Supplementary Figure 5, available Mountains) at 3300 cal. yr BP (Magyari et al., 2009b). In that case
online, Figures 5 and 7, Table 2). This difference in timing is the Sordariales increase was not related to grazing but to rottening
probably real, because the increase in F. sylvatica percentages waterlogged wood because of a lake-level rise (Magyari et al.,
occurred at about the same time in both records (around 5000 cal. 2009b). In addition, dung fungal spores (e.g. Sporormiella; Baker
yr BP: the onset of LPAZ L-6b and B-5; Figure 5) implying that et al., 2013), were not found at Lake Bucura and Lake Lia. Hence,
the age–depth models are correct. we conclude that Sordariales-type spores likely indicate seasonal
One possible explanation for the earlier detection of human water-level fluctuation (shore inundation followed by low water
impact indicators at the higher lake might be the smaller represen- level) and related fungal growth on waterlogged wood between
tation of regional pollen rain in the lower lake, which was sur- 4200 and 3800 cal. yr BP at Lake Bucura. On the other hand,
rounded by denser tree cover than the upper lake (Pop et al., other lines of evidence, such as the first increase and first appear-
1965). Surface-pollen studies in the Retezat suggest that the ances of Plantago lanceolata and Plantago major/media, Adeno-
increase in Poaceae pollen and associated herbaceous pollen styles and Phyteuma-type pollen, the prominent increase in
types is a characteristic feature of Lake Bucura that is surrounded Poaceae pollen and the start of the increase in Cenococcum together
by extensive alpine meadows even today. The pollen frequencies with the decrease in LOI values suggest the start of human impact
of Poaceae far exceed the piedmont values (Pop et al., 1965). The at this time.
simultaneous increase in Poaceae, Plantago lanceolata, Plantago Our pollen data suggest that human impact likely intensified
major/media, Phyteuma-type, Adenostyles and Vaccinium-type from ~2200 yr cal BP at both sites. Increasing Poaceae and
pollen thus more likely indicate that the naturally open habitats in anthropogenic indicator herb pollen percentages coincide at Lake
the alpine zone and treeline ecotone were the first to be utilised Bucura with the disappearance of P. mugo macrofossils, suggest-
for seasonal pastural farming during the Bronze Age. ing that the local extirpation of P. mugo was likely human-
Our systematic archaeological data survey suggests that Bronze induced. This inference is also supported by the macrocharcoal
Age settlements were present in the lowland and piedmont zones results that suggest that episodic local fires continued after 2600
(up to 750 m a.s.l.) of the Hunedoara and Gorj counties, located cal. yr BP around Lake Bucura (Figures 6 and 7). The late-Holo-
north and south of the Retezat Mountains (Supplementary Figure 6, cene fires, possibly man-made, could have played an important
available online), but there is no evidence for occupancy of strictly role in the opening up of the alpine krummholz zone in the past
alpine situations. Grazing may have contributed to a decrease in the 2600 years (Finsinger et al., 2016). The intensification of grazing
P. mugo population around Lake Bucura at 4200 cal BP, but simi- around Lake Bucura, suggested by increases in Poaceae, Plan-
larly low macrofossil concentrations were found in the mid-Holo- tago lanceolata, Plantago major-media, Sedum and Juniperus
cene when human impact was probably negligible. To disentangle pollen frequencies after 2000 cal. yr BP (Figure 5b), can be attrib-
whether the percentage peak of Sordariales-type spores in Lake uted to Iron Age cultures (Luca et al., 2005).
Bucura between 4200 and 3800 cal. yr BP was derived from decay- At Lake Lia, the first human impact may have started at 2600
ing wood or dung, we examined samples of both substrates from cal. yr BP, about the time of intensified human impact around
the Retezat Mountains (goat, sheep, cow, bear faeces and decaying Lake Bucura. The expansion of pastures around Lake Lia is indi-
Picea abies and P. mugo wood). However, none of these samples cated by increases in Poaceae pollen and anthropogenic indicator
contained Sordariales-type spores (Supplementary Figure 2, avail- herbs (Table 2, Figures 5a and 7, Supplementary Figure 5, avail-
able online). Sordariales-type spores showed a prominent increase able online) and lower LOI values reflecting increased soil ero-
in the sediments of Lake St Anne (Eastern Carpathians, Ciomadul sion. The plant macrofossil–inferred timberline decrease preceded
14 The Holocene 00(0)

these changes in pollen percentages by ~400 years. This implies (Feurdean et al., 2016; Tinner, 2007; Tinner and Theurillat, 2003).
that the timberline descent was likely not directly human-induced, Our data show no evidence for the influence of Neolithic cereal
but occurred in response to decreasing summer temperatures cultivation, and Plantago lanceolata–type pollen was detected
(Tóth et al., 2015). MacroCHAR showed a significant fire epi- only after 4200 cal. yr BP. This is despite the clear presence of
sode ~50 years earlier, at 2650 cal. yr BP (Figure 7; Finsinger Neolithic settlements in the surrounding lowland and piedmont
et al., 2016) suggesting that the forest burnt at least once in this zones (Supplementary Figure 6, available online). It is possible
period. We also infer that without continuous grazing and burn- that Neolithic people did not use the high mountain pastures for
ing, scattered trees would probably still grow around Lake Lia. grazing in the Retezat Mountains. In contrast, human indicator
The survival of Picea abies and P. cembra trees not far from the pollen types in pollen diagrams of the Alps from c. 8650 cal. yr
lake is attested by the stomata record that shows decreases at 2600 BP show that Mesolithic and Neolithic cultures started to exploit
cal. yr BP, but increases afterwards (Figures 3, 4 and 7). The these mountains (Lotter, 1999; Tinner et al., 2007). Tinner (2007)
intensified human impact since 2200 cal. yr BP did not affect the points out that the regression of the timberline during the past
overall abundance of P. mugo as the macrofossil record indicates 5000 years was primarily caused by human activities. Similarly,
its persistence around the lake, probably on the steep, rocky west- late-Neolithic livestock grazing was demonstrated in the subal-
ern slope that is unsuitable for grazing. pine belt of the Rodna Mountains in the eastern Romanian Car-
At about 1000 cal. yr BP, the peaks in pollen percentages of pathians (Feurdean et al., 2016).
Artemisia, R. acetosa/acetosella, Plantago lanceolata, A. viridis In the Bronze Age, grazing activity does not seem to have
and other herbs decrease in LOI in both sites, and the decreases in resulted in the lowering of the timberline in the Retezat Moun-
Pinus diploxylon–type and Picea abies pollen at Lake Lia suggest tains, in contrast to the Alps. Our data suggest that grazing first
a short-term intensification of clearance of alpine habitats (Table focused on the alpine meadows and the naturally semi-open
2, Figure 5a). krummholz zone in the upper Bucura valley. The first shepherds
contributed to the thinning of the P. mugo thickets and some tree
cutting in the treeline ecotone. The timberline forests were not
Comparison of the Holocene timberline and treeline affected until later, when intensified human impact in the Iron Age
history in the Alps, the Carpathians and the Balkans and Middle Ages led to a considerable lowering of the timberline,
The early-Holocene species composition of the treeline on the as also attested partly by historical documents (Maderspach, 1868;
southern slopes of the Retezat Mountains was characterised by Téglás, 1888a, 1888b). Thus, the treeline ecotone was extended,
Picea abies together with P. cembra (Figures 3 and 4). These tree and the timberline is lower today than its potential elevation in the
species also expanded in the early-Holocene in the Rodna Moun- Retezat Mountains.
tains (Eastern Romanian Carpathians), but their expansion started The treeline reached higher altitudes in the Balkan Mountains
later, around 9800 cal. yr BP (Geanta et al., 2014). In the Central than in the Retezat during the early-Holocene probably because
Alps, Picea expanded much later than in the Southern and Eastern of higher growing-season temperatures. Macrofossil records sug-
Carpathians, at ~5000 cal. yr BP (Tinner et al., 1996). The timber- gest that treeline reached 2250 m in the Pirin Mountains during
line forests in the Central Alps were dominated by L. decidua and the late-glacial and was formed by Betula, Juniperus and Pinus
P. cembra at that time, while L. decidua was replaced by P. mugo peuce in the Southern Balkans (Atanassova and Stefanova, 2003;
at high altitudes in the Retezat in the early-Holocene (Magyari Stefanova et al., 2006; Tonkov et al., 2006, 2011). This species
et al., 2012). These differences may reflect different climatic composition persisted during a time of high summer insolation,
characteristics in the Southern Carpathians that favoured the early relatively warm temperatures and increased aridity until 6700 cal.
expansion and niche acquisition of Picea abies (Holtmeier, 2009). yr BP when Pinus sylvestris, P. peuce and A. alba expanded in the
Plant macrofossil records from the two lakes show that the treeline ecotone suggesting increasing moisture availability in the
maximum timberline elevation between 8000 and 3000 cal. yr BP second half of the Holocene (Tonkov and Marinova, 2005). The
was about 100 m higher than today in the Retezat Mountains. The altitudinal extent of the treeline ecotone appears to have been
maximum treeline position was also about 100 m higher than much larger in the Southern Balkans. The little available informa-
today (i.e. 2000–2050 m). By comparison, plant macrofossil stud- tion suggest that the closed forest limit ascended around 6700 cal.
ies in the Central Swiss Alps showed that the maximum elevation yr BP (Tonkov and Marinova, 2005). This mid-Holocene upward
of the timberline occurred between 10,000 and 6000 cal. yr BP at shift is not detected in the Retezat Mountains. The P. cembra
Gouillé Rion (2343 m a.s.l., Central Swiss Alps, Figure 1c) and increase around Lake Lia (Figure 3) in response to cooling post-
Lago Basso (2250 m a.s.l., North Italian Alps, Figure 1c; Wick dates this event by c. 700 years suggesting regional climatic dif-
and Tinner, 1997) and between 8500 and 4000 cal. yr BP at Lac ferentiation caused vegetation responses at different times in
de Fully (2135 m a.s.l., Figure 1c; Finsinger and Tinner, 2007). these mountains.
The Lac de Fully record is most similar to the Retezat Mountains.
The later advance of the timberline in the early-Holocene could
be because of local factors such as local dryness, topography and Conclusion
local fire events, interacting with the regional climate (Orbán The plant macrofossil, pollen, charcoal and LOI analyses from
et al., 2017). the two study sites on the southern slope of the Retezat Mountains
In the Central Alps, the timberline progressively descended by provide the first records of treeline and timberline variability in
about 180 m from 6000 to 5000 cal. yr BP (Tinner, 2007; Tinner this region and contribute to an improved understanding of the
and Theurillat, 2003). By comparison, on the southern slope of long-term impact of climate and land-use changes on these high-
the Retezat, an increase in P. cembra at the timberline occurred altitude ecosystems. The study confirms that the timberline and
suggesting timberline descent (Figure 3), but the major descent of treeline were higher in the past, as hypothesised from documen-
the timberline commenced only around 3000–3500 cal. yr BP tary evidence (Mertens and Huband, 2004). Both the treeline and
(Figure 8), later than in the Central Alps. This descent was clearly the timberline reached maximum elevations between 8000 and
connected to July temperature decrease in the Retezat Mountains 3200 cal. yr BP and their altitudinal fluctuations were in the order
(Figure 7), while in the Alps the regression of the timberline was of 100–150 m. The vegetation around 2040 m a.s.l was dominated
primarily caused by human impact (Tinner, 2007). by P. mugo during the mid-Holocene, whereas at 1910 m a.s.l.
The intensity of human activities was less in the Retezat closed mixed P. cembra and Picea abies forest developed. The
Mountains than in the Alps and in the Eastern Carpathians pollen and macrofossil evidence that human impact likely became
Vincze et al. 15

important after 4200 cal. yr BP above 2000 m a.s.l. in the Bronze Atanassova J and Stefanova I (2003) Late-glacial vegetational
Age, further back in the past than documentary evidence is able to history of Lake Kremensko-5 in the northern Pirin Mountains,
show. It probably intensified in the treeline ecotone (c. 1900 m southwestern Bulgaria. Vegetation History and Archaeobot-
a.s.l. at that time) during the Iron Age and Medieval Period, any 12(1): 1–6.
around 2600 cal. yr BP. The alpine meadows dominated by Nar- Baker AG, Bhagwat SA and Willis KJ (2013) Do dung fungal
dus stricta and Festuca spp. today developed under continuous spores make a good proxy for past distribution of large herbi-
grazing on the southern slope over the past 2200–2600 years. vores? Quaternary Science Reviews 62: 21–31.
On the basis of our results, we conclude that future changes in Barber KE, Chambers FM, Maddy D et al. (1994) A sensitive
alpine and treeline vegetation in the Retezat Mountains will be a high-resolution record of Late-Holocene climatic change
function of the amplitude and rapidity of climate change com- from a raised bog in northern England. The Holocene 4(2):
bined with forest and land-use management practices. Global 198–205.
warming with decreasing land use of the high-elevation ecosys- Bennett KD (2005) Documentation for Psimpoll4.25 and
tems would certainly lead to the upslope movement of both Picea Pscomb1.03: C Programs for Plotting Pollen Diagrams and
abies and P. cembra. However, compared with the Alps, the alti- Analysing Pollen Data. Department of Earth Sciences, Uni-
tudinal increase in the treeline would be expected to be less, in the versity of Uppsala. Available at: http://www.chrono.qub.ac.uk.
order of 100–150 m. Changing land use with reduced grazing Bennett KD and Willis KJ (2001) Pollen. In: Smol J, Birks HJ,
intensity and increased recreational use may also affect the vege- Last W, et al. (eds) Tracking Environmental Change Using
tation of the Bucura valley at lower elevations, but further Lake Sediments. Dordrecht: Springer, pp. 5–32.
research is needed to provide an overview for the whole region. Berthel N, Schwörer C and Tinner W (2012) Impact of Holocene
climate changes on alpine and treeline vegetation at Sanetsch
Acknowledgements Pass, Bernese Alps, Switzerland. Review of Palaeobotany and
The authors are grateful to Jordan Fevre for analysing the mac- Palynology 174: 91–100.
rocharcoal from Lake Lia. They appreciate the archaeological Beug H-J (2004) Leitfaden der pollenbestimmung fur Mitteleu-
information from Hunedoara county (Romania) from Dr IV Fe- ropa und angrenzende Gebiete. Munich: Verlag Friedrich
rencz (Muzeul Civilizatiei Dacice si Romane Deva, Department Pfeil, 542 pp.
of Archaeology, Deva). They are grateful to HJB Birks for help- Birks HH and Birks HJB (2000) Future uses of pollen analysis
ful comments. They also thank the Retezat National Park sup- must include plant macrofossils. Journal of Biogeography
port and the people who helped them in the field (Gergely Boros, 27(1): 31–35.
Krisztina Buczkó, Péter Eszenyi, Edit Hathi, Tamás János Juhász, Birks HH and Bjune AE (2010) Can we detect a west Norwegian
István Konyhás, Barnabás Körmöndi, Máté Mile, Judit Nédli, Ist- tree line from modern samples of plant remains and pollen?
ván Papp, Gellért Puskás, Csaba Schnitchen, László Somay, Sz- Results from the DOORMAT project. Vegetation History and
abolcs Struba, Andor Umman, Gábor Umman, Hajnalka Veres). Archaeobotany 19(4): 325–340.
IV, EKM and GJ did the plant macrofossil and macrocharcoal Birks HH, Jones VJ, Brooks SJ et al. (2012) From cold to cool
analysis of Lake Bucura; IO and HHB did the macrofossil analy- in northernmost Norway: Lateglacial and early Holocene
sis of Lake Lia and IO calculated the age–depth models of both multi-proxy environmental and climate reconstructions from
sediment sequences. IP did the pollen, stomata and microcharcoal Jansvatnet, Hammerfest. Quaternary Science Reviews 33:
analyses of Lake Lia and Lake Bucura. LOI measurement was 100–120.
done by KH on both sediment cores. MB and EKM carried out Blaauw M (2010) Methods and code for ‘classical’ age-modelling
the sediment coring. WF contributed to the macrocharcoal data of radiocarbon sequences. Quaternary Geochronology 5(5):
analyses. TB drew the 3D model of the study site in Figure 1. CD 512–518.
provided valuable information about the study site. IV, EKM, IP, Blaauw M and Christen JA (2013) Bacon Manual e V2. 2. Avail-
WF and HHB wrote the manuscript. The project that financed this able at: http://chrono.qub.ac.uk/.
research was led by EKM. The authors thank the two anonymous Bojnanský V and Fargašová A (2007) Atlas of Seeds and Fruits
reviewers for their constructive comments. All authors read and of Central and East-European Flora: The Carpathian Moun-
approved the final manuscript. tains Region. Dordrecht: Springer, 1079 pp.
Bortenschlager S (1991) Das höchst gelegene Moor der Ostalpen’
Funding Moor am Rofenberg’ 2760 m. Dissertationes Botanicae 196:
This study was supported by the Hungarian National Scientific 329–334.
Fund (OTKA NF 101362), the Humboldt fellowship to EKM and Borza A (1934) Studii fitosociologice in Munţii Retezatului.
was financially supported by GINOP-2.3.2-15-2016-00019. This Buletinul Grădinii Botanice şi al Muzeului Botanic de la Uni-
study was also supported by a short-term scientific mission pro- versitatea din Cluj 14: 1–84.
vided by the INTIMATE COST Action ES0907 awarded to IV Buczkó K, Magyari EK, Braun M et al. (2013) Diatom-inferred
and the University of Bergen, Norway, to IO. This is MTA- lateglacial and Holocene climatic variability in the South
MTM-ELTE Paleo contribution no. 231. Carpathian Mountains (Romania). Quaternary Interna-
tional (Advancing Pleistocene and Holocene climate change
research in the Carpathian-Balkan region) 293: 123–135.
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