bodnariuc-qsr-2002-21-1465

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Quaternary Science Reviews 21 (2002) 1465–1488 Holocene vegetational history of the Apuseni mountains, central Romania A. Bodnariuc a,b , A. Bouchette a , J.J. Dedoubat a , T. Otto a , M. Fontugne c , G. Jalut a, * a Laboratoire d’Ecologie Terrestre, Universit ! e Paul Sabatier, UMR 5552, 39, All ! ees. Jules Guesde, 31062 Toulouse Cedex 4, France b Faculty of Geology, Babes-Bolyai University, 1 Kogalniceanu str., Cluj Napoca 3400, Romania c Laboratoire des Sciences du Climat et de l’Environment, Laboratoire mixte CNRS - CEA, Avenue de la Terrasse, 91198 Gif sur Yvette, France Received 17 October 2000; accepted 30 September 2001 Abstract From palynological investigations in the Apuseni mountains (Transylvania, Romania), a chronology of forest development is proposed. The data are compared with others from the Romanian Carpathians and the surrounding countries. After their appearance between 11,200 and 10,190 cal BP, Picea and Corylus were dominant up to 6450 cal BP. The extension phases of Carpinus, Fagus and Abies, respectively, began at ca 6450 BP, 4500 BP and 4100 cal BP. Occurrences of their pollen are recorded from about 7800 cal BP. First evidence of human impact appeared during the 7800–7425 cal BP period and first cultivations at ca 6820 cal BP. Between 4500 and 2750 cal BP deforestation and agriculture were limited, but increased around 1935 and 695–660 cal BP. r 2002 Elsevier Science Ltd. All rights reserved. 1. Introduction In Eastern Europe, in the region of contact between the subcontinental, mediterranean and steppic regions, Romania represents a particularly interesting field of investigation for the Lateglacial and Holocene studies of vegetation and climate. Geomorphomogical studies show that despite their altitude (maximum 2500 m), the Carpathians and the Apuseni mountains (Fig. 1) were not strongly affected by glaciers during the last glacial episode (Ficheux, 1996). Only traces of nivation niches oriented to the east, in the lee of dominant winds from the west, exist on the crests of Biharia or Vladeasa (Ficheux, 1996). In the Apuseni mountains, during the last cold and dry periods, the complexity of the relief partly linked to the extension of the karstic zones, probably favoured the persistence of sheltered areas favourable for the trees that form the present forests. This country was also occupied early by humans (communities of Carcea-Gura Baciuli-Ocna Sibiului type), moving into the Transylvanian highlands, from the beginning of the Neolithic (ca 8000 cal BP) onwards (Lazarovici, 1993; Mantu, 1998). Numerous palynological studies have been carried out in Romania starting with those of Pop (1929, 1934, 1942, 1962) then Ciobanu (1948, 1958, 1965). They were based on fundamental studies of plant distribution (Donita et al., 1960; Donita, 1964, 1965; Georgescu and Donita, 1965) and peat bogs and humid zones (Pop, 1960). These studies were compared with numerous other works on Eastern and Western Europe, and a general schema for the Lateglacial and Holocene forest and climate history was proposed (Pop, 1929, 1932, 1934, 1942). Unfortunately, until the recent works of Farcas et al. (1999) and R . osch and Fischer (2000), no 14 C dates were available, and in most cases, the sampling intervals were too large to allow chronological problems to be solved. Except the study of R . osch and Fisher, there are no references to human impact. The goal of the present palynological studies is to establish a chronology of the forest history in the western Romanian Carpathians. This is a key zone for further comparison with the Carpathian mountains and Transylvania. 2. The studied area Romania occupies a transitional position in the vegetational pattern of south-east of Europe. Except in *Corresponding author. Tel.: +33-5-61-55-80-37; fax: +33-5-61-32- 83-82. E-mail address: [email protected] (G. Jalut). 0277-3791/02/$ - see front matter r 2002 Elsevier Science Ltd. All rights reserved. PII:S0277-3791(01)00117-2

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Page 1: Bodnariuc-QSR-2002-21-1465

Quaternary Science Reviews 21 (2002) 1465–1488

Holocene vegetational history of the Apuseni mountains,central Romania

A. Bodnariuca,b, A. Bouchettea, J.J. Dedoubata, T. Ottoa, M. Fontugnec, G. Jaluta,*aLaboratoire d’Ecologie Terrestre, Universit !e Paul Sabatier, UMR 5552, 39, All !ees. Jules Guesde, 31062 Toulouse Cedex 4, France

bFaculty of Geology, Babes-Bolyai University, 1 Kogalniceanu str., Cluj Napoca 3400, RomaniacLaboratoire des Sciences du Climat et de l’Environment, Laboratoire mixte CNRS - CEA, Avenue de la Terrasse, 91198 Gif sur Yvette, France

Received 17 October 2000; accepted 30 September 2001

Abstract

From palynological investigations in the Apuseni mountains (Transylvania, Romania), a chronology of forest development isproposed. The data are compared with others from the Romanian Carpathians and the surrounding countries. After their

appearance between 11,200 and 10,190 cal BP, Picea and Corylus were dominant up to 6450 cal BP. The extension phases ofCarpinus, Fagus and Abies, respectively, began at ca 6450BP, 4500BP and 4100 cal BP. Occurrences of their pollen are recordedfrom about 7800 cal BP. First evidence of human impact appeared during the 7800–7425 cal BP period and first cultivations at

ca 6820 cal BP. Between 4500 and 2750 cal BP deforestation and agriculture were limited, but increased around 1935 and695–660 cal BP. r 2002 Elsevier Science Ltd. All rights reserved.

1. Introduction

In Eastern Europe, in the region of contact betweenthe subcontinental, mediterranean and steppic regions,Romania represents a particularly interesting field ofinvestigation for the Lateglacial and Holocene studies ofvegetation and climate. Geomorphomogical studiesshow that despite their altitude (maximum 2500m),the Carpathians and the Apuseni mountains (Fig. 1)were not strongly affected by glaciers during the lastglacial episode (Ficheux, 1996). Only traces of nivationniches oriented to the east, in the lee of dominant windsfrom the west, exist on the crests of Biharia or Vladeasa(Ficheux, 1996).In the Apuseni mountains, during the last cold and

dry periods, the complexity of the relief partly linked tothe extension of the karstic zones, probably favoured thepersistence of sheltered areas favourable for the treesthat form the present forests.This country was also occupied early by humans

(communities of Carcea-Gura Baciuli-Ocna Sibiuluitype), moving into the Transylvanian highlands, fromthe beginning of the Neolithic (ca 8000 cal BP) onwards(Lazarovici, 1993; Mantu, 1998).

Numerous palynological studies have been carried outin Romania starting with those of Pop (1929, 1934,1942, 1962) then Ciobanu (1948, 1958, 1965). They werebased on fundamental studies of plant distribution(Donita et al., 1960; Donita, 1964, 1965; Georgescuand Donita, 1965) and peat bogs and humid zones (Pop,1960). These studies were compared with numerousother works on Eastern and Western Europe, and ageneral schema for the Lateglacial and Holocene forestand climate history was proposed (Pop, 1929, 1932,1934, 1942). Unfortunately, until the recent works ofFarcas et al. (1999) and R .osch and Fischer (2000), no14C dates were available, and in most cases, the samplingintervals were too large to allow chronological problemsto be solved. Except the study of R .osch and Fisher,there are no references to human impact.The goal of the present palynological studies is to

establish a chronology of the forest history in thewestern Romanian Carpathians. This is a key zone forfurther comparison with the Carpathian mountains andTransylvania.

2. The studied area

Romania occupies a transitional position in thevegetational pattern of south-east of Europe. Except in

*Corresponding author. Tel.: +33-5-61-55-80-37; fax: +33-5-61-32-

83-82.

E-mail address: [email protected] (G. Jalut).

0277-3791/02/$ - see front matter r 2002 Elsevier Science Ltd. All rights reserved.

PII: S 0 2 7 7 - 3 7 9 1 ( 0 1 ) 0 0 1 1 7 - 2

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the mountains, its forests are characterised by thepresence and the dominance of deciduous and thermo-philous trees (Ozenda, 1994). To the east and north-eastare steppes, mixed coniferousFdeciduous forests, de-ciduous forest and forestFsteppe which characterise theEurasiatic Steppic Region (Lavrenko in Walter andStraka, 1970) also called Steppic and Sarmatic Domains(Ozenda, 1994). To the west is the Atlantic Domaindominated by deciduous broad leaved trees. To the southof Romania, the Mediterranean Domain is mainlycharacterised by sclerophylous evergreen trees and shrubs.To the north, the east and the south, the Carpathian

mountains (Fig. 1) form a natural limit to the Hungar-ian plain from which emerge, in its eastern part, theApuseni mountains. (Romanian Western Carpathians).Formerly called Bihor (Ozenda, 1994), the Apusenimountains form a massif about 100 km in diameter, of

medium elevation (maximum 1848m), and occupy acentral position in Transylvania (Ficheux, 1996). Flatsurfaces are rare in the Apuseni mountains and therivers are deeply incised. Four morphological units canbe distinguished: the high Bihor, above 1000m, whichhas a complex geological structure including the karsticregion of Padis; the Metaliferi mountains; the massifsand gulfs of the western slopes, the depression ofHuedin and the Hungarian plain to the west.A cold continental climate characterises the studied

area. Rainfall is abundant from spring to autumn with amarked maximum in summer. Mean annual precipita-tion is about 1400mm and mean annual temperatureabout 4.11C. Winters are cold with absolute minimumtemperature p�301C and summers cool. Due to thecomplexity of the relief of the Apuseni mountains,important local climatic differences exist. To the west,

Fig. 1. Location of the studied area. Cited sites: Romania 1 Mluha, 2 Mohos, 3 Taul Zanogutii, 4 Rachitis, 5 Cica Mica 1, 6 Calimani Exploatare, 7

Iezerul Calimani, 8 Poiana Boilor, 9 Poiana Stiol, 10 Dupa Lunca–Voslobeni, 11 Banat; Bulgaria: 12 Sucho Ezero, 13 Kupena, 14 Tschokljovo;

Ukraina: 15 Dorjok, 16 Ogreev, 17 Kardashinski; Slovakia & Czech Republic; 18 Zlatnicka doline, 19 Tojrohe Pleso, 20 Vracov; Poland: 21 Tarnawa

Wyzna, 22 Szymbark; Hungary: 23 Batorliget; Slovenia: 24 Kaznarice.

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annual precipitation is about 300mm higher than to theeast. The maximum precipitation is concentrated in thecentre-west part of the mountains where the sites studiedare located. The present climate conditions allow thedevelopment of mountain forests in which favourabletopographic and geological conditions permit Sphagnumpeat bogs to develop.

3. The present vegetation

In the Apuseni mountains, forests are essentiallycomposed by Fagus silvatica and Picea abies. The latteris concentrated in the central part of the massif whileFagus silvatica is mainly found around it. However, dueto lumbering in the spruce forest, large areas are nowcolonised by beech which temporarily replaces spruce.Abies is rare. The strong human impact on the mountainforests has determined the extension of meadows whereFestuca rubra, Nardus stricta, Calluna and Calamagros-tis arundinacea are represented (Donita et al., 1960).Meadows with some other species occur on certain highareas. Low altitudes, depressions and corridors arecovered with oak-horbeam forest composed of Quercuspetraea, Quercus robur, Carpinus betulus, Fagus sylvati-ca, Tilia cordata, Acer pseudoplatanus, A. platanoides,Corylus avellana and Fraxinus excelsior. The abundanceof Quercus, Fagus and Carpinus depends on localecological and historical factors. Distant from themassif, thermophilous oak forests with Quercus petraea,

Q. cerris, Q. pubescens, Q. fraineto, Fraxinus ornus,Prunus mahaleb occur on limestone.

4. The selected sites

In the NW of the Apuseni mountains, five cores wereobtained using a Russian sampler (Fig. 2). The altitudeof the sites ranged between 1000 and 1300m a.s.l. All arenow ombrogenic Sphagnum peat bogs.The site of Ic Ponor (alt. 1020m a.s.l.) (Fig. 2) lies on

schists, at the foot of a slope. After a relatively shortlacustrine phase during the early Holocene, peat beganto develop. The site now corresponds to a Sphagnumpeat bog which covers 7 ha on the right side of SomesulCald river, near its confluence with the river Batrana. Tothe south, an inundated zone is colonised by Carexrostrata. The peat bog is surrounded by spruce, andscattered stands of birch (Betula pubescens and Betulapendula) are present in deforested zones. Picea andBetula are also abundant on the peat bog. At its surface,Vaccinium myrtillus and V. vitis ideae form densecommunities and Eriophorum vaginatum is abundant.Sphagnum sp. covers the whole surface of the peat bogbut Polytrichum is also present as well as Empetrumnigrum, cited by Pop (1960). Because of the large size ofthe peat bog, two cores were taken: Ic Ponor I (295 cmlength) (Fig. 3) at the summit of the peat bog, in thedeepest zone, and Ic Ponor II (165 cm length) (Fig. 4) inthe NW margin.

Fig. 2. Location of the local cited and studied sites: 1 Mlastina lui Neag, 2 Izbucu I, 3 Izbucu II, 4 La Lacuri, 5 Pietrele Onachii, 6 La Mlastina, 7 La

Mol, 8 Molhasu de la Calatele, 9 Dealu Negru, 10 Dambu Negru, 11 Ciurtuci, 12 Baita, 13 Ic Ponor I & II, 14 Bergerie, 15 Cimeti"ere, 16Padis.

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Fig.3.PollendiagramfromIcPonor1.

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Fig.4.PollendiagramfromIcPonor2.

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The three other sites are situated to the west, in thekarstic zone of Padis.Padis (alt. 1240m a.s.l.) (Fig. 2) is a Sphagnum peat

bog in one of the numerous sink holes present on thekarstic plateau. Some of these sink holes are small pondscolonised by hydrophilous communities. Others, such asthat of Padis, have been transformed into Sphagnumpeat bogs colonised by stands of spruce. The dominantshrubs are Vaccinium myrtillus and V. vitis idae.Dechampsia fluxosa, Carex echinata and C. rostrataare also well represented. Sphagnum and Polytricum arethe dominant mosses. Flat surfaces with meadowssurround the site. Picea forest occupies the hills. Incleared areas, stands of Fagus frequently occupy the hilltops. The core was collected at the centre of the peatbog. The bottom of the sequence is clay poor in organicmatter (Fig. 5).The peat bog of Cimeti"ere (alt. 1280m a.s.l.) (Fig. 2)

is situated on a slope. It has a NW exposure and islocated near a ridge in a zone strongly affected byforestry. It was surrounded by old spruce partlydestroyed by a storm. The peat bog belongs to a largepeaty complex where Sphagnum is dominant in mostplaces. Other species such as Eriophorum vaginatum,Deschampsia flexuosa and Vaccinum myrtillus areabundant or frequent while Vaccinium vitis ideae,Homogyne alpina and Carex echinata are present butrare. The base of the core is formed of coarse gravel witha low pollen content and corroded grains of uncertainorigin (Fig. 6). For these reasons, only the Sphagnumpeat deposit was taken into account for the interpreta-tion of the pollen data.The peat bog of Bergerie (alt. 1400m a.s.l.) (Fig. 2) is

a Sphagnum peat bog. Locally Sphagnum sp. dominates,in association with Carex rostrata, Eriophorum vagina-tum, Carex echinata, Juncus effusus and J. conglomer-atus. The bore was performed under 25 cm of water. Thebottom of the sequence is argilous and poor in organicmatter (Fig. 7). The site is situated above but near to thepresent upper limit of the spruce forest. The surroundingarea is devoted to pasture. Juniper (Juniperus communis)and young Picea abies colonise the surroundingmeadows.

5. Description of the cores

5.1. Ic Ponor I and II (Figs. 3 and 4)

Despite their different depths, the two cores showsimilar sedimentary facies. The bottom is composed of asandy-clay deposit showing thin intercalated peat layers.This lacustrine phase corresponds to the period 10,190–9660 cal BP (levels 298–280, Ic Ponor I; 170–140, IcPonor 2). Then, the sediments are covered by a charcoallayer with some bark fragments, leaves and seeds. The

abrupt transition corresponds to synchronous changesin pollen percentages: decrease in Betula, increase inCorylus, particularly at Ic Ponor I (Fig. 3). From theavailable radiocarbon dates and linear interpolations,the age of this transition can be estimated at ca 9650–9550 cal BP.Above, up to ca 4500 cal BP (levels 280–36, Ic Ponor I;

145–38, Ic Ponor II) a Sphagnum peat is observed. At IcPonor 1, at level 44, charcoal is present. In the twosequences, between levels 36–27 (Ic Ponor 1) and 38–20(Ic Ponor 2), mineral deposits with sand and clayindicate a flooded phase. At levels 22 (Ic Ponor 1) and15 (Ic Ponor 2), a poorly decomposed Sphagnum peatcontains charcoals. Then a recent Sphagnum peat coversthis charcoal layer. The comparison of the cores as wellas the palynological data shows that deposits betweenlevels 36–27 (Ic Ponor 1) and 38–20 (Ic Ponor 2) arecontemporaneous. However, the radiocarbon dates oflevels 34 (56807110BP, Ic Ponor 1) and 35(64607110BP, Ic Ponor 2) are different (Table 1). Inthe two cores, the beginning of this lacutrine depositcorrespond to a synchronous increase in Fagusand Abies pollen percentages. At Bergerie (Fig. 7) andPadis (Fig. 5), such increases are not synchronous anddated 4050780BP (4500 cal BP) and 37507100BP(4100 cal BP), respectively (Table 1). Out of the area,in the southwestern Carpathians (Banat mountains), thetwo events seem synchronous and are dated3880760BP (R .osch and Fischer, 2000). In all cases,these events occurred around 4500–4370 cal BP. Conse-quently, it can be assumed that the ages of levels 34 and35 of Ic Ponor 1 and 2 are too old. These dates, ageingas well as the sedimentary facies, suggests that the datedplant remains contained in the sediments were reworkedand that around levels 35–30 there is a hiatus whichmight be of about 2000yr. At Ic Ponor 1, the estimatedage of level 40 obtained by linear interpolation (6000BP:6820 cal BP) reinforces this hypothesis as does palyno-logical data. By comparison with pollen data fromBergerie and Padis, the age of the layer 35–12 can beestimated. In these sites, between 4200 cal BP and680 cal BP, the Carpinus percentages are frequently>10%. At Ic Ponor, above level 35, they do not exceed5–6%. For this reason, it can be assumed that in the twocores, the deposits between 35 and 12 cm are youngerthan 680 cal BP. Above, the pollen content of theSphagnum corresponds to the present and recentperiods.

5.2. Padis (Fig. 5)

Between levels 95 and 80, we observe a clay depositwith gravel and sand. It is covered, up to level 27, by aSphagnum peat, poorly decomposed between levels 27and 9. Sphagnum covers the site.

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Fig.5.PollendiagramfromPadis.

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Fig.6.PollendiagramfromCimeti" ere.

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Fig.7.PollendiagramfromBergerie.

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5.3. Cimeti "ere (Fig. 6)

White clay with gravel and poor in organic matterforms the bottom of the sequence. Plant macro-remainsare observed at levels 125 and 118. Between levels 90 and53 the content in organic matter slowly increases andsome undetermined plant macro-remains occur. Be-tween levels 53 and 42.5 a sandy sediment is observedand the content in organic matter slowly increases. Aftera thin grey sand layer present between levels 42.5 and 40,Sphagnum peat appears between levels 40 and 15. It iscovered by Sphagnum. This diversity of facies suggestsfrequent changes in the sedimentation and erosionprocesses. The strong variations in percentages of Pinusand Corylus between levels 130 and 80 are not observedin the other sites and support this hypothesis.

5.4. Bergerie (Fig. 7)

Between levels 230 and 90, a lacustrine depositpresents three facies: 230–190, sandy clay deposit withsome gravel; 190–145: clay with somes gravel; 145–90:clay with plant remains. Peat appears at level 90. Theabrupt sedimentary transition does not correspond tostrong pollen percentages variations which mightindicate a possible gap of sediment. If such an event

did occur, it was short and without any majorconsequences on the pollen representation. Betweenlevels 90 and 25 peat decomposition regularly. Sphag-num is observed between levels 20 and 12.

6. Pollen counting and representation of the pollen

diagram

To extract pollen grains and spores, samples of onecubic centimeter were taken using a calibrated sampler.Classical procedures were used to eliminate organicmatter and mineral fraction. To calculate the pollenconcentration, the slides were prepared according toCour (1974). The pollen concentrations are expressed asnumber of pollen grains per cm3. For each level,between 300 and 400 grains were counted, whichrepresents a statistically significant sample. After count-ing, each slide was checked for rare pollen grains.Thousands of pollen grains and spores were thusobserved per slide.Percentages were calculated from a pollen sum

including the pollen types of all the trees, shrubs andherbs in the region. Local pollen grains of hydrophilousand hygrophilous taxa were excluded, including Cyper-aceae, as well as spores of Ferns and Mosses whose

Table 1

Radicarbon dates used in pollen diagramsa

Depth (cm) Material 14CBP, sigma Ref. Lab. Ages (cal BP) d13C (%)

Berg.

95–100 Sphagnum peat 725785 Gif-11130 793–538 �25.9145–150 Sphagnum peat 3720760 Gif-11131 4240–3891 �26.19157–160 Plant macro-remains 4050780 GifA-99221 4825–4379

165–170 Sphagnum peat 56657120 Gif-11133 6691–6272 �26.88190–200 Plant macro-remains 6680780 GifA-99475 7664–7430

215–230 Plant macro-remains 70107182 GifA-99476 8177–7562

Pad.

10–15 Sphagnum peat modern Gif-11127 �26.2730–35 Sphagnum peat 445780 Gif-11128 560–308 �25.875–80 Sphagnum peat 37507100 Gif-11135 4411–3864 �26.9885–95 Sphagnum peat 4595765 Gif-11129 5470–4046 �26.8

Ic I

34 Plant macro-remains 56807110 GifA-100143 6686–6281

70 Plant macro-remains 6190790 GifA-100146 7270–6854

190 Plant macro-remains 6870790 GifA-100148 7865–7570

292–295 Plant macro-remains 8990780 GifA-99669 10268–9888

Ic II

35 Plant macro-remains 64607110 GifA-100144 7571–7177

55 Plant macro-remains 6980790 GifA-100145 7964–7659

160 Plant macro-remains 8770790 GifA-100147 9969–9550

C

90–100 Plant macro-remains 78107110 GifA-99220 8814–8406

aIc I: Ic Ponor 1; Ic II: Ic Ponor 2; Pad.: Padis; C: Cimeti"ere; Berg.: Bergerie.

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representation is frequently high and irregular (Janssen,1973). The percentages of the excluded taxa werecalculated from the sum of the other pollen grains. Inthe pollen diagrams, only the most significant taxa areindicated.Each pollen diagram was divided into Local Pollen

Assemblage Zones (LPAZ) as defined by Cushing(1963), Berglund and Ralska-Jasiewiczowa (1986).From comparisons between the pollen diagrams, Re-gional Pollen Assemblage Zones (RPAZ) were thendefined

7. Radiocarborn dates

In many cases, the dated samples were extracted fromSphagnum peat, which excludes the possibility of ageingby hard water effect (Table 1). At Padis, Cimeti"ere andBergerie, low level of organic matter at the base of thecores required p15 cm of sediment per date (Table 1).Peat samples at Padis and Bergerie ranged between 3and 5 cm. At Ic Ponor, all the dates were obtained byAMS using unidentified terrestrial plant macro-remainsfrom thin layers (Table 1).Additional estimated ages were obtained by compar-

ing the neighbouring sites and from linear interpola-tions. Radiocarbon dates were calibrated using theradiocarbon calibration program REV 4.3 (Stuiveret al., 1998). In the text, the dates are expressed incal yr BP. Their consistency is discussed in the followingchapters.

8. The local and regional pollen assemblage zones

The pollen diagrams are divided into LPAZ: Ic Ponor1, 13 LPAZ (Fig. 3, Table 2); Ic Ponor 2, 13 LPAZ(Fig. 4, Table 3); Padis, 12 LPAZ (Fig. 5, Table 4);Cimeti"ere, 13 LPAZ (Fig. 6, Table 5) and Bergerie, 13LPAZ (Fig. 7, Table 6).From the LPAZ described above and using the

sedimentary data, the local available radiocarbon datingand estimated ages obtained by interpolation, 9 RPAZ(RPAZ) were identified (Table 7):RPAZ I: Betula–Picea–Ulmus–Corylus–Pinus (10,190–

9660 calBP), only represented in Ic Ponor 1 and 2.RPAZ 2: Corylus–Picea–Ulmus–Fraxinus–Alnus

(9660–7700 cal BP), only represented in Ic Ponor 1and 2.RPAZ 3: Picea–Corylus–Ulmus (7700–7250 cal BP),

common to Ic Ponor 1 and 2 and Bergerie. Fagus pollenis rare but regularly present.RPAZ 4: Picea–Corylus–Ulmus–Alnus (7250–

6600 cal BP), common to Ic Ponor 1 and 2 and Bergerie.Pollen of Fagus, Carpinus and Abies is rare but present.

RPAZ 5: Picea–Corylus–Carpinus (6600–4500 cal BP),common to Cimeti"ere, Bergerie and the basis of Padis.Presence of Fagus and Abies.RPAZ 6: Picea–Carpinus–Fagus (4500–4100 cal BP),

not represented in Ic Ponor 1 and 2 but common toPadis, Cimeti"ere and Bergerie.RPAZ 7: Fagus–Picea–Carpinus–Abies (4100–1940

cal BP), common to Padis, Cimeti"ere and Bergerie.RPAZ 8: Fagus–Picea–Carpinus–Abies–Poaceae

(1940–480 cal BP), common to Padis, Cimeti"ere andBergerie.RPAZ 9: Fagus–Picea–Carpinus–Abies–Cerealia (480

cal BPFPresent), common to Ic Ponor 1 and 2, Padis,Cimeti"ere and Bergerie.

9. Discussion

The comparison of the five pollen diagrams showsthat the Lateglacial and the beginning of the Postglacialperiod between 11,000BP and 10,290 cal BP are notrepresented in the sites studied. This does not allow us todescribe the role of Pinus and Betula or to date thebeginning of the Picea curve.The present data concern the last 10,200yr. At Ic

Ponor, between about 10,200 and 9700–9600BP, duringRPAZ 1 (Table 7), Betula played a major role. Then,during RPAZ 2, it regressed and was replaced byCorylus and Picea. This Betula extension might be theconsequence of fires which occurred around 10,200–9650 cal BP as indicated by the presence of charcoal (IcPonor I: levels 290–285; Ic Ponor 2: level 165). Charcoalof Picea is present (M. Thinon, pers. comm.) whichdemonstrates the local presence of this tree at that time.The surrounding Picea–Corylus forest was probablystrongly modified and birch colonised. Then Corylusand Picea recovered. Except in RPAZ 1 and 2, Pinusand Betula were poorly represented everywhere. At alower stage, the oak forest was regularly representedwith Ulmus values reaching 10%. But at the altitude ofthe site, between 10,200 and 6800 cal BP (mid RPAZ 4,Table 2), Corylus and Picea were the regional dominantspecies. This period corresponds to the Picea–Corylus–Quercetum mixtum defined by Pop (1932). Goodcorrelations exist between the two cores of Ic Ponor,particularly from 10,200 to 6800 cal BP, and a referencelevel is recorded ca 7700 cal BP (6870790BP Ic Ponor 1;6980790BP, Ic Ponor 2) when the continuous Faguscurve begins.A that time, comparisons with the pollen diagram of

Bergerie becomes possible. The bottom of this sequence,dated 7010780BP (7800 cal BP), does not containpollen grains of Fagus. For this reason it might belongto upper RPAZ 2. Fagus pollen appears above, before7600–7500 cal BP (6680780BP) and the first centi-meters of the core belong to the transition between

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RPAZ 2 and 3. In RPAZ 3 most of the curves ofBergerie are similar to those of Ic Ponor 1 and 2, exceptthat of Picea. This tree is well developed at Ic Ponorfrom 10,200 cal BP upward, but it extends to Bergerieonly from 7800 cal BP. This will be discussed below.Because of the sedimentary characteristics of the cores

at Ic Ponor around 6600–6500 cal BP, comparisonsbetween Bergerie and Ic Ponor 1 are limited to RPAZ3 and 4. Using estimated dates obtained from inter-polation at Ic Ponor 1 and Bergerie, the limit betweenthe two phases might be situated around 7250 cal BP,

when Picea values increase at Bergerie and those ofFagus at Ic Ponor 1.At Bergerie, the lower and upper limits of RPAZ 4 are

defined by the extension of Picea then Carpinus at 7250and 6600–6400 cal BP (5665780BP), respectively.The Carpinus extension is also a good reference event

allowing a correlation to be established between thesequences of Cimeti"ere and Bergerie. At Cimeti"ere,before this event, in addition to the description of thecore, several palynological observations suggest distur-bances in the sedimentation processes: the abrupt

Table 2

Description of the local pollen assemblage zones in the Ic Ponor 1 profile

LPAZ/Depth (cm) LPAZ name Main features of the LPAZ

Ic 1–13 0–5 Picea–Poaceae–Betula–Cerealia Decrease in Poaceae and Picea, slight increase in Pinus and

Fabaceae, presence of Centaurea cyanus

Ic 1–12 5–15 Picea–Poaceae–Cerealia Decrease in Picea and Fagus, increase in Fabaceae.

Upper limit: decrease in Poaceae and Picea

Ic 1–11 15–20 Picea–Fagus–Corylus–Poaceae–Cerealia Decrease in Corylus and Alnus, increase in Picea, Fagus,

Poaceae, small increase in Carpinus. First presence of Cerealia

Upper limit: decrease in Fagus, Picea and Carpinus

Ic 1–10 20–25 Picea–Corylus–Fagus–Poaceae Decrease in Picea, slight decrease in Abies, increase in Poaceae,

maximum of Ericaceae

Upper limit: increase in Poaceae, Fagus and Picea, decrease in

Ericaceae

Ic 1–9 25–35 Picea–Corylus–Fagus Strong decrease in Picea, increase in Fagus, Carpinus and Alnus.

Increase in Poaceae and Ericaceae, occurrences of Plantago

lanceolata, decrease in Ulmus

Upper limit: decrease in Picea and Abies, slight increase in

Chenopodiaceae

Ic 1–8 35–45 Picea–Corylus Increase in Picea, decrease in Corylus and Quercus. Small

increase in Fagus and Abies

Upper limit: decrease in Picea, increase in Carpinus and Fagus

Ic 1–7 45–55 Corylus–Picea Increase in Corylus and Quercus, strong decrease in Picea

Upper limit: increase in Picea, decrease in Corylus and Quercus

Ic 1–6 55–70 Picea–Corylus–Ulmus Increase in Picea, decrease in Fraxinus. Regular presence of

Carpinus and Abies with low values. Fagus regularly present but

rare. Small increase in Artemisia and Humulus–Cannabis type

Upper limit: decrease in Picea, increase in Corylus

Ic 1–5 70–95 Picea–Corylus–Fraxinus Decrease in Corylus. Small increase in Fagus and decrease in

Ericaceae

Upper limit: regular presence of Carpinus and Abies, increase in

Picea

Ic 1–4 95–190 Picea–Corylus–Ericaceae Dominance of Picea and Corylus. Regular presence of Fagus

with low values. Occurrences of Abies and Carpinus. Occur-

rences of Chenopodiaceae, Urticaceae and Rumex. Increase in

Humulus–Cannabis type at the end of the phase

Upper limit: increase in Fagus, decrease in Corylus and

Ericaceae.

Ic 1–3 190–220 Corylus–Picea–Ulmus–Fraxinus–Ericaceae Dominance of Corylus and Picea, increase in Ericaceae and

Humulus–Cannabis type. Decrease in AP/T values

Upper limit: decrease in Corylus, regular presence of Fagus

Ic 1–2 220–280 Corylus–Picea–Ulmus–Fraxinus Abrupt fall in Betula values, increase in Corylus and Picea,

occurrences of Rumex, sporadic presence of Fagus and Carpinus

Upper limit: fall in Corylus, increase in Ericaceae andHumulus–

Cannabis type

Ic 1–1 280–295 Betula–Picea–Ulmus–Corylus–Pinus Abundance of Betula, Picea between 10% and 18%, decrease in

Pinus

Upper limit: fall in Betula values, increase in Corylus and Betula

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change in percentages of Pinus, Corylus and Poaceae atthe clay–peat transition; the strong variations in thepercentages of Pinus and Corylus between levels 130 and90 and, at the bottom of the sequence, the high values ofCarpinus and Fagus not observed at Bergerie and IcPonor. Both the situation of the peat bog at the summitof a western slope exposed to strong winds and rainfallsand the local human impact explain that run-off hasaffected the sedimentary processes. For these reasons,deposits from the bottom to level 40, were considered asunsuitable for radiocarbon dating and those betweenlevels 130 and 80 were rejected for paleoecologicalreconstruction. The pollen diagram of Cimeti"ere wassubdivided using data from Bergerie and Padis.

During the early Holocene represented by RPAZ 1–4,sporadic occurrences of Carpinus, Fagus and Abies areobserved. At Ic Ponor, the first Carpinus pollen wasnoted near 9500–8900 cal BP then regularly observed ca7800 cal BP at Bergerie and 7700–7600 cal BP at IcPonor. At Ic Ponor and Bergerie, Abies pollen occurredaround 7600–7500 cal BP. Fagus is found around 9400–9000 cal BP at Ic Ponor and from 7800 cal BP atBergerie.These occurrences suggest the existence of scattered

stands situated at low and mid elevation during theearly Holocene. It can be assumed that they probablyoriginate from regional Glacial and Lateglacialrefuges.

Table 3

Description of the local pollen assemblage zones in the Ic Ponor 2 profile

LPAZ/Depth (cm) LPAZ name Main features of the LPAZ

Ic 2–13 5–15 Fabaceae–Rosaceae–Picea–Corylus–Cerealia Strong decrease in Picea and Corylus. Slight decrease in

Carpinus, Fagus and Poaceae. Abundance of Fabaceae.

Urticaceae and Centaurea cyanus well represented. Presence of

Cerealia

Ic 2–12 15–20 Picea–Corylus–Fagus–Carpinus Strong decrease in Corylus, increase in Picea and Carpinus,

Poaceae and Ericaceae

Upper limit: decrease in AP/T ratio, Picea, Betula and Corylus

Ic 2–11 20–30 Picea–Corylus–Fagus–Alnus Increase in Alnus, slight increase in Cannabis–Humulus type,

occurrences of Urticaceae

Upper limit: decrease in Corylus, increase in Carpinus and Picea

Ic 2–10 30–35 Picea–Corylus–Fagus Increase in Fagus, Alnus, Betula and Picea, decrease in Corylus,

Ulmus and Fraxinus. Beginning of the regular presence of

Carpinus and occurrences of Abies. Increase in Plantago

lanceolata, Chenopodiaceae and Poaceae

Upper limit: increase in Alnus and Picea, decrease in Fagus

Ic 2–9 35–40 Picea–Corylus–Ulmus–Quercus Decrease in Picea, increase in Corylus, Quercus, Ulmus and

Betula

Upper limit: decrease in Corylus, Ulmus and Quercus, increase in

Fagus

Ic 2–8 40–55 Picea–Corylus Picea dominant, regular presence of Fagus with low values.

Slight increase in Poaceae then Ericaceae, first occurrences of

Abies

Upper limit: decrease in Picea, increase in Corylus

Ic 2–7 55–75 Picea–Corylus–Ulmus–Fraxinus Increase in Picea, synchronous decrease in Corylus. Slight

increase in Poaceae then Artemisia. Occurrences of Fagus

Upper limit: beginning of the Fagus curve, decrease in Ulmus

Ic 2–6 75–115 Corylus–Picea–Ulmus–Fraxinus Peaks of Picea in the first part of the zone, Corylus stable, AP/T

values at their maximum

Upper limit: increase in Picea, decrease in Corylus and Ulmus

Ic 2–5 115–135 Corylus–Picea–Ulmus–Fraxinus Dominance of Corylus, Picea stable around 20%, decrease in

Poaceae. Peak of Pinus at the end of the phase. First

occurrences of Carpinus and Fagus

Upper limit: increase in Picea, small decrease in Betula

Ic 2–4 135–145 Corylus–Picea–Ulmus–Betula Increase in Corylus, Fraxinus and Picea, decrease in Betula

Upper limit: decrease in Poaceae and Fraxinus

Ic 2–3 145–150 Corylus–Betula–Picea–Ulmus Increase in Corylus, decrease in Picea and Betula. Small

increase in Alnus

Upper limit: decrease in Betula, increase in Picea and Corylus

Ic 2–2 150–160 Betula–Picea–Corylus Decrease in Betula, slight increase in Corylus, increase in Picea,

small increase in Poaceae

Upper limit: decrease in Betula and Picea, increase in Corylus

Ic 2–1 160–165 Betula–Picea–Ulmus–Corylus–Pinus Increase in Betula, decrease in Picea and Pinus

Upper limit: decrease in Betula, increase in Picea

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From RPAZ 5–9 (ca 6450 cal BP–Present), the pollendiagrams of Bergerie, Padis and Cimeti"ere can becompared.At Padis, between 4650 and 4100 cal BP, in the upper

part of RPAZ 5, the percentages of Carpinus are low(2%) when comparing with Bergerie (near 5%) andCimeti"ere (near 10%). At Padis, Cimeti"ere and Bergerie,the increase in Fagus and Carpinus and the synchronousdecrease in Corylus characterise the beginning of theRPAZ 6 dated ca 4500 cal BP (4050780BP) at Bergerie.This age, deduced from that of levels 160–155 fromBergerie (4050780BP), was preferred to that of thebottom of Padis (4595765BP, 5300 cal BP) for three

reasons: at Padis the thickness of the sample is 15 cmand at Bergerie only 5 cm; at Padis the dated sedimentconcerns material situated below the beginning of theFagus curve; comparison between the dates obtained forthe Fagus expansion at Bergerie (4050780BP) and inthe southwestern Carpathians, in Banat mountains(3880760BP; R .osch and Fischer, 2000), are close to4500 cal BP.The beginning of RPAZ 7 corresponds to the Abies

development dated ca 4200 cal BP (3720760BP) atBergerie. The end of RPAZ 7 is not dated. The proposedlimit at 1940 cal BP is based on an interpolation betweenthe radiocarbon dating of Bergerie (3720760 and

Table 4

Description of the local pollen assemblage zones in the Padis profile

LPAZ/Depth (cm) LPAZ name Main features of the LPAZ

Pad. 12 5–11 Poaceae–Fagus–Picea–Carpinus–Corylus–Quer-

cus

Small decrease in Poaceae, Picea and Carpinus, slight increase

in Fagus, Quercus, Corylus and Pinus

Pad. 11 11–15 Poaceae–Fagus–Picea–Carpinus–Cerealia Decrease in Poaceae and synchronous increase in Fagus, Picea,

Carpinus, Quercus and Pinus

Upper limit: decrease in Poaceae, increase in Fagus

Pad. 10 15–20 Poaceae–Fagus–Picea–Cerealia Abrupt increase in Poaceae and synchronous decrease in Fagus

and Picea. Cerealia, Plantago, Rumex and Chenopodiaceae well

represented,

Upper limit: decrease in Poaceae, increase in Fagus and Picea

Pad. 9 20–25 Fagus–Poaceae–Picea–Corylus–Quercus Increase in Picea, decrease in Quercus, slight increase in Fagus,

decrease in Chenopodiaceae

Upper limit: increase in Poaceae, decrease in Fagus and Picea

Pad. 8 25–30 Fagus–Poaceae–Picea Decrease in Picea and Abies, increase in Quercus and Betula,

Poaceae, Chenopodiaceae, Rumex, Plantago and Cerealia

Upper limit: slight increase in Fagus and Picea, decrease in

Chenopodiaceae

Pad. 7 30–40 Fagus–Picea–Abies Fall in Carpinus, decline in Fagus, increase then decline in Picea

and Abies. Increase in Chenopodiaceae at the end of the phase

Upper limit: decline in Picea and Abies, increase in Poaceae and

Chenopodiaceae

Pad. 6 40–50 Fagus–Picea–Carpinus–Poaceae–Abies Optimum of Fagus, small variations in Picea and Carpinus and

increase in Poaceae and Corylus. Small increase in Artemisia

Upper limit: decrease in Fagus and Carpinus, increase in Picea

and Abies

Pad. 5 50–55 Fagus–Picea–Carpinus–Abies Increase in Picea, Carpinus and Chenopodiaceae. Small

decrease in Abies and small increase in Fagus

Upper limit: decrease in Carpinus and Picea, increase in

Poaceae

Pad. 4 55–70 Fagus–Picea–Abies–Carpinus Small increase in Abies and Fagus, fluctuations of Carpinus and

Corylus, small increase in Chenopodiaceae. Beginning of the

AP/T decline at the end of the phase

Upper limit: Increase in Picea, Carpinus and Chenopodiaceae

Pad. 3 70–80 Corylus–Carpinus–Fagus–Picea–Abies Decrease in Corylus, Carpinus Ulmus. Increase in Abies, Fagus

and Picea. Small increase in Juglans

Upper limit: decrease in Carpinus, small increase in Corylus

Pad. 2 80–85 Corylus–Carpinus–Picea–Fagus Decrease in Corylus and increase in Carpinus and Fagus. Small

increase in Quercus and Abies. Presence of Juglans

Upper limit: increase in Abies and Fagus, decrease in Corylus

and Carpinus

Pad. 1 85–90 Corylus–Picea–Poaceae Increase in Corylus,decrease in Poaceae. Carpinus, Fagus and

Abies present but rare

Upper limit: decrease in Corylus, increase in Carpinus and

Fagus

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725785BP) and Padis (37507100 and 445780BP).At Bergerie, Cimeti"ere and Padis this limit correspondsto a decrease in the AP/T ratio and Corylus values.At Bergerie it is also marked by a decrease inCarpinus percentages. At that time, the human impact,described below, is the determining factor of thelandscape history. The limits of the RPAZ 8 and 9correspond to major changes observed in the threesites. However, because of the difference in the local

human impact, complete analogies are difficult to befound.RPAZ 8 begins with a decrease in AP/T and an

increase in the anthropogenic indicators. It ends whenPoaceae values increase. According to the sites, Juglansis regularly present (Cimeti"ere and Padis) and thepercentages of Carpinus, Fagus and Picea decrease. AtPadis, the upper limit of RPAZ 8 is dated ca 500 cal BP(445780BP).

Table 5

Description of the local pollen assemblage zones in the Cimeti"ere profile

LPAZ/Depth (cm) LPAZ name Main features of the LPAZ

Cim. 13 0–5 Poaceae–Fagus–Picea–Carpinus–Corylus Strong decrease in Poaceae. Slight increase in Fagus, Picea,

Abies and Carpinus

Cim. 12 5–10 Poaceae–Fagus–Picea–Carpinus Decrease in Picea, small increase in Fagus, increase in Poaceae,

presence of Vitis, decrease in Cerealia

Upper limit: Fall in Poaceae, increase in Fagus, Abies, Carpinus

and Corylus

Cim. 11 10–15 Fagus–Picea–Carpinus–Abies–Cerealia Decrease in Fagus and synchronous increase in Picea, presence

of Olea and Cerealia

Upper limit: decrease in Picea, increase in Poaceae

Cim. 10 15–30 Fagus–Poaceae–Corylus–Cerealia Scarcity of Carpinus, dominance of Fagus, regular presence of

Juglans and Cerealia. Small increase in Poaceae, Chenopodia-

ceae, Urticaceae, Rumex, Plantago

Upper limit: fall in Fagus, increase in Picea

Cim. 9 30–50 Fagus–Poaceae–Picea–Abies Increase in Fagus, decrease in Carpinus. Increase in Artemisia,

regular presence of Chenopodiaceae, Rumex and Plantago

lanceolata

Upper limit: decline in Picea, beginning of Juglans

Cim. 8 50–65 Corylus–Fagus–Carpinus–Abies Beginning of the Abies curve. Strong increase in Fagus (2–20%)

and synchronous decrease in Corylus (30–10%). Decline in

Ulmus. Small increase in Chenopodiaceae and Artemisia,

presence of Rumex

Upper limit: increase in Fagus, Artemisia and Rumex, decrease

in Corylus

Cim. 7 65–70 Corylus–Carpinus–Picea Increase in Fagus and beginning of the regular presence of

Chenopodiaceae and Urticaceae. Decrease in Corylus

Upper limit: increase in Abies and Fagus, decrease in Corylus

Cim 6 70–80 Corylus–Picea–Poaceae–Carpinus–Ulmus Increase in Carpinus and Alnus, decrease in Ulmus and Poaceae.

Small increase in Picea

Upper limit: increase in Fagus, decrease in Corylus

Cim. 5 80–90 Pinus–Corylus–Poaceae–Picea–Ulmus Strong decrease in Pinus (50–2%) and Filicales, increase in

Corylus, Ulmus, Tilia and Poaceae. Beginning of the curve of

Cannabis–Humulus type

Upper limit: increase in Carpinus, decrease in Poaceae

Cim. 4 90–95 Pinus–Corylus–Picea Strong increase in Pinus, decline in Corylus, Ulmus, Alnus, Picea

and Poaceae

Upper limit: abrupt decrease in Pinus, increase in Corylus and

Poaceae

Cim. 3 95–105 Corylus–Picea–Ulmus Abrupt decrease in Pinus and increase in Corylus. Increase in

Picea and Poaceae. High values of Ulmus (16%)

Upper limit: increase in Pinus, decrease in Corylus Ulmus and

Picea

Cim. 2 105–115 Pinus–Picea Decrease in Picea and Pinus. Increase in Ulmus and high values

(15%) at the end of the phase, increase in Poaceae

Upper limit: decrease in Pinus and Filicales, increase in Corylus

and Picea

Cim. 1 115–130 Corylus–Pinus–Picea–Poaceae Picea at 15%, decrease in Corylus, increase in Pinus

Upper limit: decrease in Pinus and Picea

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RPAZ 9 corresponds to a phase of intensive humanactivity well characterised by the anthropogenic indica-tors. The AP/T value falls and strong deforestation arerecorded.

10. Human impact

The first evidence of the presence of humans isvisible at Ic Ponor around 7800 cal BP. At that time

Table 6

Description of the local pollen assemblage zones in the Bergerie profile

LPAZ/Depth (cm) LPAZ name Main features of the LPAZ

Berg. 13 25–30 Poaceae–Picea–Fagus Abrupt increase in Poaceae, increase in Chenopodiaceae,

Cichorioideae, Rumex, Plantago lanceolata and Cerealia

Berg. 12 30–70 Poaceae–Fagus–Picea–Abies–Carpinus Strong increase in Abies at the beginning of the phase. Increase

in Poaceae, Plantago, Humulus–Cannabis type and Rumex.

Regular decrease in Carpinus. Decrease then stabilization in

AP/T. First occurrence of Cerealia at the end of the phase

Upper limit: fall in Fagus, Picea,Abies and Carpinus. Strong

increase in Poaceae

Berg. 11 70–90 Poaceae–Fagus–Picea–Carpinus–Abies Increase in Poaceae, Rumex and Artemisia. Slight decrease in

Fagus and AP/T

Upper limit: increase in Abies, decrease in Fagus and Picea

Berg. 10 90–100 Fagus–Poaceae–Picea–Carpinus Small increase in Carpinus, decrease in Picea and Fagus.

Increase in Artemisia

Upper limit: increase in Poaceae, decrease in Carpinus

Berg. 9 95–110 Fagus–Picea–Poaceae–Carpinus–Abies Increase in Poaceae then Artemisia at the end of the phase.

Small increase in Abies, Betula and Chenopodiaceae, regular

occurrence of Humulus–Cannabis type and Plantago lanceolata.

Decrease in AP/T

Upper limit: decrease in Picea, Fagus and Poaceae

Berg. 8 110–120 Fagus–Picea–Carpinus–Poaceae–Abies Fall in Carpinus, increase in Poaceae, Rumex and Urticaceae.

Beginning of the regular presence of Plantago lanceolata. Small

increase in Pinus

Upper limit: increase in Poaceae and Humulus–Cannabis type,

decrease in Picea

Berg. 7 120–145 Carpinus–Picea–Fagus–Corylus–Abies Regular increase in Fagus, small decline in Picea, decrease in

Ulmus. Small but regular increase in Abies and Poaceae.

Beginning of the regular occurrence of Chenopodiaceae and

Cichorioideae. First occurrence of Plantago lanceolata

Upper limit: fall in Carpinus, increase in Ranunculaceae, Rumex

and Urticaceae

Berg. 6 145–160 Corylus–Carpinus–Picea–Fagus Decrease in Corylus, increase in Fagus, Carpinus and Picea.

Small decrease in Asteroideae, small increase in Poaceae.

Regular presence of Abies at the end of the phase

Upper limit: increase in Abies, Fagus and Alnus, decrease in Corylus

Berg. 5 160–170 Corylus–Picea–Carpinus–Ulmus Increase in Carpinus and Salix, decrease in Corylus except at the

end of the phase. Small decrease in Poaceae and Asteroideae.

Increase in Pinus at the end of the phase Presence of Abies

Upper limit: increase in Fagus, Carpinus and Picea, decrease in

Corylus and Pinus

Berg. 4 170–184 Corylus–Picea–Asteroideae–Ulmus Increase in Picea, small decrease in Asteroideae, Cichorioideae,

Poaceae, Rumex, Humulus–Cannabis type. Decline in Corylus

with abrupt changes. Increase in Poaceae, Fraxinus and Betula

at the end of the phase. Presence of Abies

Upper limit: increase in Carpinus and Salix, decrease in Poaceae

Berg. 3 184–195 Corylus–Asteroideae–Picea–Ulmus Decrease in Corylus at the beginning of the phase, increase in

Ulmus then Pinus at the end of the phase. Small increase in

Cannabis–Humulus type. Picea stable

Upper limit: increase in Picea, abrupt fall in Corylus, beginning

of the regular presence of Betula

Berg. 2 195–205 Corylus–Asteroideae–Picea Increase in Picea, decline in Corylus, small increase in

Cichorioideae and Rumex then Artemisia. First presence of Abies

Upper limit: increase in Ulmus, decrease in Artemisia

Berg. 1 205–220 Corylus–Asteroideae–Poaceae–Picea–Ulmus High values of Corylus and Asteroideae. Values of Ulmus and

Picea around 5%. Presence of Fagus and Carpinus

Upper limit: increase in Picea, Quercus and Cichorioideae

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Table 7

Time-space correlation of local and regional pollen assemblage zonesa

aA=Abies, Al=Alnus, Ast=Asteroideae, Be=Betula, Ca=Carpinus, Ce=Cerealia, Co=Corylus, E=Ericaceae, Fab=Fabaceae, F=Fagus,

Fr=Fraxinus, P=Pinus, Pi=Picea, Po=Poaceae, Q=Quercus, Ro=Rosaceae, U=Ulmus.

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Chenopodiaceae and Asteroideae became more fre-quent. This early impact is not surprising whenconsidering the situation of the site along a valleyallowing easy access to the flat areas of mid elevation.From the available archaeochronological data, this firstpresence might be attributed to the first Neolithicoccupations in Transylvania (Gura Baciului, OcnaSibiului, Starcevo–Cris III–IV Cultures) dated between7850 and 7350 cal BP (Lazarovici, 1993; Demoulle, 1998;Mantu, 1998). Settlements of these people were presento100 km from the studied area, the anthropogenicindicators suggesting that, at that time, groups hadalready begun to travel over the mountain. At Bergerie,during the same period, anthropogenic indicators suchas increases in Cichorioideae and Rumex are also foundnear 7600 cal BP. The local impact seems limited. Thesedimentation rate which was 0.6mm/yr between ca7800 and 7600 cal BP was only 0.17mm/yr between 7600and 4570 cal BP.The age of this first palynological evidence of

Romanian Neolithic husbandry agrees with the presenceof numerous dated archaeological sites in the Balkansbetween 8900 and 6800 cal BP (Willis, 1994; Willis andBennett, 1994) and with the earliest Neolithic 14C datesfrom the northern Balkans (8350–7800 cal BP) (Edwardset al., 1996).The first Cerealia pollen is noticed at Ic Ponor I

during the Neolithic, around 7100 cal BP (6190790BP).At the same time Artemisia and Poaceae are moreabundant. During the same period, several groups areknown in Transylvania near the studied zone: CheileTurzii-Lumea Noua Complex and Turdas Groups(Mantu, 1998).Around 5200–4500 cal BP at Cimeti"ere and Bergerie,

percentages of Poaceae and ruderal communities (Che-nopodiaceae, Rumex, Urticaceae) increase as well asCarpinus and Fagus. At Bergerie during the period4570–4100 cal BP the sedimentation rate is 0.71mm/yrwhich is higher than during the periods 7600–4570 cal BP (0.17mm/yr) and 4100–1935 cal BP(0.14mm/yr). The interpretation of these changes isdifficult. The changes might be due to the climaticvariations which concerned this period in Europe anddetermined the extension of beech (Huntley andPrentice, 1988; Huntley et al., 1989; Huntley, 1990a, b;Kelly and Huntley, 1991; Gardner and Willis, 1999). Butthe consequences of human impact should not beunderestimated. In the Pyrenees, at mid and lowaltitudes (Jalut et al., 1982, 1998; Jalut, 1984; Kenlaand Jalut, 1979) as well as in the plains of centralEurope (K .uster, 1997) and in the Romanian mountains,the abundance of Fagus in the forests may be partlyrelated to the successive cuttings of fir, oak or spruceforests. At Bergerie, when deforestation increases from4100 to 1935 cal BP, the sedimentation rate decreases(0.143mm/yr) then increases between ca 1935 and

680 cal BP (0.24mm/yr) which renders the interpretationdifficult.At Padis, despite very favourable topographic condi-

tions, there is no clear palynological evidence for anearly strong local human impact and the sedimentolo-gical study of the core is not informative. Thesedimentation rate stays low all along the core (between0.09 and 0.14mm/yr from level 90 to level 35). Only therecent Sphagnum peat shows a higher rate (0.78mm/yr).At Cimeti"ere, the use of the sedimentological data

might be more informative but the lack of dates does notallow calculation of the sedimentation rate. However,around 1935 cal BP a relationship exists between theabundance in anthropogenic indicators, the decrease inAP/T values and the presence of thin layers of sand inthe peat. They might be the consequence of erosionprocesses related to deforestations. The contempora-neous decline in the pollen concentration reinforcesthe hypothesis of a strong human impact on thelandscape.In the three sites, between ca 4500 and 3200–

2750 cal BP the forest cover remained stable. Then itbegan to regress. The decrease in AP values is correlatedto a rise in Poaceae, Chenopodiaceae and Plantagospecies.At Padis and Bergerie, around 2750–2550 cal BP, the

development of Poaceae and the increase in ruderalcommunities (Artemisia, Chenopodiaceae, Rumex, Ur-ticaceae, Plantago) demonstrate increased human activ-ities. They rose around 1935BP and 695–660 cal BP(presence of Cerealia at Cimeti"ere). These periodscorrespond to decreases in AP values (Bergerie, Cimeti-"ere, Padis). Humans gradually spread into the mountain(Obelic et al., 1998).At the same time, human impact affected both

elevations between 1000 and 1400m and the lowerzones. Thus, the decrease in percentages and pollenconcentration of Carpinus, Quercus, Ulmus and Tilia, issynchronous with an increase in anthropogenic indica-tors and possibly reflects the destruction of theQuerceto–Carpinetum.The massive forest clearance during the last century is

shown by the fall in AP values and the greater Poaceaeabundance in the upper levels of most peat bogs. In thestudied area the present scarcity of Abies and thenoticeable extension of Fagus are probably the con-sequences of this deforestation. In some places such as IcPonor, the pollen analysis of the fifteen upper centi-meters of Sphagnum peat shows the correlation betweenthe forest destruction and the extension of the cultivatedareas. Cerealia pollen is regularly observed and, in thesurface samples, pollen of Secale, Fagopyrum, Centaureacyanus, Plantago lanceolata, Plantago coronopus andFabaceae are well represented. The abundance ofOnobrychis pollen type indicates cultivated zones, fallowland and pathways in the close vicinity (Figs. 3 and 4).

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11. Comparisons with published local data

In a work focussed on the same area of the Apusenimountains, Ciobanu (1965) studied five sites includingPietrele Onachii (2 cores) and La Ic (called here IcPonor). At the bottom of Pietrele Onachii I, the fall ofPinus and increase in Picea and Corylus values arerecorded. At Pietrele Onachii II, Abies and Carpinus arepresent with low values during the Picea–Corylus phase,before the Fagus and Carpinus extension. Then Carpi-nus, Fagus and Abies seem to extend at the same timewhile Picea decreases. The apparent synchronism in theextension of Carpinus, Fagus and Abies is due toexcessively long sampling intervals.In the Ic Ponor sequence (La Ic of Ciobanu, 1965)

Fagus, Abies and Carpinus appear sporadically duringthe phase with a maximum of Picea. This confirms ourobservations at Ic Ponor, where sporadic occurrences ofFagus, Abies and Carpinus occur during the earlyHolocene.In other sites of the Apuseni mountains (Fig. 2), La

Mlastina and La Mol (Ciobanu, 1967), Calatele(Ciobanu, 1968), Dealul Negru (2 cores) (Lupsa,1972), Dimbul Negru (4 cores; Lupsa, 1973), Baita(Ratiu and Boscaiu, 1971) and Mluha (2 cores)(Ciobanu, 1958) (Fig. 2), a first extension of Carpinusfollowed by the development of beech and fir isobserved. This corresponds to the chronology describedat Padis, Cimeti"ere and Bergerie. The same succession isalso recorded at Dimbul Negru-La Pod and at DealulNegru I. In the latter site, at level 270, the firstpercentage of Fagus (about 8%) indicates that thebeginning of the curve is situated lower, between levels280 and 270, which agrees with our data. In the otherdiagrams, the beginning of curves of Abies and Fagusare indistinct because of the size of the samplinginterval.At Ciurtuci (Fig. 2) (Lupsa, 1971, 1974) several cores

were extracted in different peat bogs. In the earliest work(Lupsa, 1971), the Abies curve is lacking in the pollendiagrams but the extension of Fagus and the decrease ofPicea are synchronous. From our results, it can beassumed that this event probably occurred around4500 calBP. In the second study, (Ciurtuci I, Lupsa,1974) the decline of Corylus and the first increases inCarpinus and Fagus values are observed. In the pollendiagram, these events seem to be simultaneous, bothoccurring between 6450 and 4500 cal BP. Then, a secondand greater extension of Carpinus and Fagus occurred,correlated with the decline of Picea and the developmentof Abies which reached 17%. At Bergerie and Padis theseevents are dated 4030BP.In conclusion, most of the sites studied in the Apuseni

mountains, including Padis, Cimeti"ere and Bergerie,show a comparable history of Picea, Carpinus, Fagusand Abies.

The present and past studies of Ic Ponor confirm thepresence of Carpinus, Fagus and Abies during the earlyHolocene, between 10,190 and 6820BP.Beyond the Apuseni mountains, in the Eastern

Carpathians (Iezerul Calimani, alt. 1650m, CalimaniMts.) and Central Carpathians (Fig. 1) (Taul Zanogutii,alt. 1840m, Retezat Mts.) (Pop and Lupsa, 1971;Mitroescu-Farcas, 1995; Farcas, 1996; Farcas andTantau, 1998; Farcas et al., 1999) the Lateglacial periodis well represented with a strong development ofArtemisia and Chenopodiaceae followed by successiveextensions of Pinus and Picea. The fall in Artemisiavalues and the beginning of the increase in Picea valuesare dated ca 13,140 cal BP (11,140775BP) and latter,Younger Dryas seems to be represented (Farcas et al.,1999).For the period 10,190 cal BP–Present, noticeable

differences are observed with our data from the Apusenimountains. At Iezerul Calimani Pinus remains abundantup to about 5110 cal BP. At Taul Zanogutii (1840m) itstops being well represented around 9660 cal BP due tothe competition between Corylus and Picea. A similar,but un-dated evolution, is recorded in Banat mountains,to the west of Taul Zanogutii (R .osch and Fischer, 2000)(Fig. 1). During the Lateglacial and the beginning of theHolocene, Betula is poorly represented and, as pre-viously discussed, its abundance at Ic Ponor is theconsequence of forest fires favouring heliophilousspecies. During the early Holocene, frequent naturalfires were also noticed by R .osch and Fischer (2000) inthe Banat mountains.During the early Holocene, at Iezerul Calimani and

Taul Zanogutii (Farcas and Tantau, 1998), low-altitudeoak forest is represented with Ulmus percentagesfrequently higher than that of Quercus. Such highUlmus percentages are also noticed in the Apusenimountains (Ic Ponor) and the Banat mountains. InEastern and Central Carpathians as at Apuseni moun-tains, Corylus and Picea are the two dominant trees inthe mountains. After which, Picea declines, whileCarpinus, Fagus and Abies expand. On the contrary,In the Banat mountains, only Corylus is well representedduring the early Holocene and the values of Picea staylow (ca 5%).During the early Holocene at Taul Zanogutii (Farcas

and Tantau, 1998) as well as in the Banat mountains(R .osch and Fischer, 2000), Carpinus is sporadicallypresent and its curve becomes continuous before itsextension phase. In a more recent study of TaulZanogutii (Farcas et al., 1999), occurrences of isolatedhornbeam pollen are very rare during the earlyHolocene and the development of Carpinus is dated7560 cal BP (6645765BP). This date can be comparedwith the dating of the beginning of the regular presenceof Carpinus at Ic Ponor I (7670 cal BP: 6870790BP)and at Ic Ponor II (7800 cal BP: 6980790BP). However,

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at Iezerul Calimani, the first regular occurrences ofCarpinus are noticed only between 5110 and 4500BPand in the Banat mountains, hornbeam spreads between7600 cal BP (6780760BP) and 4350 cal BP (3880760BP). In these sites, the history of Carpinus is similarto that described at Bergerie, Cimeti"ere and Padis. Toexplain the different timing of Carpinus colonisation,Diaconeasa and Farcas (1998) suggested the existence oftwo simultaneous patterns of Carpathian colonisationfrom northern Yugoslavia, one via the west, the otherfrom the south and the east. This hypothesis isquestionable because of the early presence of Carpinusin the Apuseni mountains, the Central Carpathians andthe Banat Mountains which suggests the existence ofglacial and Lateglacial refuges in these areas and,consequently, a process of Holocene recolonisationfrom numerous sheltered zones in the Carpathiansrather than a single area. On the basis of this latterhypothesis, the late development of Carpinus in the NECarpathians might be explained not by the absence ofrefuges but by climate conditions. It was only during the5110–4500 cal BP period that Calimani Mountains, closeto the Ukraine forest-steppe and the present south-eastern lowland Carpinus limit (Ozenda, 1994), experi-enced climatic conditions favourable for the extension ofhornbeam from isolated stands.Differences exist between our dates and the previously

published dates for the extension of Fagus and Abies.Because of the presence of a hiatus, the extensions ofFagus and Abies are not dated at Taul Zanogutii. AtIezerul Calimani, the first regular occurrences of Fagusare observed during the period 5110–4000 cal BP.Despite the inversion of two comparable radiocarbondates, it can be assumed that this phase of developmentbegins ca 4500 cal BP. It is contemporaneous with thatobserved at Padis, Cimeti"ere and Bergerie as well as inmost of the other studied sites. Regular occurrences ofAbies are also noticed from about 4500 cal BP at IezerulCalimani, but a real increase is only noticed around2875 cal BP, later than in our western sites. The recentdata from the Banat mountains show a synchronousextension of Fagus and Abies dated around 4350 cal BP(3880760BP). Here, the distinct interval between thetwo events, visible in the Apuseni mountains and atIezerul Calimani, is not observed.Prior to the studies of Farcas et al. (1999), palyno-

logical studies were devoted the Eastern Carpathians.They had concerned the sites of Poiana Stiolului, DupaLunca-Voslobeni, Mohos (Pop, 1962; Pop and Diaco-neasa, 1967; Ratiu, 1969; Farcas and Tantau, 1998);Calimani Exploatare II, alt. 1700m; Poiana Boilor, alt.1300m; Rachitis, alt. 1700m (Mitroescu-Farcas, 1995)and Cica Mica 1; alt. 1700m (Farcas, 1995). Theobservation of the related pollen diagrams shows thatthe representation of the trees is dependent on localecological conditions or pollen transport but the most

important chronological data agree with our presentresults.In the eastern Carpathians (Poiana Stiolului, Dupa

Lunca–Voslobeni, Mohos) (Pop, 1962; Pop and Diaco-neasa, 1967; Ratiu, 1969; Farcas and Tantau, 1998), thepollen data show the same succession: Pinus phase,Pinus–Picea phase, Picea–Quercetum mixtum–Corylusphase, Picea–Carpinus phase, Picea–Fagus phase. In thefirst two sites Abies is not represented. At Mohos itappears after Fagus development but it stays rare.Concerning human impact, by comparison with the

available pollen data from the Romanian Carpathians,the data from the Apuseni mountains reveal an earlyhuman impact on the mountain. In the EasternCarpathians, at Iezerul Calimani (Farcas et al., 1999),the regular presence of Plantago is only noticed from ca3430 cal BP and is synchronous with the extension ofFagus. High values of Juglans are dated around1580 cal BP, after a strong decrease in Pinus values andan increase in Poaceae, Asteroideae and Carpinuspollen. The latter might have been favoured by thelocal deforestations.In the Banat mountains, the first presence of humans

is recorded earlier, around 4450 cal BP. It is charac-terised by the presence of grains of Triticum-type andPlantago lanceolata and by an increase in charredparticles (R .osch and Fischer, 2000). Then, during theIron Age around 4650–4250 cal BP, increases in Planta-go, Cereals and charcoal are recorded. Increases incharcoal are noticed from the late medieval to themodern period.

12. Regional comparisons

The forest history of the Apuseni mountains describedabove is similar to that of the surrounding countries(Ukraine, Bulgaria, Hungary, Czechia, Slovakia, Polandand Slovenia) (Fig. 1) There is no difference concerningthe history of Quercus and Ulmus. They were present inall these countries from the beginning of the Holocene.Their abundance at 10,190BP in the Apuseni mountainssuggests that they were also present.

Picea, present in Romania around 11,165–10,870 cal BP (Farcas et al., 1999) is also regularlyobserved from the beginning of the Holocene in Czechiaand Slovakia (Zlatnicka dolina, Tojrohe Pleso, Vracov:Rybnickova and Rybnicek, 1996), Slovenia (Sercelj,1996; Culiberg and Sercelj, 1996), and Poland (Ralska-Jasiewiczowa and Latalowa, 1996). However, from theobservation of the pollen diagrams it seems that itsmaximum values are not synchronous.

Carpinus is regularly observed from the beginning ofthe Holocene in Bulgaria (Kupena, Lake Sucho Ezero 2,Tschokljovo: Bozilova and Smit, 1979; Bozilova et al.,1990, 1989; Tshchalova et al., 1990; Tonkov and

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Bozilova, 1992; Willis, 1994; Bozilova et al., 1996;Tonkov et al., 1998) and Hungary (Batorliget: Williset al., 1995) and sporadically present before 7800 cal BPin Ukraine (Dovjok, Orgeev, Kardashinski: Kremenets-ki, 1991, 1995; Kremenetski et al., 1999). These earlyoccurrences support the hypothesis of the existence ofnumerous regional glacial and Lateglacial refuges. In theApuseni mountains, hornbeam occurrs sporadicallyaround 9450–8875 cal BP (Ic Ponor). Its pollen isregularly present at Bergerie from ca 7800 cal BP andits percentages increase strongly ca 6400 cal BP. Thischronology is coherent with data from the Balkans(Willis, 1994), but in the surrounding countries, theCarpinus extension phase is not always synchronous.

Fagus appears in the Apuseni mountains ca 9450–9200 cal BP (Ic Ponor), as well as in Hungary, Slovenia,Czechia and Slovakia where it is observed around10,435, 8900 and before 7800 cal BP, respectively. AtPadis, Cimeti"ere and Bergerie its extension occurred ca4500 cal BP, as in the western Mediterranean (Jalut,1984; Jalut et al., 1982, 1998; Reille and Lowe, 1993). InSlovenia, Czechia, Slovakia and Poland, the extensionphases were not synchronous. They occurred around8340, 7800, 6820 and 5360 cal BP, respectively.In the Apuseni mountains, Abies is present around

6820 cal BP (Ic Ponor) as well as in Czechia andSlovakia (Rybnickova and Rybnicek, 1996) and Poland(Ralska-Jasiewiczowa and Latalowa, 1996), but earlierin Slovenia (>7800 cal BP) (Sercelj, 1996; Culiberg andSercelj, 1996). The extension phases dated 4030 cal BP atBergerie and Padis, around 4500 cal BP in Czechia andSlovakia (Rybnickova and Rybnicek, 1996), and 5110–4500BP at Iezerul Calimani and Taul Zanogutii (Farcaset al., 1999), as well as the contemporaneous extensionof Fagus, emphasises the importance of the 5110–4030 cal BP period for the installation of the mountainforests. Comparisons with vegetation changes occurringat the same time in the Western Mediterranean (Jalutet al., 1997, 2000) area show that these changes wereessentially controlled by climate. When considering thathuman impact increased around 5100–4500 cal BP, wecan consider, like Willis (1994) that, for Romania andthe surrounding areas, it was a critical period for thedevelopment of the present day landscape.At a geographic scale which includes a large part of

the Balkan region (Willis, 1994) and covers a greatdiversity of ecological situations, correspondences be-tween major vegetation changes in the Balkans and inthe Apuseni mountains can be observed. Between 10,190and 8875 cal BP there was expansion of Pistacia in theBalkans and development of Quercus and Ulmus in theApuseni mountains. Between 7800 and 5730 cal BP,Carpinus betulus and Fagus appeared in the Balkanswith the regular presence then development of Carpinusin the Apuseni mountains. Around 5100 cal BP, impor-tant changes occurred in the landscape in the Balkans,

contemporaneous with: the extension of Fagus, theincrease in Poaceae and ruderal communities in theApuseni mountains. This correlation of natural envir-onmental and vegetation changes suggests that theywere determined by climatic changes and possibly thatsuch changes influenced human activities.

13. Conclusions

Studies in the Apuseni mountains date the classicalHolocene phases described in earlier palynologicalinvestigations. In the study area Pinus and Betula werenever important forest components between10,190 cal BP and the Present. At Ic Ponor, the peakof Betula near 9850 cal BP corresponds to a localdevelopment associated to natural forest fires. Thebeginning of the Picea extension occurred prior to10,190 cal BP and around 11,180 cal BP (Farcas et al.,1999).Between 10,190 and 6450 cal BP, Corylus and Picea

were dominant at mid altitude (the Picea–Corylus–Quercetum mixtum phase. Carpinus occurred at low andmedium elevation ca 6450 cal BP. The Picea–Carpinusphase ended around 4500 cal BP when Fagus spread.The Abies development occurred slightly later ca4100 cal BP. Then the Picea–Carpinus–Fagus–Abiesphase began.Around 2540–1935 cal BP Carpinus, decreased and

around 680–660BP the montane forest was submitted tostrong human impact. Before their extension phases, theregional presence of Carpinus, and Fagus is attested bysporadic presence around 8875 cal BP then by noticeableoccurrences from about 7800 cal BP. Abies is observedlater at 7545–7425 cal BP. These occurrences suggest theexistence, during the last cold phases, of regional refugessituated in the deep valleys of the Apuseni mountains.They might have favoured the survival of some of thepresent tree species during the glaciation.More generally, it can be assumed that during the

Last Glacial and Lateglacial period numerous refugesexisted in the Carpathians. These isolated standsfavoured colonisation during the Holocene. Localclimate conditions were major limiting factors andexplain many of the chronological differences.These new palynological investigations demonstrate

the early role of humans on the forest. The firstevidences for settlements is recorded around 7800 cal BPthen 7570 and 7425 cal BP.The first Cerealia pollen is found ca 6820 cal BP.

During the Bronze Age, between about 5100 and 3200–2750 cal BP, human impact seems stable and limited.Then it increased at all elevations, particularly around1935 and 695–660BP. It is during the Last Century thatthe most extensive forest destructions occurred. At

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medium elevation, beech was favoured at the expense ofPicea during the recent recolonisation phases.

Acknowledgements

This research was supported by the Minist"ere Fran-

-cais de l’Education Nationale, de la Recherche et de laTechnologie ‘‘R!eseau Formation Recherche Pays Eur-ope Centrale et Orientale–R!eseau Franco–Roumain’’(Contract 4778836 A), Coordinator Dr. Ch. Causse, andby the Minist"ere Fran-cais des Affaires Etrang"eres(Grant no. 268230C). We express our gratitude to Dr.E. Silvestru (Emil Racovita Speological Institute, ClujNapoca, Romania), for his determining help during thefield work; to Pr. L. Ghergari, for her support at theBabes-Bolyai University of Cluj Napoca; to Pr. Dr. C.Radulescu, Speological Institute of Bucharest for hissupport and his welcome; Dr. M. Bakalowicz and Dr. A.Mangin for their helpful comments on the field; to Dr.M. Thinon for determination of Picea charcoal, Ms D.Dejean for her help in bibliography.Thanks are due to M. Arnold, head of UMS 2004

Tandetron, L.S.C.E., Gif sur Yvette and to the L.S.C.E.Radiocarbon team, especially M. Paterne, N. Tisnerat,E. Kaltnecker, C. Noury and C. Hatt!e.We thank Dr K. Willis, Dr M. Magny and Prof. J.

Rose for their helpful comments on the manuscript.

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