# The beech-fir forest, the baseline natural forest ecosystem in the montane belt of temperate Europe: questioning an ecological myth

Vanessa Py-Saragaglia, Mélanie Saulnier, Laurent Larrieu, Sylvain Burri, Cécile Brun, Mihaela Danu, Didier Galop, Sarah Parrilla, Florence Mazier, Sylvie Ladet

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## To cite this version:

Vanessa Py-Saragaglia, Mélanie Saulnier, Laurent Larrieu, Sylvain Burri, Cécile Brun, et al. The beech-fir forest, the baseline natural forest ecosystem in the montane belt of temperate Europe: questioning an ecological myth. Journal of Archaeological Science: Reports, 2025, 64 (33 p.), pp.105133. 10.1016/j.jasrep.2025.105133. hal-05049406

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Submitted on 28 Apr 2025

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# The beech-fir forest, the baseline natural forest ecosystem in the montane belt of temperate Europe: questioning an ecological myth

Vanessa Py-Saragaglia¹*, Mélanie Saulnier²*, Laurent Larrieu³,⁴, Sylvain Burri S⁵, Cécile Brun²,⁶,
Mihaela Danu⁷, Didier Galop²†, Sarah Parrilla²,⁵,⁸, Florence Mazier², Sylvie Ladet³

*Vanessa Py-Saragaglia and Mélanie Saulnier are jointly first and corresponding authors.

vanessa.py-saragaglia@utbm.fr

melanie.saulnier@univ-tlse2.fr

1. IRAMAT-LMC UMR 7065 CNRS-Université de Technologie Belfort-Monbéliard (France)

2. GEODE UMR 5602 CNRS-Université de Toulouse (France)

3. DYNAFOR UMR 1201 INRAE-Université de Toulouse (France)

4. CNPF-CRPF Occitanie (France)

5. TRACES UMR 5608-Université de Toulouse (France)

6. Nantes Université (France)

7. Faculty of Biology, Alexandru Ioan Cuza University of Iasi (Romania)

8. LASA-Università degli studi di Genova (Italy)

## Abstract

The present study aims to challenge and refine the existing paradigm of relict primary forests located in temperate Europe and to clarify their long-term dynamics. We focussed on the beech-fir (Fagus sylvatica L.-Abies alba Mill.) forest located in the montane belt of Romanian Carpathian Mountains and French Central Pyrenees. An hierarchical sampling strategy was employed, encompassing a multi-proxies study of two cores extracted from one peat bog (Romania) and one lake (France) in proximity to six best-preserved Old-Growth Forests (OGFs), complemented by an archaeological survey on 40 ha, the study of historical archives (16th-20th c.), the soil charcoal analysis of 16 pits, and the charcoal analysis and radiocarbon dating of former charcoal kiln platforms (n = 41). The results enabled the reconstruction of the Holocene vegetation history of these OGFs from the postglacial forest recovery and to highlight the main anthropogenic phases since c. 4000 BP. At the local scale, in the heart of the OGFs, we detected direct traces of human activities from the Bronze Age, with a sharp increase in the Modern Times, influencing forest composition and dynamics. Our results upset the paradigm of relict primary forests in temperate Europe. The montane 'climax' _beech-fir forest used as baseline for natural forest conservation is the result of a co-construction between natural and anthropogenic processes. The biodiversity observed in contemporary OGFs can be attributed to the characteristics of maturity that
have emerged in the absence of human economic activities for several decades. This underscores the necessity for biodiversity-friendly forest management strategies.

**Key-words:** baseline forest ecosystem; long-term old-growth forest history; ecological history; palaeoecology; archaeology; societies-environment interaction

**Highlights:**

(i) Historical ecology reveals the long-term co-construction of the current European beech-fir old-growth forests

(ii) European temperate OGFs are not “primary”

(iii) Insights for the implementation of a biodiversity-friendly forest management

# 1. Introduction

Global change and the worldwide loss of biodiversity have led to a renewed interest in ‘natural’ forests (Watson et al., 2018), also called ‘primary’ forests (Py-Saragaglia et al., 2020), even in Europe, where they now represent less than 3% of the total forest cover (Sabatini et al., 2021) and do not include the full diversity of forest types (Larrieu et al., 2023). The latest inventories of ‘primary’ forests in Europe show that less than 0.1% of these forests remain in France (4% of the total forest cover of the French Pyrenees), while they represent about 2% of the total forest cover of Romania and are concentrated in the Carpathian ecoregion (Knorn et al., 2013; Sabatini et al., 2018, 2021). In general, the most well-preserved natural forests, which show significantly higher levels of biodiversity than forests currently being exploited, are found in valleys with deep, steep slopes, at higher elevations, far away roads and settlements (Sabatini et al., 2018; Larrieu et al., 2023). They are predominantly concentrated in the mountain belt, which ranges from 600-900 to 1200-1700 m.a.s.l. depending on latitude and mountain orientation (Ozenda, 1994), where the climate provides optimal conditions for the growth of beech (Fagus sylvatica L.) mixed with silver fir (Abies alba Mill.) and spruce (Picea abies L.) (only in Romania) in greater proportions depending on local conditions (European Environment Agency, 2007).

Forest ecologists and [nature conservationists](https://nature conservationists.org/) believe that diversified old-growth beech-fir stands are the baseline natural forest ecosystem and the benchmark for biodiversity conservation and mitigate climate change in the montane belt (Larrieu et al., 2023). Indeed, despite the wealth of palaeoenvironmental and archaeological data showing the early human-induced changes of high-altitude environments in Europe (E.g. Walsh et al., 2007, 2008; Galop and Catto, 2014; Rendu et al., 2016; Schumacher et al., 2016; Gassiot Ballbè et al., 2017), these 'relict' stands are still considered to be 'primary' (i.e. they have grown by spontaneous colonisation without human intervention since the end of the last ice age or a very long unspecified time) or close to a 'natural' state (i.e. naturally regenerated forests of native species without visible indications of human activities). In addition to the dominance
of long-lived and shade-tolerant tree species, this assumption is primarily based on the observation of old-growth features in existing stands (Bauhus et al., 2009), i.e. structural heterogeneity and tree species diversity, many large and presumably old living trees, many types of tree-related microhabitats (TreMs), and large amounts of dead wood, lying down or standing, in various stages of decay, which provide resources for rich biodiversity. The study and monitoring of these features (Knorn et al., 2012) is rarely combined with high spatial resolution research into the historical depth of these ecosystems (Pokorný et al., 2022).

A

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C

<figure><img src="image_4.png" /><figcaption>Fig. 1. A: General map of the study area, with the six studied OGF sites, the Tăul Roșia peat bog in Romania and the Espingo Lake in Pyrenees. In the Romanian Carpathians: (1) Băiuț and (2) Poiana Botizii within the Strâmbu Băiuț forest, and (3) Bistra in the valley of Bistra. In the French Pyrenees: (4) Burat natural forest reserve in Marignac, (5) Bois-Neuf in Saint-Mamet, (6) Barrada in Gèdre-Gavarnie. B: Detailed maps of the three studied OGF sites in Romania. At the top, the general map of the Romanian Carpathians OGF sites with Băiuț and Poiana, within the Strâmbu Băiuț forest, and the forest of Bistra. At the bottom, detailed maps of the three study areas: Băiuț and Poiana Botizii together, and Bistra. C: Detailed maps of the three studied OGF sites in French Pyrenees. At the top, the general map of Pyrenean OGF sites: the fir forest of Barrada in Gèdres-Gavarnie, the forest of Bois-Neuf in Saint-Mamet and the Burat natural forest reserve in Marignac. In B and C, the dark green areas represent forest cover and the light green areas represent agro-pastoral zones (only Pyrenees). The red crosses represent the pits for soil charcoal analysis, the black triangles the charcoal kiln platforms, the green dots the pastoral huts and the mine symbols (hammer and pick crossed) the main mines. OGF sites are outlined in black and the circular plots surveyed are outlined in grey.</figcaption></figure>

B

European mountain forests have for a long time provided many resources to past societies (timber, fuelwood, charcoal, grazing resources, etc.) and have been intensively managed through wood-pasture (Rotherham, 2013), coppice (Buckley and Mills, 2015) and high forest systems (Savill, 2015).
Moreover, the impact of human occupation has often been very ancient, sometimes coinciding with the re-colonisation of the present-day tree dryads (Galop and Jalut, 1994; Ralska-Jasiewiczowa et al., 2004; Tinner and Lotter, 2006). Therefore, we postulate that these so-called ‘primary’ forests, which are considered ‘natural’ because they are no longer being exploited, also potentially have a long history of societies interacting with nature (Kirby and Watkins, 2015; Latałowa et al., 2015). This past interaction needs to be studied in several relict ‘primary’ forests to better understand how they have maintained their natural features and dynamics. Therefore, we applied a historical ecology study combining palaeoecological approaches and archaeology at different scales in 6 best preserved Old Growth Forest sites (called after OGF sites) distributed both in the northern Pyrenees (France), which has the highest density of OGFs in Western Europe (Savoie et al., 2015; Sabatini et al., 2021), and in the north-western Carpathian Mountains of Romania (Fig. 1), where the United Nations Educational, Scientific and Cultural Organisation (UNESCO) have classified some forests as “primeval beech forests”.

<figure><img src="image_5.png" /></figure>

<figure><img src="image_6.png" /><figcaption>Fig. 2. Simplified pollen diagram, charcoal accumulation rate (CHAR, only for Espingo lake, expressed in mm⁻².cm⁻².yr⁻¹), and selected pollen and NPP percentage for (A) the Tăul Roșia peat bog and (B) the Espingo lake. (A) For the Tăul Roșia peat bog the exaggerated curves (light colour) are × 5. The pollen diagrams are divided into three 'pollen zones', which have been computed using CONISS zonation in Tilia. (B) For the Espingo lake, the exaggerated curves (light colour) are × 10. The pollen diagrams are divided into five 'pollen zones', which have been computed using CONISS zonation in Tilia.</figcaption></figure>

Through a multi-proxy analysis of sedimentary archives, a systematic archaeological survey, and detailed charcoal analysis, we aim to provide a comprehensive spatio-temporal reconstruction of
the long-term dynamics of these ecosystems, and to explore how interactions between human and natural processes have shaped the exceptional forest ecosystems that remain today.

Our goal is therefore, firstly, to challenge and refine the existing paradigm surrounding relict primary forests in Europe and, secondly, to clarify the long-term dynamics of these forests. Specifically, we seek to demonstrate that the 'climax' beech-fir forest is not merely a natural relic, but the result of a long co-construction between human and natural dynamics.

## 2. Materials and methods

### 2.1. Study areas (Fig. 1 A-C)

In the Romanian Carpathians, the three OGF sites are composed of two sites within the Strâmbu-Băiuț forest, Băiuț and Poiana Botizii, and one forest site in Bistra valley. The site of Poiana is located in the core zone of the UNESCO protected forest area of Strâmbu-Băiuț (Maramureș) created in 2017 (Py-Saragaglia et al., 2020; Nap et al., 2022) (Fig. 1 A, 1B, Table 1). In the Pyrenees, the three OGF sites in the French Pyrenees are Barrada, Bois-Neuf and the natural reserve of Burat (Fig. 1A, 1C, Table 1).

<table><caption>**Table 1**
Main features of the OGF sites including the country, the location, the name of OGF sites and number on the map (Fig. 1), the latitudes and longitudes of the centroids of the sites, the altitudinal ranges (m a.s.l.), the total forest surface (ha), the number of 1 ha-Plot surveyed, and the main tree species present in plots.</caption><thead><tr><th>Context type</th><th>Country</th><th>Location (commune)</th><th>Forest site</th><th>Site id</th><th>Lat</th><th>Lang</th><th>Altitude range (m.a.s. L)</th><th>Surface ha</th><th>Number of 1 ha-plots surveyed</th><th>Main tree species (dominant in bold)</th></tr></thead><tbody><tr><td>Peat bog of Taul Rosia Espingo lake OGFs</td><td>Romania</td><td>Băiuț</td><td>—</td><td>—</td><td>47,661025</td><td>24,027391</td><td>1066</td><td>&lt; 1</td><td>—</td><td>Abies alba, Picea abies, Fagus sylvatica, Sphagnum, Polytricum, Poaceae, Carex Heathland, Nardus grasslands, Pinus ulyssis</td></tr><tr><td></td><td>France</td><td>Oò</td><td>—</td><td>—</td><td>42,72815</td><td>0,4961333</td><td>1882</td><td>7,6</td><td>—</td><td>Abies alba, Picea abies, Fagus sylvatica, Acer pseudoplatanus, Populus alba, Carpinus betulus</td></tr><tr><td></td><td>Romania</td><td>Băiuț</td><td>Strâmbu-Băiuț (1-2)</td><td>1</td><td>47,6388465</td><td>24,0223137</td><td>560-1300</td><td>598 (protected area)</td><td>10</td><td>Abies alba, Picea abies, Fagus sylvatica, Abies alba, Acer pseudoplatanus, Populus alba</td></tr><tr><td></td><td>Romania</td><td>Poiana Botizii</td><td>—</td><td>2</td><td>47,6220659</td><td>24,0711179</td><td>—</td><td>—</td><td>10</td><td>Fagus sylvatica, Abies alba, Acer pseudoplatanus, Populus alba</td></tr><tr><td></td><td>Romania</td><td>Bistra</td><td>Bistra (3)</td><td>3</td><td>47,8910988</td><td>24,2876549</td><td>960-1150</td><td>36</td><td>5</td><td>Fagus sylvatica, Abies alba, Picea abies, Acer pseudoplatanus, Sorbus aucuparia, Ulmus glabra, Sambucus nigra, Praxinus excelsior, Betula pendula, Acer platanoides</td></tr><tr><td></td><td>France</td><td>Marignac</td><td>Burat (4)</td><td>4</td><td>42,8745061</td><td>0,66448723</td><td>1000-1700</td><td>286</td><td>5</td><td>Abies alba, Betula pubescens, Fagus sylvatica, Sorbus aucuparia</td></tr><tr><td></td><td>France</td><td>Saint-Mamet</td><td>Bois-Neuf (5)</td><td>5</td><td>42,8745061</td><td>0,66448723</td><td>1000-1800</td><td>133</td><td>5</td><td>Abies alba, Fagus sylvatica, Salix, Acer platanoides, Betula pubescens, Prunus padus, Sorbus aucuparia</td></tr><tr><td></td><td>France</td><td>Gedre-Gavarnie</td><td>Barrada (6)</td><td>6</td><td>42,8085904</td><td>0,0343462</td><td>1400-1800</td><td>45,9</td><td>5</td><td>Abies alba, Fagus sylvatica, Sorbus aucuparia, Sorbus aria, Quercus petraea, Salix caprea, Betula pendula, Praxinus excelsior</td></tr></tbody></table>

All the OGF sites present several common features (Table 1): (i) they are mainly located in the montane belt, (ii) they belong to the European beech-silver fir, (iii) they have old-growth features, (iv) they are considered as "natural" and well preserved from logging and human activities since a long time, but (v) some of them are also close to or within ancient mining areas (Table 1, Fig. 1 A-C). Indeed, the Maramureș hosts the Baia Mare metallogenic district, one of the most important polymetallic mining districts in the country (Kovacs and Fülöp, 2009). The Strâmbu-Băiuț forest extends over the Herja-Băiuț sector where mining and smelting activities have prospered at least from the 14th c. to 2006
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(Zsamboki, 1989; Borcoş and Udubaşa, 2012). The mining area encompassed multiple zones, including two situated within the UNESCO-classified forest (Poiana Botizii and Cizma) and three located in the Băiuţ valley (Breiner, Johan Hell Mine, and Văratec) (Fig. 1A, 1B). The Bistra site is located close to the Saulyak gold deposit (Dilove village, Ukraine), which contains many other ores including pyrite, and the Solotvyno salt mine (Tiachiv Raion, Zakarpattia Oblast, Ukraine). The Gavarnie area and the Burat forest have zinc and/or silver-lead deposits that were mined to a lesser extent in the 19th-20th centuries. Some forests are also located along rivers used for timber rafting (Garonne in the Pyrenees) and to a lesser extent Bistra in the Carpathians (Fig. 1A, 1C) (Turnock, 1990; Minovez, 1999).

## 2.2. *Hierarchical sampling strategy*

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We applied a spatially hierarchical survey combining palaeoecological and archaeological approaches (Py-Saragaglia et al., 2020).

### Regional and landscape levels

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Multi-proxies analyses, encompassing the study of pollen and Non Pollen Palynomorphs (NPPs), and the calculation of CHarcoal Accumulation Rates (CHAR) were applied in peaty and lacustrine archives to reconstruct the main regional vegetation trajectories (Table 1).

In Romania, the Tăul Roșia peat bog area (1066 m.a.s.l., Văratec Massif, Lăpuș Mountains, Table 1) is 4 km away as the crow flies from the Băiuț forest and 13 to 25 km from the Poiana forest (Fig. 1A). The site, composed of three peat bogs, is described in detail by Mîndrescu et al. (2023). The peat bog area is located in the heart of an OGF area inventoried by WWF Romania (https://padurivirgine.ro/wp-content/uploads/2021/06/Brosura-Baiut-Ultimii-Giganti-ai-Maramuresului.pdf), consisting mainly of *Abies alba*, *Fagus sylvatica* and *Picea abies*. Just above the peat bog zone, this last tree species forms pure subalpine stands (*Homogyno-Piceetum*) with *Vaccinium myrtillus* L., *Vaccinium vitis-idaea* L., *Sphagnum* sp. and *Sorbus aucuparia* L.. The core taken in 2017 from the ombrotrophic and deepest peat bog of Tăul Mare has been used for the study of charcoal particles (Mîndrescu et al., 2023). In 2018, a second 420 cm-long peat core was extracted using a Russian corer with a length of 50 cm and a diameter of 5 cm. This entire core was subsampled at 5 cm intervals for pollen analysis.

In France, the Lake of Espingo (1882 m.a.s.l., Haute-Garonne, Table 1) is situated 12 km from Bois-Neuf, 20 km from Burat, and 60 km from Barrada (Fig 1A). This natural lake covers an area of 7.6 ha and receives water from the Lake of Saussat, 500 m southeast. It is located in the heart of the Oô summer pasture, a 'Natura 2000' site. The vegetation consists mainly of alpine and subalpine meadows and heathlands (*Vaccinium* sp. and *Rhododendron ferrugineum* L.), *Nardus* grasslands and clumps of mountain pines (*Pinus uncinata* Ramond ex DC.). Lake sediment was cored in 2015 using a UWITEC coring device with a diameter of 63 cm. The 100 cm-long lake core was subsampled at 4 cm intervals for pollen and NPP analysis, 1 cm intervals for CHAR analysis.
## OGF sites level

At a finer spatial resolution, we randomly distributed 5 to 10 circular 1ha plots in each OGF sites, in those parts of the forest where the drone could fly with visual contact (Fig. 1B, 1C). LiDAR was used to detect and map any archaeological sites. In addition, a systematic archaeological survey was carried out on every plot from 2017 to 2022. All identified Charcoal Kiln platforms (CKs) were sampled using a soil auger (Py-Saragaglia et al., 2017) or simply by sampling the charcoal horizon when this was located just below the litter. We also consult preserved historical archives (16th to 20th c.) about mining activities and forest use.

We also dug two to four pedoanthracological pits, for a total of 16 pits (4 in Băiuț and Poiana Botizii in two different sectors, and 2 in each other sites). The location of the pits was chosen to avoid local disturbances that could affect the recording. A general description of all pits is presented in the Supplementary files-A.1-Methods. In each pit, soil levels were delineated based on their textural and structural homogeneity (Saulnier et al., 2020). Approximately 5 liters of soil was then extracted from each delineated level (Carcaillet and Thinon, 1996).

## 2.3. Laboratory treatment

### 2.3.1. Multi-proxy study of peat bog and lake

For the pollen core, the samples were processed following the standard techniques (Faegri and Iversen, 1989). They were subjected to treatment with 10% HCl, 10% NaOH, 40% HF, and ZnCl2 acetolysis for 8 minutes. In each sample, a minimum of 400 pollen grains from terrestrial plants was
counted to ensure statistical significance; samples with low pollen concentration were excluded. Pollen and spores were identified using pollen keys (Reille, 1992; Moore et al. 1991) and the nomenclature of pollen types follows Beug (2004). The percentage values were calculated based on terrestrial pollen grains counted in the sample. The pollen diagrams were constructed with Tilia software v. 3.0.1 and CONISS zonation were used to account for patterns in pollen composition along the temporal sequences (Grimm, 2011).

To reconstruct fire regimes at Espingo, contiguous subsamples of 1 cm³ from each 1 cm-thick sediment were analysed for their charcoal content. Subsamples were shaken for 24 hours in an aqueous solution containing 5% sodium hexametaphosphate, 5% KOH, and 10% NaClO to facilitate deflocculation and to distinguish black charcoal from bleached organic matter (Bamber, 1982; Schlachter and Horn, 2010). Charcoal particles were sieved with a 160 µm mesh, then counted and measured using a stereoscope coupled to an image analysis system (WinSEEDLE, Regent Instruments Inc.), enabling the calculation of charcoal accumulation rates (hereafter CHAR, i.e., mm².cm⁻² year⁻¹), with sediment accumulation rates derived from the age-depth models.

### 2.3.2. Charcoal analysis

Soil samples were first air-dried and weighed, except for the Pyrenean pits, where the combination with paleoentomological approaches (Moret et al., 2022)

involved some adjustments. Only in this case, the soils were not dried prior to being weighed and sieved. A fraction of each soil was taken and dried in order to calculate the ratio between the weight of wet and dry soil and thus recover the initial weight of dry soil for the calculation of anthracomasses (see below). Dried or wet samples were wet-sieved using a column of sieves with 4-, 2- and 0.8-mm. The mineral elements larger than 4mm were then removed and weighted to obtain the dried soil weight without coarse fraction. Charcoals were extracted from the remaining organic fraction under a binocular microscope. The total amount of charcoals from each sieve were weighted for the calculation of anthracomasses expressed in milligram of charcoals per kilogram of dry sediment (Talon et al., 1998).

<table><caption>Table 2 Radiocarbon dates of plant remains from the Taul Rosia peatbog and from the Espingo lake used to calibrate the age depth model.</caption><thead><tr><th>Material</th><th>Country</th><th>Mountain</th><th>Location (commune)</th><th>Forest_name</th><th>Pot</th><th>Alt (m a. s.l.)</th><th>Sample depth</th><th>Identification</th><th>Code labo</th><th>Uncalibrated dates BP</th><th>Date cal BP/AD (2s, 95.4%)</th><th>Date cal BP (2s, 95.4%)</th></tr></thead><tbody><tr><td rowspan="10">Plant remains (Sphagnum leaves)</td><td rowspan="10">Romania</td><td rowspan="10">Carpathian</td><td rowspan="10">Baint</td><td rowspan="10">Taul Rosia peat bog</td><td rowspan="10">NA</td><td rowspan="10">1066</td><td>26.3</td><td rowspan="10">NA</td><td>GdA-5830</td><td>1.6990 ± 0.0066</td><td>1965-1966 calAD</td><td>-15 - -16 cal BP</td></tr><tr><td>34.4</td><td>GdA-5961</td><td>300 ± 50</td><td>1463-1666 calAD</td><td>488-284 cal BP</td></tr><tr><td>47.6</td><td>GdA-5832</td><td>190 ± 30</td><td>1725-1813 calAD</td><td>226-138 cal BP</td></tr><tr><td>75.3</td><td>GdA-5833</td><td>1325 ± 45</td><td>641-778 calAD</td><td>1310-1173 cal BP</td></tr><tr><td>100.3</td><td>GdA-5962</td><td>1560 ± 30</td><td>426-575 calAD</td><td>1525-1375 cal BP</td></tr><tr><td>150.5</td><td>GdA-5835</td><td>1750 ± 35</td><td>249-408 calAD</td><td>1704-1543 cal BP</td></tr><tr><td>199.2</td><td>GdA-5836</td><td>2100 ± 35</td><td>200-39 calBC</td><td>2149-1998 cal BP</td></tr><tr><td>230</td><td>GdA-5837</td><td>2270 ± 35</td><td>312-326 calBC</td><td>2261-2155 cal BP</td></tr><tr><td>299.7</td><td>GdA-5838</td><td>2415 ± 50</td><td>596-597 calBC</td><td>2854-2246 cal BP</td></tr><tr><td>331</td><td>GdA-5839</td><td>2700 ± 35</td><td>910-903 calBC</td><td>2862-2752 cal BP</td></tr><tr><td rowspan="5">Plant remains (vermiculite remains)</td><td rowspan="5">France</td><td rowspan="5">Pyrenees</td><td rowspan="5">Où</td><td rowspan="5">Espingo Lake</td><td rowspan="5">NA</td><td rowspan="5">1880</td><td>381.1</td><td>GdA-5963</td><td>11.0 ± 3.4</td><td>1700-1607 calBC</td><td>364-3556 cal BP</td></tr><tr><td>9</td><td>Perx</td><td>10.3 ± 0.39calM</td><td>1061-1929 cal AD</td><td>209-12 cal BP</td></tr><tr><td>32.5</td><td>Perx</td><td>395 ± 30</td><td>1440-1628 cal AD</td><td>511-322 cal BP</td></tr><tr><td>52.5</td><td>Perx</td><td>770 ± 30</td><td>1222-1282 cal AD</td><td>729-666 cal BP</td></tr><tr><td>62</td><td>Perx</td><td>1330 ± 30</td><td>649-775 cal AD</td><td>1301-1176 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>72.5</td><td></td><td>Perx</td><td>1720 ± 30</td><td>250-411 cal AD</td><td>1700-1539 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>88</td><td></td><td>Perx</td><td>1740 ± 30</td><td>245-402 cal AD</td><td>1705-1549 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>97.5</td><td></td><td>Perx</td><td>2015 ± 30</td><td>95 cal BC-109 cal AD</td><td>2044-1841 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>70980</td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>
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We randomly selected 50 charcoals, if available, from each sieve fraction of each layer for taxonomic identification. We calculated the total anthracomasses per pit (SAP), i.e. the weight of charcoal per pit divided by the total dried soil weight, the anthracomasses per layer (SAL), i.e. the weight of charcoal per layer, and, finally, the specific anthracomasses in each layer per taxa (SAT), i.e. the weight of charcoal per taxa per layer.

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The sampled materials extracted from CKs were wet sieved using 4 mm and 2 mm mesh sieves and then dried in the shade. For this preliminary charcoal analysis, which consisted of identifying the main charred tree-species, we randomly selected 30 to 50 charcoal fragments from each CK sample selected for radiocarbon dating. We have mainly used 4mm mesh charcoal, except for a few samples that contain less charcoal.

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All taxon identifications of charcoal from both contexts (soil and CKs) were done using a reflected light microscope, with magnifications of 100 to 1000, reference collections and xylological atlases (Schweingruber, 1990).

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### 2.3.3. Radiocarbon dating

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Age-depth models were constructed with eleven 14C dates for the Tăul Roșia core (various plant remains), and six 14C for the Espingo. Plant remains (*Sphagnum* sp., leaves) were radiocarbon dated using Accelerator Mass Spectrometry (AMS) in the Radiocarbon and Mass Spectrometry Laboratory of Gliwice (Poland) and in the Poznań Radiocarbon Laboratory (Poland).

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We selected 58 soil charcoals and 44 samples from CKs for radiocarbon dating using AMS in the Poznań Radiocarbon Laboratory or in the Beta analytic laboratory (Table S3). For soil charcoal, samples were chosen based on (i) their weight, greater than 2mg, (ii) their identification, from one of the four main species (fir, beech, spruce, sycamore), and (iii) extracted from a deeper level. For archaeological samples, we selected mainly branches and twigs from the main tree-species identified in each CK selected. Conventional radiocarbon 14C ages were then calibrated using Oxcal v. 4.4 (Bronk Ramsey, 2009; Bronk Ramsey and Lee, 2013) and the calibration curve IntCal20 (Reimer et al., 2020) and Bomb21NH1 (Hua et al., 2022). We used a standard deviation of 2 σ (95% probability).

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## 3. Results

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### 3.1. Multi-proxy study of peat bog and lake (Fig 2A and B, Table 2)

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The core extracted from the Taul Rosia peat bog consists mainly of relatively homogeneous peat sediment (dark or light peat with or without fibrous part). The pollen diagram obtained for the Romanian site reveals three distinct phases (Fig 2A) spanning the period (3650 cal. BP to present) (Table 2). During the first phase (3650-2400 cal. BP), the landscape was largely dominated by forests, with *Fagus*, *Picea*, and *Carpinus* as the predominant pollen types, supplemented by contributions from lower altitude species like *Quercus* and *Corylus*. The second phase (2400-500 cal. BP) is characterised by a significant increase in *Abies* and *Pinus* pollen, along with marked fluctuations in the major forest taxa
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(*Fagus* and *Picea*). This period also exhibits a substantial rise in herbaceous pollen types, such as Poaceae and *Artemisia*, along with a growing presence of pollen linked to human activities, particularly Cerealia, which becomes increasingly prominent during the early Middle Ages. In the final phase, beginning around 500 cal. BP, there is a gradual increase in *Abies*, *Picea*, and Ericaceae pollen, occurring alongside a concurrent decline in *Fagus* and *Carpinus*.

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The entire core extracted from the Espingo lake consists of relatively homogeneous lacustrine sediment, with the exception of a few sand inclusions (several sand inclusions between 14,3 and 35.5 cm; 74-75 cm and 77-78 cm) and some areas of large plant remains (67-68 cm). The 2000 yrs reconstruction of regional vegetation changes show five main pollen zones (Fig. 2B, Table 2). The period 2000-1350 cal. BP is dominated by trees (*Pinus*, *Corylus*, *Abies*, *Betula*, *Quercus*) with low levels of pastoral indicators. Around 1300 cal. BP, *Pinus* and *Abies* start to decrease while Poaceae, pastoral indicators (pollen and NPPs) and CHAR particles increase. The period 900-550 cal. BP shows a decrease in percentages of *Abies* and increase in *Juniperus*, Poaceae and pastoral indicators (*Rumex acetosa* L., *Plantago lanceolata* L., *Urtica* and coprophilous fungi). Between 550-450 cal. BP, pine pollen as well as coprophilous fungi increased while Poaceae declined, suggesting wood-pasture practices development. After this phase, *Abies* and *Betula* pollen decreased again, and the tree curve reached its lowest level while CHAR particles increased. From 0 cal. BP, this trend is reversing with the increase of all tree taxa and the sharp decrease of anthropogenic indicators.

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We analysed 3370 charcoals (2167 from Carpathian pits and 1203 from Pyrenees pits) (Fig. S1). The SAPs varied greatly from 0,03g/kg dry soil in Poiana S1-pit 2 to more than 3g/kg dry soil in Bois-Neuf 1 (Fig. 3). Except for the latter, the SAPs of the two mountain areas are almost comparable, with values ranging from 0.03 to 0.07 g/kg. The maximum SAP in France, in Bois-neuf 1, is about 8 times higher than the maximum SAP in Romanian, in Băiuţ S1-pit 1. Most of the lower SAP were obtained in Romania, especially in Poiana S1-pit 2 and 1, Poiana S2- pit 2, Bistra 2, Baiut S1-pit 1, just one SAP from a Pyrenean soil is similarly very low in Barrada 2, 0.05 g/kg.

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The SAL also varied greatly between layers of the same pits, with the uppermost layer usually characterised by the higher SALs, but sometimes the SALs increase as we reach deeper horizons, as in the two pits of Barrada in Pyrenees and the pits of Băiuţ S1-pit 1 and Poiana S1-pit2 (Fig. 4).

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We identified 3338 charcoals at various taxonomic levels (family, genus, species, etc), while 332 charcoals remained indeterminable, meaning that the charcoal does not contain any anatomical features that would allow its identification (Table 3). Indeterminate taxa whose anatomy is observable, but which are either from immature wood (one- or two-year-old twigs) or from wood in poor preservation are grouped within the sub- phylum as either "Angiospermae" or "Gymnospermae" based on the few criteria identified. Unidentifiable charcoal is grouped under the term "indeterminable".

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(charcoal in very poor condition with no visible anatomical features). Among the others, we classified charcoal with bark but without enough wood anatomical features as "bark", and we grouped together some taxa with common features that we could not separate: *Betula/Corylus*, *Abies/Picea*, and *Salix/Populus*.

Based on the ecological niche of the different species, we assume that some genus may be undoubtedly associated with a species (specified in brackets in Table 3).

In the Carpathians, the main taxa identified is beech (Fig. 4, Table 3). In the Pyrenees, soil charcoals were predominantly dominated by fir, with the frequent presence, even with very weak anthracomasses, of pioneering and post-pioneering species, such as *Acer* and Ericaceae, indicating some canopy opening events. Beech charcoal remains were poorly represented, except in the Barrada pit 2.

<figure><img src="image_7.png" /><figcaption>Fig. 3. Soil Anthracomasses per Pits (SAP) in forest soils in France (left) and in Romania (right). The SAPs are expressed as the ratio between the weight of charcoals in grams and the weight of the dry soil in kilograms.</figcaption></figure>

<figure><img src="image_8.png" /><figcaption>Fig. 4. Specific anthracomasses per taxa and layer (SAT in g/kg) in the 16 pits from Pyrenees (A-F) on the left and from the Carpathians on the right (G-P).</figcaption></figure>

We selected 28 charcoal remains from the Pyrenean pits and 30 from the Carpathian pits for radiocarbon dating. The radiocarbon dates indicate the existence of three principal periods of forest fires, though not contemporaneously (Fig. 5, Table 4). In the Carpathians (Fig. 5A), the initial period of detected fires dates back to the Upper Pleistocene. The second period aligns with the transition between
<table><caption>**Table 3**
The number of soil charcoal remains identified by major taxonomic groups in provided below. In parentheses are the species to which the charcoal most certainly belongs, given the local ecological conditions.</caption><thead><tr><th>Taxonomic level</th><th>Taxon/Group</th><th>Nb charcoal in carpathian pits</th><th>Nb charcoal in pyrenean pits</th></tr></thead><tbody><tr><td rowspan="3">Sub-phyllum</td><td>Angiosperma</td><td>190</td><td>181</td></tr><tr><td>Gymnosperma</td><td>198</td><td>133</td></tr><tr><td>Ericaceae</td><td>-</td><td>38</td></tr><tr><td rowspan="2">Family</td><td>Fabaceae</td><td>-</td><td>2</td></tr><tr><td>Rozaceae</td><td>-</td><td>2</td></tr><tr><td rowspan="6">Genus</td><td>Abies (Silver fir)</td><td>179</td><td>665</td></tr><tr><td>Acer (Sycamore/ Norway maple)</td><td>2</td><td>24</td></tr><tr><td>Betula (Silver/White birch)</td><td>5</td><td>4</td></tr><tr><td>Corylus</td><td>3</td><td>3</td></tr><tr><td>Fagus (European beech)</td><td>1241</td><td>12</td></tr><tr><td>Praxinus (Ash)</td><td>1</td><td>1</td></tr><tr><td rowspan="10">Others classification</td><td>Picea (Spruce)</td><td>170</td><td>-</td></tr><tr><td>Pinus</td><td>58</td><td>-</td></tr><tr><td>Prunus (Wild/Bird cherry)</td><td>-</td><td>1</td></tr><tr><td>Quercus</td><td>1</td><td>4</td></tr><tr><td>Rhododendron (Alpenrose)</td><td>-</td><td>15</td></tr><tr><td>Silix</td><td>14</td><td>1</td></tr><tr><td>Sorbus (Mountain ash/ Whitebeam)</td><td>-</td><td>1</td></tr><tr><td>Ulmus</td><td>1</td><td>-</td></tr><tr><td>Indeterminable</td><td>225</td><td>107</td></tr><tr><td></td><td>Bark</td><td>37</td><td>7</td></tr><tr><td></td><td>Abies/Picea</td><td>147</td><td>-</td></tr><tr><td></td><td>Betula/Corylus</td><td>-</td><td>2</td></tr><tr><td></td><td>Picea/Pinus</td><td>3</td><td>-</td></tr><tr><td></td><td>Populus/Silix</td><td>8</td><td>-</td></tr><tr><td>Total</td><td></td><td>2167</td><td>1203</td></tr></tbody></table>

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the end of the Neolithic era and the advent of the Metal Age. The final fire phase, for which we also have the most extensive chronological data, spans from the Bronze Age to the present. In the Pyrenees (Fig. 5B), the initial period aligns with the early Holocene. The subsequent period of fire activity is situated in the middle of the Holocene. Finally, the most extensive period, which encompasses the majority of the obtained dates, commences with the Bronze Age and the advent of human activities in the region.

### 3.3. Archaeology and CK charcoal analysis

The number of points extracted by LiDAR was limited by the conifer dominated forest and did not allow the detection of archaeological sites. However, the pedestrian

survey revealed numerous former CKs at every OGF site (Fig. 6 A, B). The area occupied by the CK (platform) rarely exceeds 5x5m and is generally 3x2.5m (2x2 on steep slopes) (Table S1). In both the Pyrenees and the Carpathians, the rich charcoal horizon is rarely more than 10 cm thick. In most cases it is in the A horizon, just below the litter.

A total of 93 CKs were inventoried and mapped, including 73 in Romania (25 ha surveyed) and 20 in France (15 ha surveyed) (Fig. 1B, 1C, Table S1). The highest concentration of CK remains was found at the Băiuț site with an average of 4,5 CKs/ha. The plot, which is located near the mine entrances in the Johan Hell mine area (Băiuț), has a maximum of 16 CKs/ha (Fig. 1B). A King Robert diploma on the rights of local communities in the forest of Baia Sprie/Baia Mare confirms charcoal burning and logging activities in the *silva nigra* in the 14th c. (May 29th, 1329). The charcoal supply to the Băiuț industry is documented by accounting registers in the early 20th c. (1912-1929), but without specifying its origin.

In the Pyrenees only, we also found the remains of 4 huts made of dry-stone wall and perishable cover (2 Barrada, 2 Bois-Neuf, Fig. 1C), probably related to pastoral activity. In fact, in this region, as in central Europe (Hutt, 2023), the huts of lumberjacks and charcoal burners were usually made of perishable materials only (Py-Saragaglia, 2024). The pastoral activity is also documented by texts from the 19th c. (access of the herds to the forest). Texts document also the installation of hydraulic saws and the sale of wood cuttings in the modern and contemporary times.

We dated 23 CKs found in Romania and 18 in France (Fig 6, Table 4). The majority of CKs worked during the Modern-Contemporary times (Table 4). The oldest CK has been found in the Bistra site and dated from 2284-2037 cal. BC, i.e the Bronze Age. It is noteworthy that specimens of CK this
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advanced in age are exceedingly rare in Europe (Krebs et al., 2017, Table 4). In the Pyrenees, the oldest CK dates back to the Carolingian period (689-890 cal. AD). In the Strambu-Băiuţ area, the oldest dated CK is Medieval (1280-1395 cal. AD) and coincides with the beginning of the Medieval mining activity in the Băiuţ sector. In the core zone of the UNESCO forest, the oldest CK found dates from the late Medieval-early Modern period (1436-1624 cal. AD).

1360 charcoal fragments from 41 CKs were identified. A total of 13 taxa were recognised, ranging from the sub-branch level (e.g. conifer/Gymnospermae), for the poorly preserved pieces, to the species level (e.g. *Corylus avellana*) (Fig. 7). We identified shade tolerant trees (*Abies alba*, *Fagus sylvatica*, *Picea abies*, *Taxus baccata*), shrubs and bushes (*Corylus avellana* L., cf. *Hedera helix* L. and *Lonicera* spp.); shade to heliophilous bush (*Vaccinium* spp.) and pioneer and post-pioneer trees and shrub (*Betula* spp., *Rhododendron ferrugineum* and *Acer* spp.).

The main tree taxa used for charcoal manufacturing were beech and fir, in very different proportions depending on the site (Fig. 7; Table S1). Some mountain Ericaceae, found only in the Pyrenees, were probably used with other plants (leaves) to cover the wood pile before carbonisation. The yew, identified only in three CKs of the Bistra site and dated to the Bronze Age and the Late Medieval periods, is currently absent from the vegetation cover. All the other taxa still exist in today's vegetation. A complementary ongoing historical investigation and study using dendroanthracological tools aim to characterise charcoal production and forest management practices.

# 4. Discussion

## 4.1 Divergent holocene trajectories led to comparable fir-beech OGFs (Fig. 8 A and B)

In Romania, all OGF sites showed the same Late Glacial and Early Holocene history with an early colonisation by pine species between 14000-11000 cal. BP, followed by the first evidence of beech in soil charcoal around 3800-3600 cal. BP, and then the first evidence of fir around 3,000 cal. BP (Fig. 9A). This charcoal-based Holocene reconstruction of tree expansion is consistent with the pollen-based reconstruction from Tăul-Roșia and, more globally, with the other regional pattern of Holocene vegetation expansion and fire dynamics in this part of the Carpathians (Fig. 8A) (Haliuc et al., 2016; Tanțău et al., 2011; Magyari et al., 2018; Grindean et al., 2019; Lestienne et al., 2023). In this area, the expansion of beech was dated to the late Holocene, i.e. during the sub-Atlantic period (Fărcaș et al., 2013; Tanțău et al., 2003) even if its presence is already confirmed from the Atlantic-Subboreal Period (Tanțău et al., 2011). The first detections of beech in Poiana Botizii, Bistra and Băiuţ are consistent with all of these previous studies with three beech charcoals from soils radiocarbon dated to 3821-3574 cal. BP, 3480-3373 cal. BP, and 3070-2863 cal. BP, respectively. These three dates, clustered as the second fire period in the Carpathians, also coincide with episodic increases in fire activity in this region that
have been documented for the periods 4000 and 3700 cal. BP, 3400 and 3200 cal. BP, and around 2700 cal. BP (Haliuc et al., 2016).

<figure><img src="image_9.png" /></figure>

## Soil charcoal radiocarbon datings

<figure><img src="image_10.png" /><figcaption>Fig. 5. Radiocarbon dates of charcoal pieces sampled in soil pits from Carpathians (A) and Pyrenees (B).</figcaption></figure>

The arrival and subsequent expansion of fir in the Carpathians is frequently cited as occurring at a much later point in time than in other European massifs (Tanţău et al., 2011; Magyari et al., 2012; Cheddadi et al., 2014; Grindean et al., 2019). Our regional reconstruction, in conjunction with the majority of dates obtained from fir charcoal extracted from soils, corroborates this late arrival. Nonetheless, in Bistra, we also dated one fir charcoal for the period of beech expansion. Surprisingly,
one charcoal of fir was dated between 13587-13327 cal. BP, which confirms the presence of the species before the late glacial period during the Younger Dryas cooling. As already highlighted by palynological and genetic studies, the Carpathian Arc probably provided microrefugia for many woody species during the last glacial maximum (Varga, 2009; Magyari et al., 2012, 2018). It is highly likely that fir populations maintained in such refugia but that this former population does not contribute to the modern gene pool exhibits, as already observed for beech (Magri et al., 2008; Gömöry et al., 2012). A new genetic study of the Maramureş fir population could provide confirmation that fir populations still present at the last interglacial did or did not contribute to the gene pool of current populations. Indeed, recent syntheses on the Holocene expansion of the species (Liepelt et al., 2009; Cheddadi et al., 2014) do not integrate new evidence of fir presence attested by different macro-remains recently discovered in the southwestern part of the Romania (Magyari et al., 2018).

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<table><caption>Table 4
Radiocarbon dates of charcoal remains from soil and former charcoal kiln platforms.</caption><thead><tr><th>Material</th><th>Country</th><th>Mountain</th><th>Location (commune)</th><th>Forest_name</th><th>Plot</th><th>Alt (m a. s.l.)</th><th>Pit /CK_number</th><th>Identification</th><th>Code labo</th><th>Uncalibrated dates BP</th><th>Date cal BP/AD (2σ, 95,4%)</th><th>Date cal BP (2σ, 95,4%)</th></tr></thead><tbody><tr><td rowspan="10">Soll charcoal</td><td rowspan="10">Romania</td><td rowspan="10">Carpathian</td><td rowspan="10">Polana</td><td rowspan="3">Strâmbu-Bălut</td><td>5</td><td>976</td><td>Pit 1,51</td><td>Fagus</td><td>Poz.131940</td><td>210 ± 30</td><td>1642-1923 cal AD</td><td>255-34 cal BP</td></tr><tr><td>2</td><td>850</td><td>Pit 2,51</td><td>Pinus</td><td>Poz.132335</td><td>10000 ± 50</td><td>9777-9320 cal BC</td><td>11726-11269 cal BP</td></tr><tr><td>1</td><td>950</td><td>Pit 1,52</td><td>Pinus</td><td>Poz.131903</td><td>11850 ± 60</td><td>12030-11564 cal BC</td><td>13973-13514 cal BP</td></tr><tr><td rowspan="7">Bălut</td><td rowspan="7">Strâmbu-Bălut</td><td>2</td><td>1100</td><td>Pit 2,52</td><td>Fagus</td><td>Poz.132336</td><td>3425 ± 30</td><td>1873-1625 cal BC</td><td>3621-3574 cal BP</td></tr><tr><td>6</td><td>1100</td><td>Pit 1,52</td><td>Fagus</td><td>Poz.132529</td><td>320 ± 30</td><td>1484-1644 cal AD</td><td>467-307 cal BP</td></tr><tr><td>4</td><td>890</td><td>Pit 2,52</td><td>Fagus</td><td>Poz.132337</td><td>11600 ± 60</td><td>11639-11379 cal BC</td><td>13567-13327 cal BP</td></tr><tr><td>1</td><td>846</td><td>Pit 1,51</td><td>Fagus</td><td>Poz.132511</td><td>195 ± 30</td><td>1646-1916 cal AD</td><td>303-0 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.132339</td><td>11220 ± 70</td><td>11351-11042 cal BC</td><td>13298-12974 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.132340</td><td>2850 ± 35</td><td>1122-915 cal BC</td><td>3070-2863 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.160044</td><td>145 ± 30</td><td>1669-1940 cal AD</td><td>281-4 cal BP</td></tr><tr><td rowspan="15">France</td><td rowspan="15">Pyrenees</td><td rowspan="15">Marignac</td><td rowspan="15">Bistra</td><td rowspan="3">Bistra</td><td>3,2</td><td>1120</td><td>Pit 1</td><td>Fagus</td><td>Poz.131905</td><td>75 ± 30</td><td>441-154 cal BP</td><td>258-31 cal BP</td></tr><tr><td>7,1</td><td>1145</td><td>Pit 2</td><td>Fagus</td><td>Poz.131911</td><td>100 ± 30</td><td>1683-1930 cal AD</td><td>266-14 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.131904</td><td>745 ± 30</td><td>1225-1295 cal AD</td><td>724-655 cal BP</td></tr><tr><td rowspan="7">Saint-Mamet</td><td rowspan="7">Bois-neuf</td><td rowspan="7">Burat</td><td>91</td><td>1394</td><td>Pit 1</td><td>Fagus</td><td>Poz.131907</td><td>805 ± 30</td><td>1179-1278 cal AD</td><td>771-673 cal BP</td></tr><tr><td></td><td></td><td></td><td>Picea</td><td>Poz.131908</td><td>810 ± 30</td><td>1178-1276 cal AD</td><td>772-675 cal BP</td></tr><tr><td></td><td></td><td></td><td>Fagus</td><td>Poz.159553</td><td>335 ± 30</td><td>1475-1640 cal AD</td><td>473-312 cal BP</td></tr><tr><td>94</td><td>1177</td><td>Pit 2</td><td>Fagus</td><td>Poz.159554</td><td>330 ± 30</td><td>1490-1640 cal AD</td><td>471-310 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.159555</td><td>710 ± 30</td><td>1262-1387 cal AD</td><td>687-564 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.159563</td><td>700 ± 30</td><td>1267-1388 cal AD</td><td>682-563 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.160043</td><td>650 ± 30</td><td>1281-1395 cal AD</td><td>669-555 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.131906</td><td>560 ± 30</td><td>2740-2493 cal BP</td><td>640-524 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.131910</td><td>620 ± 30</td><td>1296-1400 cal AD</td><td>662-551 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.132341</td><td>465 ± 30</td><td>1410-1464 cal AD</td><td>540-482 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.131904</td><td>635 ± 30</td><td>1299-1397 cal AD</td><td>661-554 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.172732</td><td>10060 ± 60</td><td>9874-9372 cal BC</td><td>11823-11321 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.172771</td><td>2945 ± 30</td><td>1260-1048 cal BC</td><td>3209-2997 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.172772</td><td>3215 ± 30</td><td>1531-1424 cal BC</td><td>3480-3373 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.172773</td><td>160 ± 30</td><td>1663-1905 cal AD</td><td>267-45 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.172770</td><td>2905 ± 30</td><td>1209-1009 cal BC</td><td>3158-2958 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.172769</td><td>1840 ± 30</td><td>124-311 cal AD</td><td>1827-1640 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.157310</td><td>1085 ± 30</td><td>992-1021 cal AD</td><td>1069-930 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.147649</td><td>2655 ± 30</td><td>999-787 cal BC</td><td>2648-2736 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.148015</td><td>2850 ± 30</td><td>1114-924 cal BC</td><td>3063-2873 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.147650</td><td>1500 ± 30</td><td>430-556 cal AD</td><td>1530-1394 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.159400</td><td>8590 ± 50</td><td>7732-7537 cal BC</td><td>9601-9406 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.160040</td><td>8930 ± 50</td><td>7128-6701 cal BC</td><td>9691-9650 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.148016</td><td>8580 ± 80</td><td>7935-7480 cal BC</td><td>9684-9429 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.153156</td><td>3310 ± 30</td><td>1669-1504 cal BC</td><td>3618-3453 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.159481</td><td>6250 ± 40</td><td>5312-5066 cal BC</td><td>7261-7015 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.153672</td><td>1990 ± 30</td><td>46 cal BC-117 cal AD</td><td>1995-1834 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.153673</td><td>6095 ± 35</td><td>5206-4053 cal BC</td><td>7157-6802 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.153675</td><td>9390 ± 50</td><td>8001-9491 cal BC</td><td>10750-10440 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.160038</td><td>3160 ± 35</td><td>1506-1315 cal BC</td><td>3454-3264 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.160039</td><td>3270 ± 30</td><td>1616-1456 cal BC</td><td>3656-3405 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.160042</td><td>1380 ± 30</td><td>601-758 cal AD</td><td>1349-1192 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.160041</td><td>1435 ± 30</td><td>580-655 cal AD</td><td>1371-1295 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.148017</td><td>1950 ± 30</td><td>37 cal BC - 201 cal AD</td><td>1986-1750 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.147651</td><td>1355 ± 30</td><td>610-774 cal AD</td><td>1340-1176 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.148018</td><td>1215 ± 30</td><td>609-890 cal AD</td><td>1262-1061 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.159552</td><td>2940 ± 35</td><td>1260-1017 cal BC</td><td>3209-2966 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.157309</td><td>275 ± 30</td><td>1509-1796 cal AD</td><td>441-154 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.156505</td><td>2525 ± 30</td><td>791-544 cal BC</td><td>2740-2493 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz.157254</td><td>2075 ± 30</td><td>172 cal BC - 8 cal AD</td><td>2121-1942 cal BP</td></tr></tbody></table>

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<table><caption>Table 4 (continued)</caption><thead><tr><th>Material</th><th>Country</th><th>Mountain</th><th>Location (commune)</th><th>Forest_name</th><th>Plot</th><th>Alt (m a.s.l.)</th><th>Pit</th><th>Pit /CK_number</th><th>Identification</th><th>Code labo</th><th>Uncalibrated dates BP</th><th>Date cal BC/AD (2σ, 95,4%)</th><th>Date cal BP (2σ, 95,4%)</th></tr></thead><tbody><tr><td rowspan="10">CK charcoal</td><td rowspan="10">Romania</td><td rowspan="10">Carpathian</td><td rowspan="10">Bistra</td><td rowspan="10">Bistra</td><td rowspan="2">2,2</td><td>1624</td><td>Pit 1</td><td></td><td>Abies</td><td>Poz-157307</td><td>2435 ± 30</td><td>751–406 cal BC</td><td>2700–3355 cal BP</td></tr><tr><td>1595</td><td>Pit 2</td><td></td><td>Abies</td><td>Poz-157308</td><td>3150 ± 30</td><td>1500–1314 cal BC</td><td>3449–3263 cal BP</td></tr><tr><td>4</td><td></td><td></td><td>Abies</td><td>Poz-156561</td><td>1630 ± 30</td><td>382–541 cal AD</td><td>1556–1409 cal BP</td></tr><tr><td></td><td></td><td></td><td>Fagus</td><td>Poz-159651</td><td>55 ± 30</td><td>1694–1917 cal AD</td><td>257–33 cal BP</td></tr><tr><td></td><td></td><td></td><td>Fagus</td><td>Poz-157261</td><td>111.42 ± 0.36</td><td>Port-1945</td><td>/</td></tr><tr><td></td><td></td><td></td><td>Fagus + Abies</td><td>Poz-174148</td><td>375 ± 30</td><td>1450–1632 cal AD</td><td>501–318 cal BP</td></tr><tr><td></td><td></td><td></td><td>Fagus</td><td>Poz-174076</td><td>250 ± 30</td><td>1522–1940 cal AD</td><td>429–114 cal BP</td></tr><tr><td></td><td></td><td></td><td>Acer</td><td>Poz-174075</td><td>110 ± 30</td><td>1689–1938 cal AD</td><td>270–11 cal BP</td></tr><tr><td></td><td></td><td></td><td>Picea</td><td>Poz-174201</td><td>100 ± 30</td><td>1683–1930 cal AD</td><td>267–21 cal BP</td></tr><tr><td></td><td></td><td></td><td>Abies</td><td>Poz-174074</td><td>175 ± 30</td><td>1658–1910 cal AD</td><td>293–41 cal BP</td></tr><tr><td></td><td></td><td></td><td>Picea</td><td>Poz-174073</td><td>165 ± 30</td><td>1661–1907 cal AD</td><td>289–44 cal BP</td></tr><tr><td></td><td></td><td></td><td>Taxus</td><td>Poz-174072</td><td>3750 ± 30</td><td>2284–2037 cal BC</td><td>4233–3906 cal BP</td></tr><tr><td></td><td></td><td></td><td>Abies</td><td>Poz-174071</td><td>185 ± 30</td><td>1653–1915 cal AD</td><td>297–35 cal BP</td></tr><tr><td></td><td></td><td></td><td>Taxus</td><td>Poz-174160</td><td>345 ± 30</td><td>1470–1637 cal AD</td><td>480–313 cal BP</td></tr><tr><td></td><td></td><td></td><td>Abies alba</td><td>Poz-153373</td><td>155 ± 30</td><td>1665–1905 cal AD</td><td>265–46 cal BP</td></tr><tr><td></td><td></td><td></td><td>Fagus sylvatica + Corylus</td><td>Poz-153127</td><td>75 ± 30</td><td>1691–1920 cal AD</td><td>259–31 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>824</td><td>CK3</td><td></td><td>Fagus sylvatica</td><td>Poz-153374</td><td>655 ± 30</td><td>1280–1395 cal AD</td><td>671–555 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>838</td><td>CK20</td><td></td><td>Abies alba</td><td>Poz-153372</td><td>55 ± 30</td><td>1694–1917 cal AD</td><td>257–33 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Abies alba</td><td>Poz-153380</td><td>105 ± 30</td><td>1683–1936 cal AD</td><td>266–14 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Fagus sylvatica</td><td>Poz-153153</td><td>200 ± 30</td><td>1643–1917 cal AD</td><td>307–34 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Fagus sylvatica</td><td>Poz-153154</td><td>100.25 ± 0.38</td><td>1695–1917 cal AD</td><td>265–22 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Abies alba</td><td>Poz-153154</td><td>80 ± 30</td><td>1691–1921 cal AD</td><td>260–29 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>893</td><td>CK6</td><td></td><td>Abies alba</td><td>Poz-153129</td><td>135 ± 30</td><td>1674–1943 cal AD</td><td>276–6 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>902</td><td>CK7</td><td></td><td>Abies alba</td><td>Poz-153155</td><td>405 ± 30</td><td>1436–1624 cal AD</td><td>515–327 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>916</td><td>CK6</td><td></td><td>Fagus sylvatica + Abies</td><td>Poz-153131</td><td>140.21 ± 0.44</td><td>1682–1936 cal AD</td><td>278–6 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Abies alba</td><td></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>927</td><td>CK12</td><td></td><td>Fagus sylvatica</td><td>Poz-153389</td><td>115 ± 30</td><td>1690–1940 cal AD</td><td>270–10 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>926</td><td>CK14</td><td></td><td>Abies alba</td><td>Poz-153130</td><td>150 ± 30</td><td>1667–1903 cal AD</td><td>284–48 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>916</td><td>CK10</td><td></td><td>Fagus sylvatica</td><td>Poz-153151</td><td>185 ± 30</td><td>1653–1915 cal AD</td><td>297–35 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Fagus sylvatica</td><td>Poz-153293</td><td>5 ± 30</td><td>1695–1916 cal AD</td><td>255–34 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>971</td><td>CK16</td><td></td><td>Fagus sylvatica</td><td>Poz-153379</td><td>115 ± 30</td><td>1680–1940 cal AD</td><td>270–10 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>954</td><td>CK17</td><td></td><td>Fagus sylvatica</td><td>Poz-153273</td><td>240 ± 30</td><td>1525–1929 cal AD</td><td>425–21 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>769</td><td>CK2</td><td></td><td>Fagus sylvatica</td><td>Poz-167142</td><td>60 ± 30BP</td><td>1694–1918 cal AD</td><td>257–33 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1672</td><td>CK5</td><td></td><td>Abies + Fagus</td><td>Poz-167143</td><td>105 ± 30BP</td><td>1683–1936 cal AD</td><td>266–14 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1682</td><td>CK4</td><td></td><td>Abies + Fagus</td><td>Poz-167144</td><td>205 ± 30BP</td><td>1643–1920 cal AD</td><td>306–31 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1601</td><td>CK7</td><td></td><td>Abies + Fagus</td><td>Poz-167149</td><td>295 ± 30BP</td><td>1695–1908 cal AD</td><td>450–291 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1619</td><td>CK6</td><td></td><td>Abies + Fagus</td><td>Poz-167146</td><td>100 ± 30BP</td><td>1683–1930 cal AD</td><td>267–21 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1663</td><td>CK1</td><td></td><td>Abies + Fagus</td><td>Poz-167147</td><td>100 ± 30BP</td><td>1683–1930 cal AD</td><td>267–21 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1667</td><td>CK2</td><td></td><td>Abies + Fagus</td><td>Poz-167148</td><td>150 ± 30BP</td><td>1667–1903 cal AD</td><td>284–48 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1696</td><td>CK3</td><td></td><td>Abies + Rhododendron</td><td>Poz-167156</td><td>1215 ± 30BP</td><td>609–890 cal AD</td><td>1252–1061 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1726</td><td>CK6</td><td></td><td>Abies + Betula</td><td>Poz-167157</td><td>90 ± 30BP</td><td>1687–1926 cal AD</td><td>263–25 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1785</td><td>CK7</td><td></td><td>Abies</td><td>Poz-167159</td><td>905 ± 30BP</td><td>1041–1217 cal AD</td><td>909–734 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1794</td><td>CK8</td><td></td><td>Conifer + Lonicera</td><td>Poz-167160</td><td>140 ± 30BP</td><td>1672–1908 cal AD</td><td>270–61 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1695</td><td>CK2</td><td></td><td>Abies + Fagus + Vaccinium</td><td>Poz-167158</td><td>35 ± 30BP</td><td>1694–1917 cal AD</td><td>256–33 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1684</td><td>CK3</td><td></td><td>Vaccinium</td><td>Poz-167161</td><td>1025 ± 30BP</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>Abies + Rhododendron</td><td>Poz-167150</td><td>250 ± 30BP</td><td>902–1150 cal AD</td><td>1049–800 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1684</td><td>CK4</td><td></td><td>Abies</td><td>Poz-167152</td><td>465 ± 30BP</td><td>1410–1464 cal AD</td><td>541–407 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1525</td><td>CK1</td><td></td><td>Abies</td><td>Poz-167153</td><td>85 ± 30BP</td><td>1680–1924 cal AD</td><td>262–26 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1515</td><td>CK2</td><td></td><td>Abies</td><td>Poz-167154</td><td>170 ± 30BP</td><td>1660–1908 cal AD</td><td>291–61 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1499</td><td>CK3</td><td></td><td>Abies</td><td>Poz-167155</td><td>115 ± 30BP</td><td>1680–1940 cal AD</td><td>270–10 cal BP</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1502</td><td>CK4</td><td></td><td>Abies + Fagus</td><td>Poz-167153</td><td>115 ± 30BP</td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td>1290</td><td>CK1</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>

A comparative analysis of the Carpathians and the Pyrenees reveals notable differences in both vegetation succession and fire intensity over time (Fig. 8B). Indeed, there is a notable shift in the timing of peak fire activity, specifically the third fire period for which we have obtained the maximum radiocarbon dates. This shift spans approximately 3000 years and can be observed between the Carpathians, which begin around 770 cal. BP, and the Pyrenees, which began around 3500 cal. BP. In contrast to the Pyrenees, where the number of fires per 250-year period reaches its highest point between 3750 and 1500 cal. BP, subsequently declining in the recent period, in Romania this number is consistently rising, reaching its peak in the current period (Fig. 8A, B). Our reconstruction of the Holocene vegetation dynamics and fire activity in Pyrenees, as evidenced by pollen and charcoal analysis, are nevertheless aligned with the prevailing syntheses available for this part of the massif (Jalut et al., 1988, 1996, 1998; Galop and Jalut, 1994; Heinz and Barbaza, 1998). Notwithstanding the absence of pine trees, which would have marked the initial post-glacial recolonisation phase, and the identification of birch charcoal dating from this period (9077-8650 cal. BP) is consistent with known reconstructions. The presence of rhododendron (9681-9486 and 9884-9429 cal. BP) and maple (10750-10440 cal. BP) indicates the existence of open environments and open heathland, which may align with the ecosystems of the pine-oak-birch phase of the early Holocene. Prior research has established the preeminent role of *Acer* trees in mountainous regions during the post-glacial recolonisation of forests of the early Holocene (Carcaillet and Brun, 2000). The rhododendron has persisted in the Pyrenees during

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glacial periods in a variety of refugia, including lowland refugia in the eastern portion of the mountain range and nunataks at high elevations (Charrier et al., 2014). This adaptability likely facilitated its early recolonisation following the glacial retreat. The second fire phase reveals the presence of deciduous oak (7157-6802 cal. BP) and hazel (7261-7015 cal. BP), in line with previous vegetation reconstructions (Jalut et al., 1988, 1992, 1998; Galop and Jalut, 1994; Heinz and Barbaza, 1998). Indeed, the *Quercus-Corylus* phase in the Pyrenees has been documented to extend from approximately 10000 to 7250 cal. BP (Jalut et al., 1992, 1998; Heinz and Barbaza, 1998). The dates obtained for both species are slightly more recent than the known end of this phase. However, it should be noted that vegetation successions are not sudden phenomena, but rather slow, responding to changes in the environmental context. Furthermore, time lags between different phases are well documented in the Pyrenees, due to differences in climate and ecological contexts along the east-west gradient (Jalut et al., 1992, 1998). The third phase of fire activity appears to demonstrate an acceleration in fire frequency, as evidenced by 22 radiocarbon dates. As previously stated, although comparable to the phenomenon observed in the Carpathians, this third phase began much earlier in the Pyrenees at the advent of the Early Bronze Age (started around 4150 BP) and spanned a considerably longer period, exceeding 4000 years. The third phase also demonstrates the early appearance of fir at approximately 3500 cal. BP. This represents a further significant divergence in vegetation trajectories with the Carpathians, namely the advent of fir precedes that of beech. As with rhododendron, a substantial body of evidence from numerous studies indicates that fir established glacial refuges in the Pyrenees (Cheddadi et al., 2014; Scotti-Saintagne et al., 2021), thereby enabling it to reclaim the montane level as soon as climatic conditions became favourable. The principal Holocene reconstructions of vegetation in the Central Pyrenees indicate that the expansion of fir started approximately 5000 yr BP. Although beech also survived in glacial refuges, it was only after 4000 years BP that beech became a dominant component of Pyrenean forests (Jalut et al., 1992, 1998, Galop and Jalut, 1994).

## 4.2 Current European OGF Mountain Fir-beech forest inherited from long-term human-forest interaction

The oldest direct evidence of human activity was found in the Carpathians, at the site of Bistra, one of the least accessible nowadays. It dates back to the Early Bronze Age (c. 2000 cal. BC) and precedes the locally recorded fire events of 1200-1000 cal. BC. This very old CK platform, the topography of which is relatively well preserved (Py-Saragaglia, 2024), also testifies to the past use of yew for charcoal manufacturing, although this species is mainly known to be used to make bows and tools during prehistoric/protohistoric times (Pérez-Díaz et al., 2013; Uzquiano et al., 2015). Despite its well-known toxicity (Thomas and Polwart, 2003), this tree species with specific mechanical properties was of particular interest to past societies and could make yew-rich stands attractive for humans (Hageneder, 2007). Throughout Europe, during the Bronze Age, in search of various resources such as copper and tin ore deposits, people explored and seasonally occupied remote and mountainous areas to
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record heights (Marembert, 2000; Rendu, 2003; Pelisiak et al., 2018; Saint-Sever & Remicourt, 2019).

570

Despite its remote location, this Bronze Age CK may be the evidence of human mobility in search of raw materials, and yew charcoal may have been produced for the thermal treatment of ores or in

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## Romania

<figure><img src="image_11.png" /></figure>

<figure><img src="image_12.png" /></figure>

## France

<figure><img src="image_13.png" /><figcaption>Fig. 6. Radiocarbon dates obtained with charcoals from CKs in Romania (A) and in France (B).</figcaption></figure>
metalworking. Although direct archaeological traces of such activity are very rare in the area under consideration, and the low number of mines with undoubted evidence of Bronze Age exploitation (Kiss and Romhányi, 2023), its significant endowment of polymetallic ores, very rich in base and precious metals (Heinrich and Neubauer, 2002), may have led to rudimentary mining (ores collected on the surface of outcrops) and smelting practices. In the Northern Carpathians, the Bronze Age, which is globally characterised by a forest-dominated landscape, is also known for small-scale agricultural activity at lower altitudes, evidenced low levels of *Cerealia* and *Secale*-type pollen, and the increasing use of higher mountain areas for pasture (Feurdean et al., 2013; Geantă et al., 2014). Despite the low level of human practices and the moderate fire activity (between 3000 and 2300 cal. BP) recorded in the Tăul Roșia peat bog (Mîndrescu et al., 2023), several fire events recorded at the heart of Poiana, Bistra and Băiuț OGFs, occurring c. 1700 and 1000 cal. BC and burning beech trees, may be related to the use of fire to increase grazing resources (Feurdean et al., 2012).

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<figure><img src="image_14.png" /><figcaption>Fig. 7. Relative frequencies of taxa identified in CKs, classified from oldest to most recent. Before the CK code (site, n° plot, n° CK), the prefix BA means: Bronze Age, M = Medieval, M-EMT = Medieval-Early Modern Times, Mo = Modern.</figcaption></figure>

Our results also reveal the unsuspected place of yew around 4000 cal yrs BP in the mountainous spruce-dominated forest where beech and fir were already present (Farcaş et al., 2013; Geantă et al., 2014), as attested by soil charcoals. Yew is a shade-tolerant and long-lived species generally poorly detected by palynological studies (Pérez-Díaz et al., 2013) while both its distribution and ecological ranges are wide (Caudullo et al., 2017; Dumé et al., 2018; Kaiser et al., 2024). It was still present in the beech-fir forest of Bistra in the late Middle Ages-early modern times and used with fir to make charcoal.
This result suggests that yew was readily available in the local environment and that the exploited forest had preserved some maturity attributes. In this same OGF site, yew is found again in a Modern times CK, but it occurs in small proportions, mixed with fir, spruce and beech. As highlighted elsewhere in Europe, it probably disappeared from the forest during this period as a result of increasing human pressure (charcoal making, forest management strategies favouring more fast-growing species, livestock grazing). Historical sources also show that between the 13th and 16th centuries, yew trees were exported in large quantities from Central Europe to England for the manufacture of longbows (Hilf, 1925; Hageneder, 2007). A similar phenomenon has been observed in the northern central Pyrenees, where yew has been found in both soil and archaeological charcoal dating from the Bronze Age to modern times, in areas that have long been shaped by human activities and are currently devoid of this species (Py-Saragaglia et al., 2017; Saulnier et al., 2020; Fouédjeu et al., 2021).
<figure><img src="image_15.png" /></figure>

<figure><img src="image_16.png" /><figcaption>Fig. 8. Schematic representation of forest vegetation dynamics since the superior Pleistocene (mid and Late dryas) to the current period for the Carpathians (A) and Pyrenees (B) based on pollen reconstruction and charcoal particles.</figcaption></figure>

In the northern Carpathians and particularly in the Maramureş region, human pressure have significantly intensified during the late Iron Age, around 2500 years ago (Vulpe, 2001; Feurdean et al., 2010, 2013; Fărcaş et al., 2013; Mîndrescu et al., 2023). Archaeological and palaeoecological studies indicate that this period witnessed a marked increase in land use intensity, with extensive clearings and the subsequent spread of plant species associated with human activity. But the massive expansion of human occupation and the associated clearing in lowlands has occurred from the Roman period (c. 2000-
1700 cal. BP), and especially from the Middle Ages onwards. Although societies have occupied and used forests for thousands of years, human activities have intensified in mountain areas and, as a result, the pressures on forest resources (Geantă et al., 2014). From this period onward, historical archives reports large-scale clearance, due to harvesting for construction and mining, but also for the settlement of land for agriculture (Giurescu, 1980; Csucsuja, 1998; Kaplan et al., 2009; Feurdean et al., 2010; Py-Saragaglia et al., 2020). The increasing demand of wood probably modified forest features throughout more regular harvesting and forest management strategies (planting) favouring fast-growing conifers at the expense of beech (Wollmann, 1996; Feurdean et al., 2010), particularly from the 17th-18th c. as elsewhere in western and central Europe (Savill, 2015). The Austro-Hungarians, who controlled the area and the management of the mines, belonged to the German classical school of forestry (Von Carlowitz, 1713) and are known to have favoured spruce plantations, especially for mining (firesetting and timbering). In the studied OGFs, spruce was occasionally used to make charcoal, but fir and beech were preferred. The high number of radiocarbon dates from soil charcoal obtained in OGFs as well as the presence of light demanding species such as *Acer* spp. and *Salix* spp. supports the intensification of forest use by widespread forest grazing of livestock and wood harvesting. Finally, from the last two hundred years, the development of industrial wood exploitation is also evidenced by the increasing anthropogenic disturbance events detected in mature stands (Kaplan et al., 2009; Feurdean et al., 2010; Py-Saragaglia et al., 2020). In addition, the increasing number of radiocarbon dates obtained from CK's charcoals may also reflect the large-scale charcoal making practices to supply forging and smelting activities. All of these human practices raise issues about their consequence on the overall functioning of forest and may explain the delay in biodiversity recovery (Retrogression; see paper of Pokorny et al., 2022).

In the northern Pyrenees, the Bronze Age is an important period of anthropisation of mountain and subalpine areas with an increase of agropastoral use (Galop et al., 2007, 2013; Carozza & Galop, 2008; Rendu et al., 2013; Saulnier et al., 2020) and human pressure on forests, related both to mining activities (Galop et al., 2001; Monna et al., 2004) and to an increase in fire events associated with land use changes (Rius et al., 2009; Bal et al., 2010). In the OGFs studied, the Middle Bronze Age and the Bronze Age-Iron age transition (c. 1600-1500 and 1000 cal. BC) is also characterised by a significant increase in fire activity, affecting the fir-dominated forest, probably related to human practices, possibly patchy land clearance for livestock grazing. As it was highlighted elsewhere in the northern central and eastern Pyrenees, increasing human activity in fir-dominated forests probably contributed to the expansion of beech, especially from historical periods (Galop and Jalut, 1994; Py-Saragaglia et al., 2017; Saulnier et al., 2020; Fouédjeu et al., 2021, 2022). The Espingo pollen diagram shows that the fir-beech forest was already present in the region during the Roman period, when anthropogenic activities are well characterised. In the OGFs studied, fire events affecting beech, maple and fir occurred regularly from the beginning of the Iron Age until Late Antiquity. Charcoal-making activities have been documented since the early Medieval period, as in the Vicdessos valley (Ariège), where local
communities exploited iron ore deposits (Fouédjeu et al., 2022). This practice intensified in Modern times with the increased use of beech mixed with fir, particularly in Barrada and, to a lesser extent, Burat and Bois-Neuf. At the same time, fir and beech fires have become more frequent since the Middle Age. On a regional scale, the sharp increase in human practices since 1000 cal. AD has led to a reduction in forest cover at all altitudes and an intensification of the fire regime. In the heart of the OGFs, despite increasing forest use and local canopy opening events, also evidenced by soil insect remains (ongoing study), local communities have globally maintained the continuity of forest cover to provide fuel and timber resources until the 20th c. At least from the 19th c., the fir was probably favoured by high forest management, without totally excluding cattle grazing. For example, in the Burat forest, historical archives show in the 19th c. a major shift in stand composition, with a transition from a beech-dominated forest to a fir-dominated forest, which is clearly related to harvesting (Borde, 2024). The Bois-Neuf forest was already a fir forest in the 19th c., which means that this species has been managed as a high fir forest for much longer.

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## Conclusion

The combination of palaeoecological proxies reveals divergent Holocene trajectories of the fir-beech OGF between French central Pyrenees and Romanian Carpathians, yet the current OGF sites present high similarity. Pollen data reconstructions offer a relatively brief temporal record that aligns closely with established regional trajectories. In contrast, charcoal analysis has provided insights into vegetation succession and fire episodes across a broader temporal range, also corroborating regional and local reconstructions. The convergence of these disparate pathways, culminating in the emergence of contemporary forest ecosystems, gives rise to inquiries concerning their purely ecological heritage. Archaeological and palaeoecological data also suggest that past societies have played a significant role in the formation of the mixed beech and fir forests that are currently regarded as the baseline of natural forest ecosystems in the mountain belt of temperate Europe. The influence of human societies on natural dynamics is evident in both areas under study from the Bronze Age onwards. From the Middle Ages onward, the increasing demand for timber and fuel wood has resulted in the proliferation of fast-growing conifers (spruce, fir) favored by high forest management systems. Additionally, human practices have led to significant alterations in vegetation composition, including the removal of yew. In some cases, such as at Poiana, the current dominance of beech may be the result of past practices that have favored beech over fir and spruce, including grazing and coppice systems. The OGFs under examination are thus the result of this interaction and cannot be considered relics of the primeval forest that preceded the advent of farming. The current high level of maturity can be attributed to several factors, including the long-term continuity of the forest cover despite human activities and small-scale canopy opening events. Additionally, despite charcoal burning practices and logging activities, a certain tree diversity has been preserved. Furthermore, mature elements and deadwood have been conserved in the stands. Most importantly, the cessation of practices sometimes associated with the implementation of rigorous
protective measures has contributed to the current state of the forests. This is evident in the examples of
682
Burat, Poiana Botizii, and Bistra. The last 200 years thus appear to be of pivotal importance, and we are
683
currently engaged in a process of studying them using appropriate methodologies, including dendrology
684
and dendrochronology on both living trees and deadwood, in conjunction with a detailed examination
685
of historical archives. The results demonstrate that the composition of OGFs is closely associated with
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historical human activities. To effectively conserve biodiversity, it is essential to consider the maturity,
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stand complexity, and landscape heterogeneity of the forest. This necessitates the implementation of
688
appropriate management strategies, including the use of natural disturbance regimes to inform practices,
689
the reintroduction of yew, and the expansion of protected areas.
690

691

692 Acknowledgements

The French National Research Agency (ANR) funded this research as part of the BENDYS project (The
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last European old-growth fir-BEech forests: a loNg term and global stuDy for their better underStanding,
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conservation and management - ANRJCJC-19-CE03-0010) coordinated by Vanessa Py-Saragaglia. This
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research also benefited from the support of the FORETEXIL project, also coordinated by Vanessa Py-
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Saragaglia and funded by the Mission for Interdisciplinary and Transdisciplinary Initiatives (MITI) of
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the National Centre for Scientific Research (CNRS, France). The SAFE NATURE project, coordinated
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by C'ecile Brun and funded by the Maison des Sciences de l'Homme et de la Société de Toulouse
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(MSHS-T, University of Toulouse, France), supported the field work for the Taul Roşia sampling. We
700
would like to thank all the BENDYS team and in particular Antoine Brin, Sandrine Buscaino, François
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De Vleeschouwer, Evan Fisher, Vincent Labbas, Gaël Le Roux, Théo Maisonhaute, Marcel Mîndrescu,
702
Ancuţa Petraş, Magali Philippe, Nicolas Poirier, Alexandre Raimbault, Coralie Roy-Camille, Mélanie
703
Roy, Agnes Ruskal, Călin Gabriel Tămaş and Ruppert Vimal for their fruitful help during fieldwork
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and/or laboratory analyses. We would also like to express our gratitude to the Remote Primary Forest
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research group, particularly Miroslav Svoboda and Martin Mikoláš, for their invaluable guidance and
706
the provision of their comprehensive database. We would like to thank Ocolul Silvic Strâmbu Băiuţ
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(Romsilva), especially Florin Mârzac, but also WWF Romania, especially Călin Ardelean, Parcul
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Natural Munţii Maramureşului, especially Cătălina Bogdan, and ONF (Office National des Forêts) for
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all the fieldwork permits and their efficient cooperation.
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Figures and Tables

Figure 1: Maps of the study area, with the six studied OGF sites, the Tăul Roșia peat bog and the Espingo Lake. In the Romanian Carpathian: (1) Tăul Roșia peat bog, (3) Baiut and (4) Poiana Botizii within the Strâmbu Băiuț forest, and (5) Bistra in the valley of Bistra. In the French Pyrenees: (2) the Espingo lake, (6) Burat natural forest reserve in Marignac, (7) Bois-Neuf in Saint-Mamet, (8) Barrada in Gèdre-Gavarnie and the lake of Espingo. More detailed locations of all data census are presented in supplementary materials (Figure S1A and B).

Figure 2: Simplified pollen diagram, charcoal accumulation rate (CHAR, only for Espingo lake, expressed in mm⁻².cm⁻².yr⁻¹), and selected pollen and NPP percentage for (A) the Tăul Roșia peat bog and (B) the Espingo lake. (A) For the Tăul Roșia peat bog the exaggerated curves (light colour) are ×5. The pollen diagrams are divided into three ‘pollen zones’, which have been computed using CONISS zonation in Tilia. (B) For the Espingo lake, the exaggerated curves (light colour) are ×10. The pollen diagrams are divided into five ‘pollen zones’, which have been computed using CONISS zonation in Tilia.

Figure 3: Soil Anthracomasses per Pits (SAP) in forest soils in France (left) and in Romania (right). The SAPs are expressed as the ratio between the weight of charcoals in grams and the weight of the dry soil in kilograms.

Figure 4: Specific anthracomasses per taxa and layer (SAT in g/kg) in the 16 pits from Pyrenees (A-F) on the left and from the Carpathians on the right (G-P).

Figure 5: Radiocarbon dates of charcoal pieces sampled in soil pits from Carpathians (A) and Pyrenees (B).

Figure 6: Detailed maps of the six studied OGF sites in Romania (A) and Pyrenees (B). (A) At top, the general map of the Romanian Carpathians OGF sites with Băiuț and Poiana, within the Strâmbu Băiuț forest, and the forest of Bistra. At the bottom, detailed maps of the three study areas: Băiuț S1.4; Poiana Botizii S1.2; Bistra S1.4. (B) At top, the general map of Pyrenean OGF sites: the fir forest of Barrada in Gèdres-Gavarnie, the forest of Bois-Neuf in Saint-Mamet and the Burat natural forest reserve in Marignac. The dark green areas represent forest cover and the light green areas represent agro-pastoral zones. The red crosses represent the pits, the black triangles the charcoal kiln platforms, the green dots the pastoral huts and the mine symboles (hammer and pick crossed) the main mines. OGF sites are outlined in black and the circular plots surveyed are outlined in grey.

Figure 7: Radiocarbon dates obtained with charcoals from CKs in Romania (A) and in France (B).

Figure 8: Relative frequencies of taxa identified in CKs, classified from oldest to most recent. Before the CK code (site, n° plot, n° CK), the prefix BA means: Bronze Age, M=Medieval, M-EMT=Medieval-Early Modern Times, Mo=Modern.

Figure 9: Schematic representation of forest vegetation dynamics since the superior Pleistocene (mid and Late dryas) to the current period for the Carpathians (A) and Pyrenees (B) based on pollen reconstruction and charcoal particles.
1089 **Table 1:** Main features of the OGF sites including the country, the location, the name of OGF sites and number on the map (FIG. 1), the latitudes and longitudes of the centroids of the sites, the altitudinal ranges (m a.s.l.), the total forest surface (ha), the number of 1ha-Plot surveyed, and the main tree species present in plots.

1093 **Table 2:** Radiocarbon dates of plant remains from the Taul Rosia peatbog and from the Espingo lake used to calibrate the age depth model

1095 **Table 3:** The number of soil charcoal remains identified by major taxonomic groups is provided below. In parentheses are the species to which the charcoal most certainly belongs, given the local ecological conditions.

1098 **Table 4:** Radiocarbon dates of charcoal remains from soil and former charcoal kiln platforms.
1099
1100
1101
1102