# Neolithic societies and landscape transformations in northwestern France: A high-resolution multi-proxy study including sedimentary DNA

Laurence Lemer, Agnès Gauthier, Laurent Lespez, Antony Brown, Samuel

Hudson, Marta Garcia, Kazuyo Tachikawa, Edouard Bard, François Giligny,

José Antonio López-Sáez

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## To cite this version:

Laurence Lemer, Agnès Gauthier, Laurent Lespez, Antony Brown, Samuel Hudson, et al. Neolithic societies and landscape transformations in northwestern France: A high-resolution multi-proxy study including sedimentary DNA. Quaternary Science Reviews, 2025, 369, pp.109644. 10.1016/j.quascirev.2025.109644. hal-05312490

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# Neolithic societies and landscape transformations in northwestern France: A high-resolution multi-proxy study including sedimentary DNA

Laurence Lemer<sup>a,b,c,*</sup>, Agnès Gauthier<sup>d</sup>, Laurent Lespez<sup>a,e</sup>, Antony Brown<sup>f,g</sup>,

Samuel Hudson<sup>f</sup>, Marta Garcia<sup>h</sup>, Kazuyo Tachikawa<sup>h</sup>, Edouard Bard<sup>h</sup>, François Giligny<sup>b,c</sup>,

José Antonio López-Sáez<sup>i,b</sup>

## ARTICLE INFO

Handling Editor: Dr Donatella Magri

Keywords:

Palaeoenvironments

Anthropization

Geoarchaeology

Paleogeography

Continental biomarkers

Vegetation dynamics

Palynology

Ancient DNA

Holocene

Western europe

## ABSTRACT

To understand the impact of human practices on Mid-Holocene vegetation in northwestern France, more pollen records and multi-proxy studies with accurate chronologies located near excavated archaeological sites are needed. In northwestern Normandy, the Caen Plain is particularly suitable for gaining further insight into the impact of early human activity on the landscape, due to the high density of recognized and excavated archaeological sites dating back to Neolithic times. Therefore, in this area, we developed a high temporal resolution approach using multiple palaeobiological proxies (pollen, non-pollen palynomorphs (NPPs), charcoal analysis, and sedimentary ancient DNA (sedaDNA)), along with sedimentological proxies to analyze a new sequence from the Vey valley, located close to the excavated Neolithic site of Cairon, and which could be compared to the archaeobotanical and archaeozoological data from archaeological excavations. Our specific objective was to propose a reconstruction of the landscape changes under human influence at a local scale, in order to gain a precise understanding of the processes of anthropization in the Caen Plain at the end of the Mesolithic and during the Neolithic. The impact of human activity during the Mesolithic and Early Neolithic appears to have been limited. Nevertheless, the end of the Late Mesolithic (5900–5200 BCE; 7850–7150 cal. BP) showed the first increase in local fire frequency. These fires were responsible for temporary clearings of the forest along the wetland, and sedaDNA indicated that wild boar were present in the wetland. Most notably, the Cairon site offered the first opportunity to precisely determine the impact of changing land use in an area that had been inhabited since the Middle Neolithic (4400–3475 BCE; 6350–5425 cal. BP). The combined use of sedaDNA, NPPs, and charcoal analysis demonstrated the importance of pastoral practices and fires in opening up environments and transforming alluvial wetlands during this period.

## 1. Introduction

Over the past few decades, pollen-based reconstructions of past vegetation in quantitative terms —under both climatic and human

influences—have been produced at regional scales (e.g. David et al., 2012; Fyfe et al., 2013; Lechterbeck et al., 2014; Nielsen et al., 2012) and continental scales (e.g. Davis et al., 2015; Fyfe et al., 2015; Githumbi et al., 2022; Marquer et al., 2014, 2017; Roberts et al., 2018;

<footnote><sup>a</sup> Laboratoire de Géographie Physique, UMR CNRS 8591, 2 rue Henri Dunant, 94320, Thiais, France</footnote>

<footnote><sup>b</sup> Trajectoires, UMR CNRS, 8215, 9 rue Malher, 75004, Paris, France</footnote>

<footnote><sup>c</sup> Université de Paris1 Panthéon-Sorbonne, Institut d'art et d'archéologie, 3 rue Michelet 75006, Paris, France</footnote>

<footnote><sup>d</sup> Laboratoire de Géographie Physique UMR CNRS 8591 - Universités Paris I - UPEC, 2 rue Henri Dunant, 94320, Thiais, France</footnote>

<footnote><sup>e</sup> Université de Paris-Est-Créteil (UPEC), Département de Géographie, 61 avenue du Général De Gaulle, 94010, Créteil, France</footnote>

<footnote><sup>f</sup> Geography and Environmental Science, University of Southampton, Southampton, UK</footnote>

<footnote><sup>g</sup> ArcEcoGEN Research Centre, The Arctic University Museum of Norway,UiT - The Arctic University of Norway, Tromsø, Norway</footnote>

<footnote><sup>h</sup> Aix Marseille Univ, CNRS, IRD, INRAE, Coll France, CEREGE, Aix-en-Provence, France</footnote>

<footnote><sup>i</sup> Environmental Archaeology Research Group, Institute of History, CCHS, CSIC, Albasanz 26-28, 28037, Madrid, Spain</footnote>

<footnote>* Corresponding author. Laboratoire de Géographie Physique, UMR CNRS 8591, 2 rue Henri Dunant, 94320, Thiais, France.</footnote>

<footnote>E-mail addresses: laurence.lemer@cnrs.fr (L. Lemer), agnes.gauthier@lgp.cnrs.fr (A. Gauthier), Laurent.lespez@lgp.cnrs.fr (L. Lespez), antony.g.brown@uit.no (A. Brown), S.Hudson@soton.ac.uk (S. Hudson), garcia@cerege.fr (M. Garcia), kazuyo@cerege.fr (K. Tachikawa), bard@cerege.fr (E. Bard), Francois.Giligny@univ-paris1.fr (F. Giligny), joseantonio.lopez@cchs.csic.es (J.A. López-Sáez).</footnote>
Trondman et al., 2015; Woodbridge et al., 2018) using databases such as the European Pollen Database (Fyfe et al., 2009), Neotoma Paleoecology Database and Pangea (Fyfe et al., 2015). In northwestern Europe, the first half of the Middle Holocene (around 6650–4050 BCE; 8600–6000 cal. BP) was a period of broad-leaf forest development, which constituted the dominant land cover. Locally, indicators of arable cultivation or changes in woodland composition may have been linked to an early wave of Neolithic agriculture (e.g. Brown, 1997, 1999; Kalis et al., 2003; Leroyer et al., 2004; Richard, 2000; Visset et al., 2002). Nevertheless, a progressive decline in forest cover became apparent on a broader scale from around 4050 BCE (6000 cal. BP), when the Neolithic was established across much of mid-latitude Europe. Subsequently, forest loss varied in both intensity and timing, but much of the landscape had already been cleared during the Bronze Age and Iron Age due to forest conversion to farming and grazing from approximately 2050 BCE (4000 cal. BP). Therefore, we needed to consider more precisely the impact of early human activity on the landscape prior to these large-scale clearings.

Pollen analysis is increasingly coupled with other bioindicators, such as non-pollen palynomorphs (NPPs), charcoal analysis, and sedimentary ancient DNA (sedaDNA). When combined both complementary and orthogonal (or Sankey diagram sensu Bell and Blais, 2021) proxies can provide deeper and more robust insights into the role of past societies in environmental dynamics (e.g. Albert et al., 2021; Glais et al., 2016, 2023; Gouriveau et al., 2020; Hudson et al., 2022, 2023; Innes et al., 2010, 2013; Jouffroy-Bapicot et al., 2013; Revelles and van Geel, 2016). However, much of the pollen data used in regional overviews is outdated, based on a small number of radiocarbon dates with large margins of error and/or derived from wetlands located far from known Neolithic settlements (Giesecke et al., 2014; Leroyer, 2006; Woodbridge et al., 2018). Therefore, to better understand the impact of human practices on mid-Holocene vegetation, more pollen records and multi-proxy studies with accurate chronologies located near excavated archaeological sites are needed. This is challenging but possible in northwestern France, where the best-dated pollen sequences are old and concentrated downstream of the Loire valley and its tributaries, and in the central part of the Paris basin (e.g. Barbier et al., 2002; Cyprien et al., 2004; Joly-Delanoë and Visset, 2011; Leroyer, 2006; Ouguerram and Visset, 2001; Visset et al., 2001, 2002, 2005). Finally, although still rare in northwest France, some existing studies demonstrate the great potential of multiproxy palaeobiological analyses—combining the analysis of pollen, NPPs and/or charcoal—to reconstruct vegetation dynamics under anthropogenic impact (De Milleville et al., 2023; Gauthier, 1995; Reinbold, 2017).

In this context, the aim of this paper was to provide new data to further understand the impact of early human activity on the landscape of the plains and plateaus of northwestern France since the Neolithic. The Caen Plain was particularly suitable, being located close to a high density of recognized and excavated archaeological sites. In fact, the colonization of the plateaus by farmers from the Linear Pottery culture has been documented from the end of the 6th millennium BCE (Billard and Alix, 2014; Billard and Legris, 2010; Le Saint-Allain et al., 2021) and has stimulated research on the impact of the earliest agricultural activities. Nevertheless, the first anthropogenic pollen indicators in sedimentary sequences was dated to the Middle Neolithic onwards (Lespez, 2012b; Lespez et al., 2005), but the low-resolution timescale did not allow us to discern the impact of agropastoral activities and their changes during the Neolithic. Therefore, we adopted a high temporal resolution approach using multiple palaeobiological proxies (pollen, NPPs, charcoal analysis and sedaDNA) to analyze a new sequence from the Vey valley, located close to the excavated Neolithic site of Cairon, and compared it to the archaeoenvironmental data from archaeological excavations. Our specific objective was to propose a reconstruction of landscape changes under human influence at a local scale, in order to gain a precise understanding of the processes of anthropization in the Caen Plain at the end of the Mesolithic and during the Neolithic.

## 2. Study area

### 2.1. Environmental setting

The Caen Plain is located in the westernmost part of the Paris basin, where the Jurassic (Bathonian) limestone bedrock is highly permeable, which explains the low density of the hydrographic network (Fig. 1). Mainly regulated by groundwater, the low-energy rivers meander through the bottoms of incised valleys, where sedimentary records have been documented (Lespez, 2012a; Lespez et al., 2005, 2008). Wetlands are dominated by meadows and woodlands, contrasting with the monotonous landscape of the plateaus, where the highly fertile loamy loessic soils have favored the development of vast cereal and oilseed fields.

The Vey is a tributary of the Mue river, which in turn is the final tributary of the Seulles river (Fig. 1). The hydrological and sedimentary functioning of the Seulles river and its tributaries is well documented (Beauchamp et al., 2017a, 2017b; Lespez et al., 2015; Viel et al., 2014a, 2014b, 2024). In the Mue catchment, sedimentary archives from valley bottoms have been investigated and dated at several sites (Lespez et al., 2008, 2012). Geomorphological studies have frequently been combined with palaeoecological and palynological analyses (Lespez et al., 2005, 2008, 2012). The rivers are supplied by carbonate-rich water from the Bathonian aquifer, and during the first half of the Holocene, the dissolved load led to the formation of tufa dams in the alluvial plains. At the beginning of the Middle Holocene, organic sequences that accumulated preferentially behind these dams provided well-preserved pollen records.

The Vey is a small river approximately 1.3 km long, flowing through a narrow valley (maximum width of about 100 m) (Fig. 2A). On the hillsides, rendzina soils (Fig. 2B) develop over limestone bedrock, whereas the plateaus are mainly covered by luvisols derived from loess dating from the Last Glacial (Viel et al., 2024). In the valley bottom, redox-dominated soils are exposed to seasonal water excess, resulting in hydromorphic conditions below 50 cm from the soil surface. The open landscape is dominated by meadows in the valley bottom and cereal crops on the hillsides and plateaus.

### 2.2. Previous research and archaeological setting

In western Normandy, following the pioneering work of Elhai (1963), pollen data has been used to define regional vegetation changes during the Holocene. However, this research has been limited by poorly dated or undated sequences (Clet-Pellerin, 1985; Clet-Pellerin et al., 1977, 1981, 1993; Clet-Pellerin and Verron, 2004; Huault, 1972). A resurgence of palaeoenvironmental research from the 2000s onwards has produced new data, driven by the development of an interdisciplinary approach in connection with archaeological findings (Coutard and Clet-Pellerin, 2006; Lemer et al., 2018; Lespez et al., 2004, 2005, 2008, 2010; Lespez, 2012a). Indeed, the open-field landscape of the Caen Plain was favorable for identifying archaeological remains, and furthermore, urban development projects around Caen have stimulated numerous preventive archaeological interventions. Thus, many archaeoenvironmental studies (mainly anthracological, carpological, archaeozoological and pedological) have provided valuable information on plant and animal exploitation from the Neolithic to the historical period. In this context, the first geoarchaeological investigations have been conducted as part of the *Landscape archaeology research program on the Caen Plain from the Neolithic*, and outlined initial reconstructions of past landscape dynamics (Germain-Vallée and Lespez, 2016, 2018; Lespez et al., 2013; Lespez and Germain-Vallée, 2011).

The Vey valley is an archaeological area characteristic of the Caen plain (Lespez et al., 2013). Within a 1-km radius of the valley bottom, several Neolithic sites have been identified (Fig. 2A), including two permanent domestic settlements on the hillside between the Mue and Vey valleys, which have been assigned to the Villeneuve-Saint-Germain
<figure><img src="image_3.png" /><figcaption>Fig. 1. Archaeoenvironmental and paleoenvironmental data in the Caen Plain (modified from Germain-Vallée and Lespez, 2016).</figcaption></figure>

(VSG) culture (Early Neolithic: 5000–4700 BCE; 6950–6650 cal. BP) (Germain-Vallée et al., 2017; Giazzon, 2012). The most emblematic and best-studied site, La Pierre Tourneresse, is located around 250 m from the valley bottom. Successive archaeological excavations have revealed the remains of three distinct phases of occupation, which were dated using the radiocarbon method (Ghesquière and Marcigny, 2011). The construction of a funerary monument led to the preservation of a subsoil with remains interpreted as part of a religious area, dated to the Middle Neolithic I period (4400 BCE; 6350 cal. BP). There were also remains associated with a domestic settlement dated to the Early Middle Neolithic II period (4200–4000 BCE; 6150–5950 cal. BP).

Archaeozoological and archaeobotanical data from this settlement pointed to the development of agro-pastoral activities. After this settlement was abandoned, the funerary monument was constructed around 3900 BCE (5850 cal. BP) and was used until the end of the Middle Neolithic II period (3500 BCE; 5450 cal. BP). A first palynological investigation (core Cairon: Fig. 2C) was undertaken in relation to the excavation, but it failed to provide clear evidence of human impact on the vegetation during the settlement periods (Lespez et al., 2005).
<figure><img src="image_4.png" /><figcaption>**Fig. 2.** Study site. A: Archaeological setting within a 1-km radius of Cairon (IGN topographic database and *carte archéologique* database, 2022); B: Soils (Calvados soil reference system (2016), available on https://www.geoportail.gouv.fr/donnees/carte-des-sols (Technical manager of the study: P. Le Gouée)); C: Location of cores and borehole.</figcaption></figure>

## 3. Materials and methods

We have developed a high temporal resolution approach using multiple palaeobiological proxies to reconstruct landscape changes under human influence during the Neolithic, using a multi-source and multi-scalar approach. It provided the opportunity to compare off-site information with on-site archaeological and palaeoenvironmental data (within a radius of 1–3 km). Sedimentological analyses (magnetic susceptibility, X-ray fluorescence, carbon and nitrogen analyses) were primarily used to characterize changes in the depositional environment, in order to discuss the taphonomic processes that led to the fossilization of the bioindicators. Palynology was employed to reconstruct vegetation changes at both wetland and catchment scale. SedaDNA and NPPs complemented pollen, as they were used to reconstruct wetland dynamics and vegetation patterns, as well as local changes in grazing pressure. Charcoal analysis was conducted to reconstruct the local fire history in relation to vegetation changes and human practices.

### 3.1. Coring and chronology

A 3 m deep coring (Cairon, 2016) was extracted using a percussion drilling machine (Cobra TT), carried out near the previous cores and boreholes (Cairon and Cairon 2: Lespez et al., 2005, 2008). In 2019, a second core (Cairon, 2019) was taken 5 cm from Cairon (2016) (Fig. 2C) to perform sedaDNA analysis (See Fig. A in supplementary data 1). The chronology of the organic deposits of Cairon (2016) is based on 7 radiocarbon dates (AMS) (See Table A in supplementary data 1) performed at the Poznan Radiocarbon Laboratory and Beta Analytic. Radiocarbon ages were calibrated using OxCal v4.4 software (Bronk Ramsey, 2017) based on the IntCal20 atmospheric curve (Reimer et al., 2020). From the 7 dates obtained, one age-depth model (Fig. 4) was created using the Bayesian model in the Bacon package in R (Blaauw and Christen, 2011; version 2.5.7).

### 3.2. Magnetic susceptibility analysis

Volumetric magnetic susceptibility (MS) measurements (*10⁻⁶ CGS; SI units = CGS*4π) were performed continuously over the entire core at 1 cm intervals, in a double pass, using a Geotek multi-parameter bench fitted with a Bartington MS2E probe, at the Geosciences Laboratory of Rennes. Values were averaged and then multiplied by two to correct for the effect of the probe used. According to Dearing (1994), negative or zero MS values (diamagnetic material) are characteristic of quartz or carbonate rocks that do not contain metals, but also of sediments rich in water or organic matter. In the Caen Plain, we assumed that these values were related to carbonate detrital sediments resulting from the erosion of the limestone substratum and/or highly organic sediments. When MS values are strongly (ferrimagnetic or ferromagnetic) or moderately positive (paramagnetic or antiferromagnetic), the sediments contain materials charged with magnetic elements (Dearing, 1994); some of these elements are very common in sedimentary rocks and soils, such as iron (Fe), titanium (Ti) or manganese (Mn) (Rothwell and Croudace, 2015). In the Caen Plain, we assumed that these values were characteristic of detrital sediments from the loamy soils developed on the plateaus.

### 3.3. Carbon and nitrogen analyses

Total carbon (TC), total organic carbon (TOC) (after 3N HCl mineralization of the samples) and total nitrogen (TN) were measured on 39 samples (between 254 and 32.5 cm depth) at the Normandy Isotope Platform and at the Interdisciplinary Laboratory of Continental Environments (UMR 7360 CNRS - University of Lorraine). Values were used to calculate the percentage of inorganic carbon (TC-TOC = TIC) and the carbon-to-nitrogen ratio (TOC/TN = C/N). We hypothesized that high TOC indicated swampy or peat deposits, that high TIC was associated with carbonate detrital inputs from the limestone substratum of the Caen Plain, and that a decrease in the C/N ratio indicated soil mineralization due to a lowered water table and/or increased detrital inputs
(e.g. Ballut et al., 2008; López-Merino et al., 2010).

### 3.4. X-ray fluorescence analysis of major elements

The X-ray fluorescence analysis (XRF) was performed using an ITRAX scanner (Cox Analytical Systems) at the European Centre for Research and Education in Environmental Geosciences (CEREGE). A molybdenum tube was used as the x-ray source at 30 KV, 40 mA and 15 s count time. This rapid and nondestructive analytical technique was enabled to estimate the relative concentration of selected elements (mainly Ca, Sr, Ti, K, Mn, Fe, Zr) at the surface of the sediment core (0.5 cm spatial resolution between 244.5 and 5.5 cm depth), providing semi-quantitative values expressed in XRF intensity (Crouace and Rothwell,

2015; Longman et al., 2019). Titanium (Ti) is one of the main elements used to trace the evolution of terrigenous inputs (Rothwell and Crouace, 2015). Moreover, Ti is regarded as a good proxy for tracing aeolian inputs because it is a high field-strength element (Itambi et al., 2010). Therefore, in this study, variations in Ti have been used to trace the evolution of silty detrital inputs from the loamy soils developed on the plateaus, while the Ca/Ti ratio has been used to trace the evolution of carbonate detrital inputs. Following Schittek et al. (2015, 2016, 2021): first, under anoxic conditions and when humification is minimal, the proportion of manganese (Mn) as Mn²⁺ increases in the sediment; second, frequent oxic conditions caused by water table fluctuations lead to strong precipitation of Fe³⁺ oxides in the upper peat layer (Damman et al., 1992). In this study, we assumed that a high Mn/Fe ratio indicated

<figure><img src="image_5.png" /><figcaption>**Fig. 3.** Main NPPs identified from Cairon (2016). A *Sporormiella*-t. (HdV-113), B *Cercophora*-t. (HdV-112), C *Glomus* (HdV-207), D *Anthostomella* fuegiana (HdV-4), E Xylariaceae cf. *Helicogermslita* (EMA-55), F *Sordaria*-t. (HdV-55A), G HdV-11, H cf. *Persiciospora* (HdV-124), I *Clasterosporium* caricinum (HdV-126), J Coniochaetaceae-Xylariaceae, K HdV-713, L HdV-123, M HdV-201, N *Gelasinospora* (HdV-1), O *Melanconis alni* (EMA-3), P HdV-361, Q Scalariform perforation plates (HdV-114), R HdV-128A, S *Zygnema*, T HdV-224, U *Spirogyra*.</figcaption></figure>
anoxic conditions linked to a stable water table, while fluctuations and more frequent oxic conditions increased Fe proportions, lowering the Mn/Fe ratio (Schittek et al., 2015, 2016, 2021). Iron (Fe) was normalized to Ti, which is considered to be allochthonous and immobile in peat (Muller et al., 2008; Rothwell and Croudace, 2015), and a sharp increase in the Fe/Ti ratio was used as an indicator of more frequent water table fluctuations (Schittek et al., 2015, 2016, 2021).

### 3.5. Pollen and non-pollen palynomorphs analyses

Pollen and non-pollen palynomorphs (NPPs) analyses were performed on 36 sediment samples weighing 2–5 g (between 195.5 and 54.5 cm depth: average resolution of 4 cm intervals). The same standard protocol was applied for both pollen and NPPs analysis, adapted from Moore et al. (1991) and Fægri et al. (1989): 160 µm mesh sieving (using distilled water), hot 10 % KOH for 15 min (repeated until a clear supernatant is obtained), acetolysis, and densimetric separation for siltier samples using sodium polytungstate (SPT, density 1.9) to remove mineral particles. Pollen and spores were identified using pollen keys, pollen atlases (Beug, 2004; Fægri et al., 1989; Moore et al., 1991; Reille, 1992, 1995, 1998) and the pollen collection of the Physical Geography Laboratory (Thiais, France). NPPs were identified on the basis of the descriptions and photographic plates available in the literature (e.g. Barthelmes et al., 2006; Cugny, 2011; Cugny et al., 2010; Gelorini et al., 2011; Miola, 2012; Prager et al., 2006, 2012; Revelles and van Geel, 2016; van Geel, 1976, 1978, 1986; van Geel and Aptroot, 2006), and the non-pollen palynomorph Image Database (NPP-ID, Shumilovskikh et al., 2022). Most of the Coniochaetaceae and Xylariaceae spores described by Cugny (2011) and van Geel (1978) were grouped as Coniochaetaceae–Xylariaceae, except for Coniochaeta cf. ligniaria (HdV-172), Kretzschmaria deusta (HdV-44), and Xylariaceae cf. Heliogermslitia (EMA-55) (Barthelmes et al., 2006), which were more easily distinguished. Similarly, most Sordariaceae spores distinguished by Cugny (2011) were grouped together as undifferentiated Sordariaceae. Within the algal families, the main types were attributed to Mougeotia, Spirogyra and Zygnema spores (Fig. 3). Counts were performed until a minimum of 300 pollen grains were reached, excluding pteridophyte spores, indeterminate pollen grains, and NPPs. Of these, at least 100 grains were from outside the dominant taxon, and where possible, a minimum of 20 taxa were identified. NPPs were counted in parallel with the pollen counts on the same slide. Diagrams were constructed using

Psimpoll software 4.27 (Bennett, 2009) and divided into local pollen assemblage zones (LPAZ) and NPPs zones, defined independently using the CONISS (constrained cluster analysis by the method of incremental sum of squares) function (Grimm, 1987). To allow comparison with previous palynological studies, taxonomy followed the classical botanical classification (Cronquist, 1981).

### 3.6. Sedimentary ancient DNA analysis

Cairon 2019 was subsampled from a fresh cleaned face into 24 sterile tubes (between 153.5 and 56 cm depth: average resolution of 4 cm intervals). SedaDNA extraction was performed using the QIAGEN DNeasy Powersoil Kit following protocols first documented by Zimmermann et al. (2017), but with a smaller input volume of ~0.3 g and bead-beating step, as adapted by Alsos et al. (2020), and was undertaken in a dedicated ancient DNA laboratory at the University of Southampton. PCR amplification used primers of the P6 loop of trnL UAA intro of the plant chloroplast genome (Taberlet et al., 2007) and exact PCR procedure followed Alsos et al. (2021). Briefly, DNA amplifications were in 40 µL final volumes containing 4 µL of DNA sample, 1.6 U of AmpliTaq Gold® DNA Polymerase (Life Technologies, Carlsbad, CA, USA), 1X Gold Buffer, 2.5 mM MgCl2, 0.2 mM each dNTP, 0.2 µM each primer and 160 ng Bovine Serum Albumin. All PCR samples (DNA and controls) were randomly placed on PCR plates. Following the enzyme activation step (10 min at 95 °C), PCR mixtures underwent 45 cycles of 30 s at 95 °C, 30 s at 50 °C and 1 min at 72 °C, plus a final elongation step (7 min at 72 °C). Four negative extraction controls, two PCR negative controls and one positive control were carried out. Eight individually dual tagged PCR replicates were made for each sample to increase the chance of detecting taxa represented by low quantities of DNA, as well as to increase confidence in the taxa identified. We pooled PCR products and then cleaned the resulting pool following Rijal et al. (2021), using a MinElute PCR purification kit, and quantified them using Qubit (Qubit™ dsDNA HS Assay Kit, Thermofisher). Each amplicon pool was then converted into a DNA library at Tromsø using the Illumina TruSeq DNA PCR-Free protocol (Illumina Inc., CA, USA) with unique dual indexes, the only modification being that the magnetic bead clean-up steps were altered to retain short amplicons. Each library was sequenced on ~10 % of 2 × 150-cycle mid-output flow cells on the Illumina NextSeq platform at the Genomics Support Centre Tromsø (GSCT) at The Arctic University of Norway in Tromsø.

All next-generation sequence data were aligned, filtered and trimmed using the OBITools software package (Boyer et al., 2016) using the same criteria as Rijal et al. (2021) with custom R scripts (available at https://github.com/Y-Lammers/MergeAndFilter). Briefly, paired-end reads were aligned using SeqPrep (https://github.com/jstjohn/SeqPrep/releases, v1.2). Merged reads were demultiplexed according to their 8 bp unique primer tags and identical sequences were collapsed. Singleton sequences and those shorter than 10 bp were removed, and putative artifactual sequences were identified and removed from the dataset. Resulting barcodes were assigned to taxa using the ecotag program and four independent reference datasets. One reference contained arctic (Sønstebø et al., 2010) and boreal (Willerslev et al., 2014) vascular plants as well as bryophytes from the circumpolar region (Soininen et al., 2015) (ArcBorBryo, n = 2280 sequences of which 1053 are unique), one the NCBI nucleotide database (January 2021 release), and finally the PhyloNorway p6loop database (Alsos et al., 2022). The resulting identifications were merged and filtered, retaining barcode sequences if they were identified to >98 % in at least two reference sets and had at least 10 reads across the entire dataset. False positives relating to common PCR errors and food contaminants were removed based on ‘blacklists’ built up from previous research at The Arctic University Museum of Norway, https://github.com/Y-Lammers/Metabarcoding_Blacklists (Rijal et al., 2021). We also removed sequences that could only be identified above family level. For the last step of filtering, the frequency of PCR repeats in samples compared to

<figure><img src="image_6.png" /><figcaption>**Fig. 4.** Stratigraphic interpretation and age-depth model of Cairon (2016).</figcaption></figure>
negative controls was examined. Sequences were retained if they had a total number of PCR repeats in samples at least twice as high as that in negative controls.

We generated the animal sedaDNA data set from the same DNA ex-
tracts used above. We used the MamP007 primers (Giguet-Covex et al.,
2014) to amplify a 60–84-bp fragment of the mitochondrial 16S locus
(excluding primer length), following the protocol described in Garcé-
s-Pastor et al. (2022) that includes the use of a dual-blocking strategy
with a forward blocking primer sequence of GGAGCTTTAATTTAT-
TAATGCA AACAGTAGG-C3 and a reverse blocking primer with a
sequence of CCCAACCGAAATTTTTAATGCAGGTTTGGTGA-C3. PCR
product cleaning, pooling, library preparation, and sequencing followed
the protocols described above. Bioinformatic procedures followed
Brown et al. (2022) and briefly are as follows: First, direct and corre-
sponding reverse reads were assembled using illuminepairedend, and
sequences having a low alignment quality score (threshold set at 40)
were filtered out. The retained reads were assigned to relevant samples
using ngsfilter, keeping sequences matching 100 per cent with tags and
allowing a maximum of three mismatches with primers. Strictly iden-
tical sequences were then merged (dereplication) using obiuniq, keeping
information on their distribution among samples. All sequences with
only a single copy and/or shorter than 12 bp were filtered out using
obigrep. Obiclean was then used to identify amplification and sequencing
errors, using a threshold ratio of 5 per cent for reclassifying ‘internal’
sequences to their relative ‘head’ sequence (Brown et al., 2022). We
identified barcode sequences that had ≥95 % identity agreement to a
reference database generated with the MamP007 primers applied to the
EMBL (rl143) database using the ecoPCR tool (Ficetola et al., 2010).
Sequences assigned to non-native taxa were blasted to check for po-
tential wrong assignments (http://www.ncbi.nlm.nih.gov/blast/). We
next collapsed barcodes with the same taxonomic assignment by sum-
ming read counts on a per PCR replicate basis and retaining the barcode
(s) with the highest identity agreement. We then collapsed barcode se-
quences with the same taxonomic assignment, as above, resulting in
only 4 animal taxa.

3.7. *Macro-charcoal analysis*

101 contiguous samples of 2 cm³ were retrieved at 2 cm intervals (between 245 and 32.5 cm depth), wet sieved with 160 µm mesh, dried in an oven at 50 °C for 24 h, and transferred to petri dishes. The number of charcoal particles was estimated under a binocular microscope at a 40× magnification to reconstruct local fire history (Blackford, 2000; Carcaillet et al., 2001; Conedera et al., 2009; Tinner et al., 1998). The decomposition approach introduced by Long et al. (1998) was applied to process the results using CharAnalysis 0.9 software (Higuera, 2009). The charcoal record was expressed as charcoal accumulation rate (CHAR) (Higuera et al., 2010), which was interpolated to a constant temporal resolution (CHARint) and filtered using the moving median smoothing method with an 800-year temporal window to model the background component (CHARback). CHARback was subtracted from the raw series (CHARint) to obtain the residual high-frequency peaks (CHARpeak). These peaks were decomposed into 2 sub-populations using a Gaussian mixture model (introduced by Gavin et al., 2006) and a locally defined threshold to separate noise (CHARnoise) from local fire events (Fire peak). Fire regimes were mainly defined according to peak magnitude (the number of charcoal particles associated with one fire event) and fire frequency within the 800-years temporal window. The signal-to-noise index (SNI) and the goodness-of-fit (GOF) metrics were used to assess whether charcoal records were appropriate for peak analysis. The choice of parameters in CharAnalysis was guided by maximizing SNI and GOF values (values greater than or equal to 3 and 0.10 respectively) (Blarquez et al., 2013; Higuera, 2009; Kelly et al., 2011).

4. Results and interpretation

4.1. *Chronology and description and interpretation of the sedimentary units*

The age-depth model (Fig. 4) which contained no stratigraphic out-
liers, was used to calculate temporal resolution of the proxy analyses
(See Table B in supplementary data 1). Sedimentary units (SU) were
described according to their depth and, where possible, the time frame
derived from the age-depth model was indicated for each SU (See Fig. 5
and Table C and Fig. B in supplementary data 1).

SU1 coarse gravel was overlain by Holocene fluvial carbonated silty
sands (SU2). SU3 consisted of carbonated gravel and detrital floodplain
deposits associated with a channelized river (Lespez et al., 2008, 2015).
SU4 was composed of organo-calcareous silt sedimentation associated
with a marshy floodplain. SU5 corresponded to peatland deposits, and
was overlain by overbank carbonated sands (SU6) and by silts from the
erosion of the silty-soils developed on the plateau (SU7). In the Mue
Valley—and more broadly across the Caen Plain—overbank silts
increased mainly between the Bronze and Iron Ages (Lespez, 2012a;
Lespez et al., 2008, 2015), although this detrital sedimentation was not
directly dated at Cairon.

4.2. *Palaeobiological analyses*

Results obtained from Cairon (2016) were described according to
ages estimated using the age-depth model.

4.2.1. *Pollen analysis results*

The number of pollen grains counted and identified per sample ranged from 305 to 584, representing a taxonomic diversity of 18–28 taxa (Fig. 6). Pollen concentration was high, with an average of 60,625 grains/g. At the base of the sequence (LPAZ1 to LPAZ2), only 3 samples were analyzed, with an average temporal resolution of 230 years. Here, pollen concentrations were the lowest (7000 to 16,000 grains/g) (Fig. 6). In the upper highly organic deposits (LPAZ3 to LPAZ6), pollen concentrations were higher (25,000 to 156,000 grains/g), and 33 samples were analyzed. These provided very high-temporal resolution results, with an estimated average of 61 years (min. 17, max. 96), offering a detailed picture of the evolution of vegetation between the end of the Mesolithic and the transition to the Late Neolithic (Fig. 7 and Table D in supplementary data 1).

4.2.2. *Non-pollen palynomorphs analysis results*

The total number of NPPs types counted and identified per sample varied between 90 and 1,263, with taxonomic diversity ranging from 4 to 28 taxa (Fig. 8). Assemblages were dominated by fungal spores and other types, followed by algal spores, with mean concentrations of 40,200 spores/g and 630 spores/g, respectively. At the base of the sequence (NPPsZ1 and NPPsZ2), concentrations of fungal spores and other types were comparatively low (700–6900 spores/g), while concentrations of algal spores were within the average (120–550 spores/g). The NPPsZ3 showed a strong increase in concentrations of fungal spores and other types (up to more than 80,000 spores/g), while algal spores concentrations decreased to zero in NPPsZ3b. In the NPPsZ4, concentrations of fungal spores and other types remained high (30,000–70,000 spores/g), but those of algae increased (3000 spores/g). From NPPsZ5 onwards, concentrations of fungal spores and other types gradually decreased (to less than 10,000 spores/g), while concentrations of algal spores remained stable. In the NPPsZ5, concentrations of both fungal spores and other types (300–1500 spores/g) and algal spores (150 spores/g) decreased. From NPPsZ4 onwards, the woody micro-remain Scalariform perforation plates (HdV-114) appeared and gradually increased in concentration, becoming the dominant taxon in NPPsZ6. Meanwhile, the concentrations and diversity of other NPPs declined sharply (Fig. 8 and Table E in the supplementary data 1).
<figure><img src="image_7.png" /><figcaption>**Fig. 5.** Sedimentological and geochemical analysis from Cairon (2016). The horizontal lines indicate the mean value of each indicator along the x-axis, serving as a baseline to assess how the curves fluctuate relative to the mean. Unreliable measurements (kilo counts per second <16,000) correspond to core intervals characterized by irregular or unconsolidated sediment surfaces, leading to reduced X-ray signal intensity and decreased data reliability. From the macroscopic description 7 main sedimentary units (SU) were distinguished and sometimes subdivided into sub-units (a, b, c, etc) according to sedimentological and geochemical results (See Table C in supplementary data 1).</figcaption></figure>

### 4.3. Sedimentary ancient DNA analysis results

The sedaDNA results obtained from Cairon (2019) were presented according to depths associated with ages interpolated from those obtained at the same depth on Cairon 2016. SedaDNA metabarcoding using the p6 loop produced 2,114,282 reads after pre-identification filtering (see methods) from 21 of the 24 samples collected, with an average of 88,113 reads per sample. In total, a list of 33 taxa was compiled representing woodland, meadow, wetland and aquatic habitats (Fig. 9). The data could be regarded as of high quality as none of the identified taxa were found in any of the three negative control samples. The assemblages reflected a closed canopy wooded environment dominated by alder (*Alnus*), willow (Salicaceae), birch/hazel (Betulaceae/Coryloid) and poplar (*Populus*), as well as wetlands and aquatic habitats (Dardillac et al., 2019). We observed the beginning of alder growth (*Alnus*) at 5160 BCE (7110 cal. BP). There were no cereals or domesticated plants, however, there were taxa associated with nutrient rich meadow or wet pasture (*Urtica* and *Galium*). The aquatics taxa indicated both running water (bur-reed/*Sparganium*) and still water (duckweed/*Lemna*) (Dardillac et al., 2019).

The 16S data was relatively sparse, with a total 120,182 reads obtained from all samples after pre-identification filtering, but does have

cattle (*Bos*) at 5180 BCE (7130 cal. BP), sheep (*Ovis*) at 4350 BCE (6300 cal. BP), and pig/wild boar (*Sus*) at 5220 BCE (7170 cal. BP), with two different sequences. The *Bos* is almost certain to be domesticated cattle (*Bos taurus*) as it was distinct enough for metabarcoding using 16S from aurochs (*Bos primigenius*) (Hudson et al., 2023). The *Ovis* is also domesticated sheep as there is no native species, and the *Sus* is most likely to be wild boar given the date, although this cannot be certain using metabarcoding data.

### 4.4. Main phases of vegetation dynamics inferred from pollen and sedaDNA

There was partial overlap between the taxa identified by sedaDNA and those found in the pollen spectra. We assumed that taxa present in the pollen spectra but absent from the sedaDNA record were not growing near the coring point, but outside the wetland. Some taxa appeared in the sedaDNA record but were absent—or very scarce—in the pollen spectra. This was likely due to the low pollen productivity of insect-pollinated species or identification challenges. Overall, sedaDNA complemented the pollen record and, in some cases, provided greater taxonomic resolution, enabling a more detailed reconstruction of vegetation formations and dynamics at both the wetland and valley
<figure><img src="image_8.png" /><figcaption>**Fig. 6.** Pollen concentration diagram from Cairon (2016) (The scale for concentrations values differs across taxa). Concentration of selected taxa representative of the different ecological associations defined in the caption of Fig. 7. Concentrations were calculated using the volumetric method (Cour, 1974). Total pollen concentration: based on the sum of AP, NAP, pteridophyte spores, and indeterminate pollen grain counts. A. Number of pollen taxa identified per sample; B. Basic pollen sum: (AP + NAP) minus *Corylus* counts; C. Main pollen sum: AP + NAP counts.</figcaption></figure>

scales.

* **6135–5105 BCE; 8085–7055 cal. BP (LPAZs 1,2,3 and 4; Figs. 6 and 10).** The sedaDNA indicated that hazel, birch, willow, viburnum, dogwood, elder, privet, and trees of the apple family may have been part of the local vegetation. Wetland vegetation was dominated by hygrophilous and aquatic formations, mainly represented by Cyperaceae, *Filipendula*, and *Sparganium-t.*, as well as common reed and other heliophilous taxa (e.g., *Asteroideae*, *Ranunculaceae*, and *Scrophulariaceae*) included in both the pollen and sedaDNA records. Upland vegetation was composed of both hazel and mixed oak forest, with mainly oak and elm, followed by lime, ivy, maple, mistletoe, and viburnum. Close to the wetland, small clearings were related to grassland and ruderal communities (*Poaceae*, *Cichorioideae*, *Apiaceae*, *Brassicaceae*, *Artemisia*, *Chenopodiaceae*, and *Polygonum aviculare-t.*). The pollen of *Viscum* in LPAZ 3 not only indicated the local presence of the obligate hemi-parasitic mistletoe plant but also in situ host tree cover, which was probably willow, ash, or poplar.

* **5105–4850 BCE; 7055–6800 cal. BP (LPAZ 5a; Figs. 6 and 10).** Both pollen and sedaDNA showed a gradual rise in riparian vegetation, mainly due to the local growth of alder. Consequently, shrubs detected by sedaDNA (viburnum, elder, privet, and apple family trees) were absent from the pollen spectra. Upland forest became denser, mainly from increased hazel, elm, oak, and lime.

* **4850–4030 BCE; 6800–5980 cal. BP (LPAZs 5b and 5c; Figs. 6 and 10).** A gradual rise in *Poaceae* (possibly reeds according to SedaDNA results) and an increase in grassland and ruderal taxa after 4520 BCE (6470 cal. BP)—mainly *Asteroideae*, *Apiaceae*, *Brassicaceae*, *Plantago lanceolata-t.*, and *Artemisia*—reflected the diversification of herbaceous formations in the wetland and the local spread of grazed meadows. From 4150 BCE onward, the first cereal pollen indicated the presence of crops near the wetland. The continued presence of *Viscum* pollen again suggested tree cover very close to the core site. In upland forest, hazel, elm, and ivy gradually decreased, while oak remained stable.

* **4030–3475 BCE; 5980–5425 cal. BP (LPAZ 6; Figs. 6 and 10).** Riparian forest, dominated by alder along with birch and willow, expanded across the valley bottom. Between 3780 and 3475 BCE (5730–5425 cal. BP), the presence of cereal pollen indicated crop cultivation near the wetland. Upland forest declined, with a sharp decrease in elm, and after 3800 BCE, ferns colonized the forest edges.

### 4.5. *Ecological successions and wetland dynamics inferred from pollen, NPPs and sedimentary indicators*

An ordination by principal component analysis (PCA) was performed on 36 samples on which bioindicators were analyzed (See Texte A and Fig. C in supplementary data 1), using a linear interpretation method to explore the fossil dataset's response to environmental gradients. Based on the PCA results, sedimentary, pollen, and NPPs indicators were selected (Fig. 10) to reconstruct wetland ecological successions in relation to changes in the water table and detrital inputs.

* **Higher water table (6135–5490 BCE; 8085–7440 cal. BP; Phase A Fig. 10).** Hygrophilous, followed by aquatic, formations spread across the valley bottom, replacing ferns and dryland trees, while both carbonate detrital and organic fractions increased, reflecting hydrological abundance in the marshy floodplain.

* **Water table stabilization (5490–4850 BCE; 7440–6800 cal. BP; Phase B Fig. 10).** Lentic conditions were associated with a decrease in the coarse carbonate detrital fraction and an increase in the organic fraction. In the peatland, aquatic plant proliferation and high silty lateral inputs corresponded to average algal spore concentrations and a sharp rise in both open and peaty wetland indicators. Subsequently, ferns and hygrophilous taxa (LPAZ 4a), meadow taxa (LPAZ 4b), and dryland and riparian forest taxa (LPAZ 5a) spread along drier edges, while open wetland indicators decreased. Silty detrital inputs sharply decreased, algal spores were no longer recorded, and peaty wetland indicators sharply increased.
<figure><img src="image_9.png" /></figure>

<figure><img src="image_10.png" /></figure>

(caption on next page)
**Fig. 7.** Pollen percentage diagram from Cairon (2016) (Curve exaggeration x10). Taxa are classified according to different ecological associations. Pine, hazel, and taxa associated with mixed oak forest originate mainly from woody vegetation on dry land and may come from both local and more regional forest communities. *Corylus* has been excluded from the mixed oak forest association because, although this taxon naturally grows in the understorey of forest communities, it is often associated with pioneer, heliophilous shrub communities (Kalis et al., 2003). It is also characteristic of forest hollows, based on the assumption that hazel pollen production is favored by increased light availability (Begeot, 1998; Feeser and Dörfler, 2014). Herbaceous plant taxa are mainly derived from local vegetation, within a radius of around 1 km—or even less in the case of cereal pollen (Diot, 1992; Heim, 1970). Taxa associated with riparian forests, hygrophilous, and aquatic species are assumed to originate from local alluvial wetland vegetation. Anthropogenic pollen indicators include cereal pollen and the main taxa considered ruderal indicators in the literature (Behre, 1981). Other herbaceous taxa are grouped in the ubiquitous association and may derive from plants in dry grasslands, wet meadows, and grazed or cultivated areas. Pteridophytes are representative of ferns growing in wetland vegetation, as well as in forest edges, undergrowth, or abandoned land. The AP/NAP (arboreal pollen/non-arboreal pollen) ratio and *Corylus* percentages were calculated based on the main pollen sum (AP + NAP counts); other individual pollen percentages were calculated using a basic pollen sum that excludes the dominant taxon ((AP + NAP) minus *Corylus* counts); percentages of pteridophyte spores were calculated from the main pollen sum plus pteridophyte spores and indeterminate pollen grains.

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

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

**Fig. 8.** NPPs concentration diagram from Cairon (2016) (The scale for concentrations values differs across taxa). According to the literature, NPPs are classified in 7 main groups as: spores of coprophilous/saprophytic fungi, spores of lignicolous fungi, spores of parasitic fungi associated with sedges, fungal spores and other types, algae (algae, hyaline NPPs often associated with algae and cyanobacteria), animal micro-remains (and protozoa) and woody micro-remains. Spores of coprophilous/ saprophytic fungi include taxa regularly considered as coprophilous, but which also have affinities with soils, wood and decaying plants (Ellis and Ellis, 1997; Perrotti and van Asperen, 2019). Lignicolous fungi are more specifically related to fungi associated with living or dead trees. Fungal spores and other types are NPPs that do not fit into other categories. Originating from fungi or unidentified organisms, their ecological meaning remains poorly defined in the literature, especially in fluvial environments. Concentrations were calculated using the volumetric method (Cour, 1974); A. Number of taxa (excluding algae) identified per sample; B. Number of algae identified per sample.

* Higher water table (4850—4520 BCE; 6800—6470 cal. BP; Phase C Fig. 10). Hygrophilous and aquatic taxa spread mainly at the expense of riparian forest taxa, and concentrations of open wetland indicators increased. Silty detrital inputs remained low, algal spores were not recorded, and peaty wetland indicators remained high.

* Rise in the silty inputs (4520—3475 BCE; 6470—5425 cal. BP; Phases D and E Fig. 10). Dryland forest taxa decreased alongside rising silty detrital inputs, while HdV-361 and algal spore concentrations were relatively high and peaty wetland indicators decreased. During phase D, ubiquitous meadow taxa spread across the peatland
<figure><img src="image_13.png" /><figcaption>Fig. 9. Plant and animal sedaDNA as several PCR replicates 1–8 and local pollen zones (LPAZ) from Cairon (2019). (Sterile samples: samples that did not provide plant sedaDNA).</figcaption></figure>

<figure><img src="image_14.png" /><figcaption>Fig. 10. Synthetic diagram of selected pollen, NPPs and sedimentary indicators of Cairon (2016) (The scale for NPPs concentrations values differs across taxa). Phases A, C and D: regressive plant succession: increase in hygrophilous and aquatic or meadow taxa. Phases B and E: progressive plant succession: increase in dryland and/or riparian forest taxa. According to PCA (See Texte A and Fig. C in supplementary data 1): Peaty wetland indicators = fungal spores (HdV-713, Coniochaetaceae-Xylariaceae, *Cercophora*-t. (HdV-112), *Sordaria*-t. (HdV-55A), *Gelasinospora* (HdV-1), HdV-182 and cf. *Brachysporium* (HdV-1036), HdV-4, cf. *Persiciospora* HdV-124, (*Glomus* (HdV-207), *Clasterosporium caricinum* (HdV-126)) clustered on axis 1 and axis 2; Mineralized wetland indicators = algal spores (*Zygnema*, HdV-224, *Spirogyra* et HdV-128A) and HdV-361 clustered on axis 1, and Scalariform perforation plates (HdV-114) opposed to a large diversity of fungal and algal spores on axis 2; Open wetland indicators = fungal spores (cf. *Helicogermslita* (EMA-55), HdV-11, HdV-201, HdV-144, HdV-16 and HdV123) clustered on axis 3 and axis 4.</figcaption></figure>
at the expense of riparian forest taxa, and open wetland indicators increased. During phase E, there was a sharp increase in riparian forest taxa and a decrease in open wetland indicators. Scalariform perforation plates (HdV-114) appeared, and from NPP Z6 onward, the concentration of this taxon increased sharply, while all other NPPs and silty detrital inputs declined.

## 4.6. Macro-charcoal analysis results

According to the GOF and SNI values, charcoal record was considered appropriate for peak analysis only for the organic sequence between 53 and 205 cm depth. Therefore, only the results from the 74 samples from this portion with a well-defined chronology were presented in Fig. 11 (For detailed description of the results see Table F in supplementary data 1). At the base of the sequence, the CHAR was very

low and no fire peaks were recorded. Then, the results highlighted two phases of increase in the CHAR and in the CHAR<sub>back</sub>, both associated with fire peaks with medium (CHARZ2) to higher magnitude (CHARZ4). Between these two phases, the CHAR and the CHAR<sub>back</sub> sharply decreased, while a single fire peak of low magnitude was recorded (CHARZ3). Towards the top of the sequence, no fire peaks were recorded, but the CHAR<sub>back</sub> remained comparatively high.

## 5. Discussion

### 5.1. Using NPPs to reconstruct wetland change under early human influence

The site of Cairon is different from the majority of those previously examined in the region, which mainly correspond to peat bogs or lakes.

<figure><img src="image_15.png" /><figcaption>**Fig. 11.** Charcoal analysis from Cairon (2016) (Charcoal zones visually defined). Charcoal accumulation rate (CHAR): based on sedimentation rate estimated by the established age-depth model and interpolated to a 16 years temporal resolution (highest temporal resolution interval); Background component (CHAR<sub>back</sub>): part of the charcoal record mainly associated with secondary charcoal and/or regional fires (Clark et al., 1998; Conedera et al., 2009; Whitlock and Larsen, 2001); Fire peak: charcoals related to the occurrence of local fire events (within a radius of 1–3 km); Peak magnitude: number of charcoals associated with one fire event; Fire frequency: number of fire peaks in the 800 years temporal window.</figcaption></figure>
In addition, all NPPs have been considered in our study, and not solely the coprophiles or the easiest to identify. Therefore, our results provide some very interesting additions to the existing comparative literature.

### 5.1.1. NNP s and wetland vegetation changes

The NPP record reflected changes in wetland vegetation and depositional environment, supporting a local origin for most palynomorphs (e.g. Cugny et al., 2010; Mulder and Janssen, 1999; van Geel, 2001; van Geel et al., 2003; van Geel and Aptroot, 2006). From 5490 BCE (7440 cal. BP) onwards, in a terrestrial peaty wetland, rise in some fungi could be related to the local development of dense fen vegetation formations, while sedaDNA attested to the presence of Cyperaceae and other helophytic plants, as well as riparian trees. Scalariform perforation plates (HdV-114) were related to forested wetland, and were only found at the top of the sequence (Fig. 10), where both pollen and high concentrations of Melanconis alni (EMA-3) (Prager et al., 2006) were indicative of the local development of a dense alder forest. In opposition, high concentrations of cf. Helicogermslita (EMA-55), HdV-201, HdV-11, HdV-123, HdV-144 and HdV-16 were related to open wetland. Our results confirmed previous palaeoecological investigations conducted in different environments, since the Scalariform perforation plates (HdV-114) occur in vessels from a variety of trees (Pals et al., 1980), and have previously been related to the local presence of riparian trees, in particular alder (Barthelmes et al., 2012; Prager et al., 2006, 2012; Revelles et al., 2016; Revelles and van Geel, 2016). Similarly, cf. Heliogermslita (EMA-55) has been related to open sedge fen (Barthelmes et al., 2006; Prager et al., 2012), HdV-201 to the presence of helophytes in temporary desiccating bottoms of pools (Kuhry, 1997), HdV-11 has been found in organic connection with a hyperaceous epidermis (van Geel, 1978), and HdV-123 in surface peat sample in common reed (Prager et al., 2012). However, spores of Clasterosporium caricinum (HdV-126) and Anthostomella fuegiana (HdV-4) are both from parasitic fungi of various Cyperaceae species (Ellis and Ellis, 1997; Pals et al., 1980; van Geel, 1978; van Geel et al., 1983; van Geel and Aptroot, 2006), some of which may occur in open and forested wetlands (Barthelmes et al., 2012). According to sedaDNA Cyperaceae were part of the local vegetation both during open and forested wetland phases at Cairon. Therefore, Clasterosporium caricinum (HdV-126) and Anthostomella fuegiana (HdV-4) were poor indicators of open and forested wetland phases.

### 5.1.2. NNP s and degree of peat mineralization and decomposition

In our study, high concentrations of HdV-713, Coniochaetaceae-Xylariaceae, and *Cercophora*-t. (HdV-112) were related to mainly authigenic deposits accumulating while forest protected the soils of the hillsides and the plateau from erosion (Fig. 10). In contrast, HdV-361 and some algal spores (Zygnema, HdV-224, Spirogyra and HdV-128A) were mainly related to partly mineralized wetland. From 4520 BCE (6470 cal. BP) onwards, the HdV-361 record and rise in concentrations of algal spores indicated peat mineralization due to detrital inputs (Fig. 10) in relation to clearings. These findings are in agreement with other palaeoecological investigations, since HdV-361 has already been related to erosive episodes due to intensive human activity in the catchment (Out, 2010; Revelles et al., 2016; Revelles and van Geel, 2016; van Der Wiel, 1982) and positively correlated with herbaceous pollen indicative of open environments (Enevold et al., 2019). As suggested by van Geel et al. (1981) and Revelles and van Geel (2016), eutrophication and higher water turbidity in ponds due to terrigenous inputs could also promote the local growth of algae (particularly green algae).

High concentrations of a diversified set of fungal and algal spores were related to peaty wetland (Fig. 10). In contrast, high concentrations of Scalariform perforation plates (HdV-114) were related to partly mineralized wetland. These micro-remains implied a high degree of decomposition of the wood and have already been related to highly degraded peat Barthelmes et al. (2006), 2012; On Cairon 2016, from

4030 BCE (5980 cal. BP) onwards, the increase in concentrations of Scalariform perforation plates (HdV-114) in parallel with the decline in most fungal and algal spores indicated a higher degree of peat decomposition. Thus, changes in the NPPs assemblage provided additional information by showing a gradual degradation of the peatland due to detrital inputs resulting from continuous human activity in the catchment area since the Middle Neolithic period.

### 5.1.3. NNP s and changes in grazing pressure

In our study, *Sporormiella*-t. (HdV-113) was the only coprophilous/saprophytic taxon always related to anthropized wetland and appeared as a robust local indicator of grazing (Fig. 12). Among the main coprophilous/saprophytic taxa determined, *Sordaria*-t. (HdV-55A) was often considered as one of the most reliable indicators of the presence of herbivore dung (Cugny et al., 2010; Perrotti and van Asperen, 2019; van Geel, 2001; van Geel and Aptroot, 2006). But *Cercophora*-t. (HdV-112) and Coniochaetaceae-Xylariaceae are from fungi growing primary on soils, wood and decaying plants, and secondary on dung (Ellis and Ellis, 1997; Perrotti and van Asperen, 2019). These three taxa were recorded with fungal spores related to wood and decaying plants (HdV-713, cf. *Persiciospora* (HdV-124) (Bakker and van Smeerdijk, 1982), *Glomus* (HdV-207) (van Geel et al., 1989, 2003) and *Gelasinospora* (HdV-1) (van Geel, 1978)), as well as the local presence of Cyperaceae (*Clasterosporium caricinum* (HdV-126) and *Anthostomella fuegiana* (HdV-4)) (Ellis and Ellis, 1997; Pals et al., 1980; van Geel, 1978; van Geel et al., 1983; van Geel and Aptroot, 2006). Therefore, from 5490 BCE onwards (7440 cal. BP), high concentrations of fungal spores were mainly related to the large amount of various dead plant material lying at the drier peatland surface at Cairon, confirming observations of previous palaeoecological investigations (e.g. Bakker and van Smeerdijk, 1982; Kuhry, 1985; Shumilovskikh et al., 2015; van Geel, 1978). Due to their ambiguous ecological affinity, most of the spores of coprophilous/saprophytic fungi were poor indicators of changes in grazing pressure. This agreed with the results obtained in modern dung samples from highland cattle grazing in the Jura Mountain, where *Cercophora* and *Sordaria* were always associated with *Sporormiella* (Jouffroy-Bapicot et al., 2013) considered to be mainly coprophilous (Cugny et al., 2010; Davis and Shafer, 2006; Perrotti and van Asperen, 2019; van Geel et al., 2003).

## 5.2. The impact of early human activity on landscape

### 5.2.1. Fire and mesolithic forest management

In the Caen Plain, previous data show that during the Early Holocene (8100–6900 BCE; 10,050–8850 cal. BP), the regional landscape was dominated by woodland interspersed with semi-open habitats where trees and shrubs were abundant (Clet-Pellerin et al., 1977, 1981; Clet-Pellerin and Verron, 2004; Elhai, 1963; Huault, 1972; Lespez, 2012b; Lespez et al., 2005, 2010). Many of these were fruit-trees, including hazel, which appears to have played a major role in the diet of Mesolithic populations (Ghesquière, 2015). The development of mixed oak forest toward the end of the Early Holocene (from 6900 BCE; 8850 cal. BP) corresponded to a decline in the woody species used for gathering wild fruit, including hazel. From this period onwards, Late Mesolithic populations likely had to adapt their subsistence practices. At the same time, archaeological remains became very scarce. They consisted mainly of lithic material collected over relatively wide areas (Artur et al., 2008; Ghesquière, 2011; Ghesquière and Chancерel, 2016), suggesting the presence of semi-permanent settlements. Trapezoidal armatures of Mediterranean influence were also discovered, and La Hoguette and Limbourg pottery has been identified at three sites (Billard and Alix, 2014; Caillaud and Lagnel, 1972; Le Saint-Allain et al., 2021). This material has sparked discussions about the early introduction of agricultural products, potentially resulting from Cardial influence or from trade networks with Early and Middle Linear Pottery culture settlers (e. g. Billard and Legris, 2010; Constantin et al., 2010; Gomart et al., 2015; Marcigny et al., 2007). In northern France, northwestern Germany, and
<figure><img src="image_16.png" /><figcaption>**Fig. 12.** Synthetic diagram of selected palaeobiological and sedimentary indicators from Cairon (2016) and correlation with the local archaeological data and palaeoclimatic data. Phase 1: short-lived clearings = fire peaks of low magnitude contemporaneous with slight changes in woodland composition. Phases 3 and 4: meadows spreading = fire peaks of greater magnitude contemporaneous with decrease in riparian or dryland forest taxa. Phase 5: clearings on a larger scale = high background component contemporaneous with decrease dryland forest taxa. According to PCA (See Texts A and Fig. C in supplementary data 1): local grazing indicator = *Sporormiella*-t. (HdV-113) the only spores of coprophilous/saprophytic fungi always related to anthropized wetland on axis 2 and axis 4. Other coprophilous/saprophytic fungi = *Podospora*-t. (HdV-368), *Apisordaria* (HdV-169), *Chaetomium* (HdV-7A), *Trichodelitschia* (HdV-546), undifferentiated *Sordariae*, *Sordaria*-t. (HdV-55B), *Sordaria*-t. (HdV-55A), *Cercophora*-t. (HdV-112), *Coniochaeta* cf. *Ligniaria* (HdV-172) and *Coniochaetaceae*-Xylariaceae. Carpological, anthracological and dendrological data from the Middle Neolithic domestic settlement (Ghesquiere and Marcigny, 2011): Mixed oak forest with areas of dense forest = mainly charcoals of oak, oak/chestnut, beech, elm, hornbeam and wood ring curvature analysis indicates that some oak charcoals come from areas of dense forest unaffected by agricultural clearance; Forest edges, shrublands and abandoned land = mainly charcoals and/or seeds of hazel, Pomoïdeae (common hawthorn (*Crataegus monogyna*), wild apple (*Malus sylvestris*), and probably the genus *Pyrus* (pear-apple)), *Prunus* (potentially wild cherry (*Prunus avium*) or blackthorn (*Prunus spinosa*)), elderberry and pine; Riparian Forest = mainly charcoals of birch, alder, willow and poplar. Palaeoclimatic data: A. Rise in the water table in the Vey valley (this study); B. Flood events probability in Great Britain (Macklin et al., 2010); C. Period of increased storminess in the North Atlantic (Goslin et al., 2018).</figcaption></figure>

Switzerland, discussions were also stimulated by the identification of cereal pollen grains—sometimes associated with a slight increase in the proportion of ruderal taxa—in off-site pollen sequences dated to the Mesolithic (Behre, 2007; Leroyer et al., 2004; Richard, 2000; Ruffaldi, 1999; Tinner et al., 2007; Visset et al., 2002, 2004).

In our results, the end of the Late Mesolithic showed the first increase in local fire frequency (CHARZ2 Fig. 11 and Phase 1 Fig. 12). The decline in woody pollen curves was mainly associated with an increase in the percentage of hygrophilous and aquatic pollen (Fig. 10). Between 5900 and 5490 BCE (7850 and 7440 cal. BP), pollen concentrations were very low (Fig. 6), highlighting that carbonate sedimentation is less favorable for pollen preservation (Moore et al., 1991). As a result, the pollen analysis sampling interval was wide (3 samples over 600 years) and did not allow us to estimate vegetation dynamics, which could be very rapid in the case of a slight hollow in the forest (Bégeot, 1998). From 5490 BCE (7440 cal. BP) onwards, pollen was well preserved with higher concentrations, and the sampling time resolution was high (around 60

years). This allowed the detection of rapid changes in vegetation. Between 5490 and 5195 BCE (7440–7145 cal. BP), the pollen curves for dryland trees were irregular, with peaks in *Corylus* and *Quercus*, while *Viburnum*, *Viscum*, and *Acer* appeared only sporadically or in low percentages (Fig. 7), indicating their presence near or at the coring point. Areas cleared by fire could have been quickly colonized by shrubs such as viburnum. Following these pioneer shrubs, hazel, oak, and maple were able to colonize the former clearings. The peaks of *Corylus* and *Quercus* may also have corresponded to minor clearings, assuming that pollen production of these taxa was favored by increased light availability (Bégeot, 1998; Feeser and Dörfler, 2014). In any case, these fire peaks of low magnitude were only associated with the development of small forest hollows at the bottom of valley or on lower hillsides.

Furthermore, on Cairon 2016, ruderal taxa (*Artemisia*, *Chenopodia-ceae*, and *Polygonum aviculare*-t.) were recorded in low cumulative percentages (0.5–2 %) (Fig. 12). Whilst they have been associated with grazed meadows (Behre, 1981), they could also grow in forest hollows or
open areas near the wetland (Behre, 2007). In these temporary open or semi-open areas, more diversified herbaceous vegetation could develop, associated with the presence of wild fruit trees (pollen and/or DNA from viburnum, elder, apple, hazel) (Figs. 7 and 9), whose fruits (at least when cooked) could be consumed. *Sporormiella*-t. (HdV-113) was recorded discontinuously with low concentrations (up to 250 spores/g). SedaDNA indicated the presence of pig/wild boar, suggesting that wild animals were regularly present in the wetland (Phase 1 Fig. 12). Clearings may have promoted the secondary growth of fruit trees and shrubs and attracted wild mammals for browsing. The wet meadow at the bottom of the valley may have also attracted wild animals for browsing or drinking. Thus, the environment during this period appeared particularly suitable for hunter-gatherer populations, who may have benefited from products that were naturally present or favored by human practices.

Our results showed that this period was characterized by a rise in the water table, which overlapped a time interval marked by episodes of significant river activity in Great Britain (Macklin et al., 2010). It was also synchronous with a period of increased storminess in the North Atlantic (Phase 1 Fig. 12), which may have been related to a persistent positive NAO-like situation (Goslin et al., 2018), resulting in higher moisture in northwestern Europe. This context suggests that the fires at Cairon were human-induced. Instead of supporting the hypotheses of the early introduction of agricultural products following contact with earlier Neolithic populations (Leroyer et al., 2004; Richard, 2000; Ruffaldi, 1999; Tinner et al., 2007; Visset et al., 2002, 2004), our data point to a transformation of the environment by local populations using fire, despite limited archaeological evidence for contemporary human settlement. According to some authors, in a denser forest context, fires could have been used to create or maintain open and semi-shaded areas to ensure the development of forest resources that Mesolithic people had consumed and that were favorable for hunting (e.g. Bishop et al., 2015; Bos and Urz, 2003; Dietze et al., 2018; Innes et al., 2013; Mason, 2000; Mithen, 2019; Moore, 2000; Sevink et al., 2018). This is consistent with the common idea in northwestern Europe about practices involving 'reasoned management' of forest resources, in opposition to the idea of an opportunistic mode of subsistence involving a high degree of population mobility.

### 5.2.2. An initial neolithization characterized by limited environmental impact

From the Early Neolithic to the beginning of Middle Neolithic I (5195–4400 BCE; 7145–6350 cal. BP; Phase 2 Fig. 12), the Cairon record shows scarce anthropogenic indicators, a significant decrease in fire frequency, and a decrease in the diversity of dryland forest trees that were densifying. The data suggest that domesticated animals were present at least periodically in the valley bottom. Indeed, *Plantago lanceolata*-t. was first recorded around 5000 BCE (6950 cal. BP), and *Sporormiella*-t. (HdV-113) was recorded discontinuously in relatively low concentrations (up to 300 spores/g). Additionally, cattle DNA—most likely from domesticated cattle—was found in one sample. Mammal sedaDNA concentrations are typically far lower than those of plant sedaDNA, even in lacustrine sequences (Brown et al., 2022), so this detection could underrepresent the true level of presence of domesticates in the landscape. The communities of the Villeneuve-Saint-Germain (VSG) culture, which settled on the hillside less than 1 km from the valley bottom (Fig. 1C) (Germain-Vallée et al., 2017; Giazzon, 2012), do not appear to have interfered with the development of the mixed oak forest. The wetland edges, as well as Mesolithic clearings, were colonized by herbaceous and shrubby vegetation, including fruit trees (pollen and/or DNA from viburnum, elder, apple, and hazel) (Figs. 7 and 9). Thus, the shrinkage of the wetland and the emergence of riparian forest led to the development of an environment suitable for gathering (fruit trees), which seems to have played an important role in the diet of the populations during this period (Billard et Alix, 2014), as well as in animal husbandry (young shoots for grazing,

and the collection of hay or leaf fodder) (Delhon et al., 2008; Innes et al., 2013; Rasmussen, 1989).

During this period, crops were located in small clearings around settlements—found only on the plateaus (e.g. Billard and Alix, 2014; Germain-Vallée, 2023; Germain-Vallée et al., 2017; Giazzon, 2012; Le Saint-Allain et al., 2021)—and were too far from the valley bottoms for pollen evidence of arable cultivation to be recorded. Cattle may have been periodically driven into the valleys near the settlements to drink and/or graze. However, our data do not suggest the regular use of a large area for breeding in the Vey valley. Thus, we can assume that livestock were mainly kept close to settlements, likely to use their manure to fertilize cultivated areas (Bogaard, 2004; Bogaard et al., 2013; Fraser et al., 2013). Therefore, Early Neolithic land use had only a minor impact on woodland, and the data from Cairon are consistent with intensive garden cultivation, ascribed to Early Neolithic farmers of the Linear Pottery culture in Europe (Bogaard, 2004; Bogaard et al., 2013; Salavert, 2017).

### 5.3. An accurate record of the impact of mid-to-late neolithic human activity on the vey valley landscape

#### 5.3.1. The emergence of a mosaic landscape (Middle Neolithic I and II (4400–3780 BCE; 6350–5730 cal. BP; phases 3 and 4 Fig. 12);

At Cairon, from 4400 BCE onwards (6350 cal. BP), a Neolithic community was present, with an area used for religious purposes between 4400 and 4200 BCE (6350–6150 cal. BP), and a domestic settlement between 4200 and 4000 BCE (6150–5950 cal. BP) (Ghesquière and Marcigny, 2011). In our record, before 4200 BCE (6150 cal. BP), the cumulative percentages of ruderal taxa were not very high (>3 %), but *Plantago lanceolata*-t. was recorded almost continuously. Most notably, concentrations of *Sporormiella*-t. (HdV-113) reached a maximum (750–800 spores/g), coinciding with the detection of sheep DNA in one sample. From 4200 to 4030 BCE (6150–5980 cal. BP), the cumulative percentages of ruderal taxa remained stable, *Plantago lanceolata*-t. was recorded continuously, and concentrations of *Sporormiella*-t. (HdV-113) decreased slightly (from 900 to 200 spores/g). The first cereal pollen was also recorded (Phase 3 Fig. 12). Archaeozoological data from the domestic settlement soil (Ghesquière and Marcigny, 2011) showed that inhabitants raised cattle, pigs, caprinae, and sheep. Although cattle dominated the bone assemblages, the proportions of the different species were relatively balanced. According to carpological data (Ghesquière and Marcigny, 2011), emmer wheat, barley, wheat, and millet were cultivated (Fig. 12).

From 4400 BCE onwards (6350 cal. BP), fire peaks of greater magnitude (Fig. 11) were recorded in Cairon (2016) and could be interpreted as evidence of more extensive fires induced by humans during a new period of increased moisture in northwestern Europe (Goslin et al., 2018; Macklin et al., 2010) (Phase 3 Fig. 12). Between 4400 and 4200 BCE (6350–6150 cal. BP), a fire peak was contemporaneous with an increase in the percentages of ubiquitous meadow taxa, while the percentages of *Alnus*, *Betula*, and other hygrophilous taxa decreased (Fig. 7). Therefore, in the valley bottom, fires were likely related to the spread of meadows. Between 4200 and 4030 BCE (6150–5980 cal. BP), riparian forest spread across the valley bottom, and pioneer species such as birch decreased in the pollen spectra (Fig. 7). Fire peaks during this period were correlated with a decrease in the percentages of *Corylus*, *Quercus*, and *Ulmus*, indicating a reduction in dryland forest on the hillsides.

Archaeobotanical data from the domestic settlement soil (Ghesquière and Marcigny, 2011) show that oak, hazel, and Pomoideae were the three main tree species used on the Cairon site, especially as fuel. Our pollen data also indicate that oak and hazel were the two main species that decreased during this period (Fig. 7). A few pieces of pine charcoal were found in the anthracological dataset (Ghesquière and Marcigny, 2011). Our data did not support a significant presence of pine in the Vey valley's forests, since this taxon — which occurs at levels typical of
long-distance transport beyond its geographical range (Bennett, 1984) — was recorded at very low percentages in the pollen sequence but not in the sedaDNA (Figs. 7 and 9). This species was a poor source of fuel and, according to Marguerie (in Ghesquière and Marcigny, 2011), may have been selected and collected as timber from areas distant from the Cairon site. From the domestic settlement soil (Ghesquière and Marcigny, 2011), charcoal and seeds of hazel, elderberry, Prunus, and particularly Pomoideae were numerous, and some were consumed as food. In addition to hazelnut, the presence of these fruit trees was also recorded in the sedaDNA from Cairon (2016) (Figs. 7 and 9). These datasets illustrated the existence of open and semi-open ecosystems—such as forest edges, shrublands, and abandoned land—that likely spread following the establishment of permanent settlements and may have been used for gathering.

According to wood ring curvature analysis, some oak charcoal from the domestic settlement soil (Ghesquière and Marcigny, 2011) also came from areas of dense forest unaffected by agricultural clearance, where wild game—represented in the archaeozoological data (Ghesquière and Marcigny, 2011) by aurochs, stag, wild boar, and roe deer—may have been hunted. The bone remains of wild horse also suggested the existence of open areas. Riparian species were scarce in the anthracological assemblages (Ghesquière and Marcigny, 2011), indicating limited use of the riparian forest, which spread across the valley bottom according to our pollen data.

Taken together, data suggest that locally, farmers used fire to clear forests that were interspersed with meadows for grazing, cultivated plots, and abandoned land, and that the domestic and farming activities of the Middle Neolithic populations from the Cairon site led to the emergence of a mosaic landscape.

On Cairon 2016, from 4030 BCE (5980 cal. BP) (phase 4 Fig. 12), anthropogenic indicators slightly decreased, showing abandonment of cultivated areas and reduced pastoral pressure. Around 3900 BCE (5850 cal. BP), a fire peak coincided with declines in dryland forest and grassland. About a century after the abandonment of the domestic settlement (Ghesquière and Marcigny, 2011), part of the forest may have been burned to build the funerary monument at Cairon. Clearings spread locally, while riparian vegetation colonized the narrow alluvial wetland where pastoral pressure remained low.

### 5.3.2. Integration of the vey valley into a wider agricultural territory (late Middle Neolithic II to early Late Neolithic; 3780–3475 BCE; 5730–5425 cal. BP; phase 5 Fig. 10)

There was no archaeological evidence of any contemporaneous settlement in the Vey valley. Some traces of settlements on the upper hillsides and plateaus were known but had not been precisely dated (Fig. 2) (Ghesquière and Marcigny, 2011). However, on Cairon 2016, the cumulative percentages of ruderal taxa remained stable (1–4 %), and Plantago lanceolata-t. was recorded continuously. Cereal pollen was also recorded despite an increase in alder, which may have acted as a filter (Leroyer and Allenet de Ribemont, 2006).

In our sequence, the decline in percentages of the main dryland tree taxa was associated with an increase in ferns and a rise in the silty fraction of the deposits (Figs. 5 and 12). Upland forest was interspersed with cultivated and grazed clearings, as well as abandoned areas where ferns proliferated. No fire peaks were recorded, but the background level was maintained. Forest reduction was one factor favoring the increase in the background component, as charcoal could have been transported by erosive runoff with detrital sediments to the wetland (Clark et al., 1998; Conedera et al., 2009; Whitlock and Larsen, 2001). It could not be excluded that some smaller charcoal particles were also transported by wind. Thus, as with pollen in a less forested context, interception of this airborne component was less impeded (Jacobson and Bradshaw, 1981; Prentice, 1985; Sugita, 1994). Along the Cairon sequence, the background component curve generally evolved in opposition to that of the dryland tree percentages (Fig. 12). During this period, we can assume that charcoal inputs (by runoff and wind) contributed to maintaining the

background component, which may be related to human-induced clearing fires on a larger scale, indicating that the Vey valley had become part of a wider agricultural territory.

### 5.4. A gradual transformation of the landscape in the plain of caen from the Middle Neolithic

In the Caen Plain, the emergence of the Cerny culture at the beginning of the Middle Neolithic marked the end of the territorial unity characteristic of the Early Neolithic (Marcigny et al., 2007). From the second half of the 5th millennium BCE onwards, the construction of huge funerary monuments and large enclosures bounded by deep ditches probably involved the participation of many individuals. These monumental constructions suggest the emergence of organized communities with collective structures and may reflect a demographic shift (Ghesquière et al., 2014; Marcigny et al., 2007). Moreover, sites were found in the valleys near the rivers along the Seulles, to the north of the Caen Plain, as well as on the limestone plateaus to the south of Caen (e.g. Chancерel et al., 2006; Dron et al., 2016; San Juan and Dron, 1997) (Fig. 1). Thus, populations occupied and used a wide range of environments. In the center of the Paris basin (David et al., 2012; Leroyer, 2003, 2006; Leroyer and Allenet de Ribemont, 2006), the first anthropogenic pollen indicators were only found in densely settled areas or in areas as close as possible to the archaeological sites. The decrease in forest in favor of grassland, ruderal and ferns was related to clearings, and the first occurrences of cereal pollen were recorded in the Mue valley (Lespez et al., 2005) and the lower Orne valley (Lespez, 2012b). At Cairon, the high-temporal-resolution results revealed a succession of human activities that had a lasting impact on the vegetation landscape since the Middle Neolithic (4400 BCE; 6350 cal. BP) (Fig. 12). In the Vey valley, farmers used fire to clear the forest, first for pastoralism, then to build their houses and the funerary monument. Our new data supports archaeological hypotheses about demographic shifts. Isolated groups of people in the first part of the Middle Neolithic (4400–4000 BCE; 6350–5950 cal. BP) had only a restricted impact on landscape around the settlement. Subsequently, when a larger and organized community was able to build the funerary monument at Cairon, populations appeared to have a wider impact on the landscape.

During the Neolithic period, the valley was almost continuously occupied (Germain-Vallée et al., 2017; Ghesquière and Marcigny, 2011; Giazzon, 2012), which prevented the forest from regenerating in the long term. As a result, both locally and at the regional scale, anthracological datasets showed a clear increase in the taxonomic richness of charcoal, along with a significant rise in the proportions of Maloideae (Chancерel et al., 2006; Dron et al., 2016; Ghesquière and Marcigny, 2011; Mar- guerie and Hunot, 2007). Anthracological and dendrological data point to the persistence of dense forests dominated by mature oaks. However, some charcoal also come from young specimens that had grown in regenerating open oak stands (Chancерel et al., 2006; Dron et al., 2016; Ghesquière and Marcigny, 2011; Mar- guerie and Hunot, 2007). Already present in the Early Neolithic, Maloideae could survive in undergrowth, and forest clearing may have promoted their development. Remains of domestic animals (cattle, followed by goats and pigs) predominated in faunal records of the Caen Plain (Chancерel et al., 2006; Dron et al., 2016; Ghesquière and Marcigny, 2011; San Juan and Dron, 1997). Wild fauna bones were mainly from large game (aurochs, deer, wild boar, and roe deer), with additional remains including a bear phalanx and horse bones (Ghesquière and Marcigny, 2011; Chancерel et al., 2006). Domestic and wild fauna both pointed to the presence of dense forests, as well as more open forests and clearings. Carpological datasets were more diversified, and data showed the presence of undergrowth, forest edges and abandoned land alongside cultivated clearings (Chancерel et al., 2006; Dron et al., 2016; Ghesquière and Marcigny, 2011; San Juan and Dron, 1997). Thus, the Cairon record in comparison with available archaeobotanical and palynological data from the Caen Plain, highlighted the use of a wide range of environments, which has resulted in
the development of a mosaic landscape with secondary woodland formations, meadows, fields, and abandoned land.

In agreement with other results obtained in northwestern Europe, some open or semi-open areas may have been preferentially exploited and subsequently maintained by Neolithic farmers, as they were suited to pastoral economies (Fyfe et al., 2015; Hudson et al., 2022). Occupation of the clearings was usually associated with temporary and localized erosion phases in the upper part of the soils (Béguier et al., 2011; Germain-Vallée and Lespez, 2011), and once the clearings were abandoned, the forest regenerated and continued to dominate regionally. However, over time, forest reduction led to the transfer of sediments from soil erosion to the bottom of the valley. After 4000 BCE (5950 cal. BP) silty inputs contributed to the mineralization and degradation of the peaty marsh at Cairon, favoring the development of a dense alder forest (Fig. 10). It was only after 3500 BCE (5450 cal. BP) that the peatland was definitively buried by detrital silts (Fig. 5).

## 6. Conclusion

The multi-proxy data here provide a new high-resolution model of the environmental context and impact of the first farmers in the plateaus of northwestern France. Over this 2500-year period, we observe.

* A late Mesolithic high-diversity mixed wet and dry deciduous woodland with gaps as well as aquatic resources and wild pig present in sedaDNA. The high charcoal in this period (5900–5195 BCE; 7850–7145 cal. BP) suggests that fire may have been used to maintain this openness.

* An Early Neolithic increase in woodland density and development of alder wet-woodland which occurred simultaneously with the appearance of cattle in the sedaDNA (5195–4400 BCE; 7145–6350 cal. BP). This and the relatively low levels of open-land indicators in both the pollen and sedaDNA suggests that the grazing might have been woodland dominated – as both taxa are well adapted to this.

* In the Middle Neolithic (4400–4200 BCE; 6350–6150 cal. BP) floristic changes in the pollen suggest grassland expansion and sheep is present in the sedaDNA. In the later Middle Neolithic (4200–3475 BCE; 6150–5425 cal. BP), there was an elm decline, and cereal cultivation evident from the pollen, along with an expansion of wet woodlands. The high charcoal in this period was related to clearing of dry deciduous woodland along with sedimentary and NPPs indicators of floodplain change.

* The comparison of plant sedaDNA with the pollen provides confirmatory evidence that the pine in the pollen is long-distance and not local and it is likely that the elm and lime were growing on the plateau. Conversely the sedaDNA suggests that viburnum, poplar and apple-family trees were growing on the floodplain or at its edge. Whilst the floodplain sequence was not as rich as a typical lake sedaDNA record, the data both add floristic depth and additional evidence to the other proxies.

The sequence reflects the closest archaeological site in.

* The archaeological comparison, which clearly shows the impact of the different types of occupation on the local vegetation, both in relation to the establishment of the religious area and the domestic settlement of Cairon, as well as vegetation change on a larger scale following the construction of the funerary monument.

* Providing more reference on fungal spores considered partly coprophilous, supporting the relevance of *Sporormiella*-t. as an indicator of changes in local grazing pressure, as opposed to taxa with stronger affinities for dead plant material related to peaty soils.

* Hydrological changes inferred from local ecological succession dynamics, related both to changes in the water table and to changes in human practices and pressure in the valley.

## Authors' contributions

**Laurence Lemer:** Field Investigations, NPPs and charcoal analyses. Writing — Original Draft. **Agnès Gauthier:** Supervision, Field Investigations, Writing — Review & Editing. **Laurent Lespez:** Supervision, Field Investigations, Writing — Review & Editing. **Antony Brown:** SedaDNA analysis. Writing — Review & Editing. **Samuel Hudson:** SedaDNA analysis. Writing — Review & Editing. **Marta Garcia:** XRF core scanner analyses. **Kazuyo Tachikawa:** XRF core scanner analyses. **Edouard Bard:** XRF core scanner analyses. **François Giligny:** Review & Editing. **José Antonio López-Sáez:** Review & Editing.

## Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

## Acknowledgements and funding

This work has received financial support from the LabEx Dynamite (ANR-11-LABX-0046), as part of the "Investissements d'Avenir" program, the University of Paris 1, the Regional Archaeology Department of Normandy and the Departmental Council of the Calvados region. Radiocarbon ages have been supported by LMC14 (Saclay, France) as part of the ARTEMIS program. We are also grateful to Axel Beauchamp and Cécile Germain-Vallée (field investigations, funding and magnetic susceptibility), Clement Virmoux (field investigations), Thierry Fonville (sedaDNA extraction), Lucile de Milleville (software) and Anne Ropars (data from the *carte archéologique* database).

## Appendix A. Supplementary data

Supplementary data to this article can be found online at https://doi.org/10.1016/j.quascirev.2025.109644.

## Data availability

A link to the data and/or code is provided as part of this submission. Pollen and non-pollen palynomorph counts are available in Supplementary data 2, and sedaDNA data is available in Supplementary Data 3. The raw sedaDNA data is held in the European Nucleotide Archive (ENA) (https://www.ebi.ac.uk/ena/). The accession code for the project is: PRJEB98234 and the tag-look up file is attached in Supplementary Data 4 which can be used for both the P6-loop and 16S libraries.

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