Quaternary Science Reviews 278 (2022) 107373 Contents lists available at ScienceDirect Quaternary Science Reviews journal homepage: www.elsevier.com/locate/quascirev Landscape dynamics and fire regime since 17,550 cal yr BP in the Cantabrian region (La Molina peat bog, Puente Viesgo, Spain) Marc Sanchez-Morales a, *, Albert Pe lachs a, Juan Carlos García-Codron b, Virginia Carracedo b, Ramon Pe rez-Obiol c a noma de Barcelona, 08193, Cerdanyola del Vall Department of Geography, Edifici B, Facultat de Filosofia i Lletres, Universitat Auto es, Bellaterra, Spain b Department of Geography, Urban Studies and Land Planning, Avenida de Los Castros s/n, Universidad de Cantabria, 39005, Santander, Spain c Department of Animal Biology, Plant Biology, and Ecology, Universitat Auto noma de Barcelona, 08193, Bellaterra, Spain a r t i c l e i n f o a b s t r a c t Article history: Our aim with this paper is to provide insight into the landscape dynamics of the Cantabrian region Received 12 October 2021 (northern Iberian Peninsula) from the Last Glacial Period to the present. We present a multiproxy Received in revised form approach performed in a sedimentary record from La Molina peat bog, located at 484 m a. s. l. in Puente 28 December 2021 Viesgo, Cantabria (Latitude: 43.26 N; Longitude: 3.97 W), that covers the last c. 17,550 years. Analyses Accepted 3 January 2022 Available online xxx were performed on the organic matter, pollen and sedimentary charcoals, which have been used to characterise the fire regime at a local level. The results revealed a steppe-like formation from the basis of Handling Editor: Yan Zhao the sequence to the Bølling-Allerød chronozone, when the spread of woody species is detected. The dry and cold conditions of the Younger Dryas caused the decrease of mesophilous taxa until the onset of the Keywords: Holocene. From that point onwards, a mixed deciduous formation composed of Betula and deciduous Paleogeography Quercus was established in the region. Besides, a series of biomass pulses was detected during the early Vegetation dynamics Holocene, probably linked to intervals of increased rainfall. Fire episodes detected at the 9.3 and the 8.2 Last Glacial Period ka events created forest openings and probably favoured the establishment of Corylus, most likely also Holocene helped by a climate shift. There was a lack of fire activity until the Neolithic, when the fire signal Palynology increased probably linked to grazing and agricultural practices and triggered deeper changes in the Sedimentary charcoal Western Europe Cantabrian plant landscape structure. Iberian Peninsula © 2022 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND Biogeography license (http://creativecommons.org/licenses/by-nc-nd/4.0/). Environmental history 1. Introduction assessing the influence of the anthropic factor as a landscape- shaping agent. The Cantabrian range (northern Iberian Peninsula) is a moun- Concerning the role of climate (Fig. 1), the northern Iberian tain system that runs parallel to the Cantabrian Sea. Due to its Peninsula has been under the influence of regional climatic vari- orography and climate features, the area is of particular environ- ability, such as the succession between Stadials and Interstadials mental interest since it is a natural border between the Euro- (Dansgaard et al., 1982; Johnsen et al., 1992; Rasmussen et al., 2014; siberian and Mediterranean biogeographical regions (Rivas- Seierstad et al., 2014), the Heinrich events (Heinrich, 1988; Bond Martínez et al., 2017) and determines the distribution pattern of et al., 1992; Hemming, 2004) and the Bond cycles (Bond et al., several terrestrial species. The northern slope of the mountain 1997; Bond et al., 2001; Mayewski et al., 2004; Wanner and range, also known as the Cantabrian region, has proved to be an Bütikofer, 2008; Isono et al., 2009; Wanner et al., 2011). Several exceptional scenario for the study of palaeoecological dynamics. A studies have examined the impact that this climate variability has large number of natural registers, such as caves, rocky shelters and had on the Cantabrian region through isotopic analyses of cave sedimentary deposits, have allowed characterising the environ- speleothems (Domínguez-Villar et al., 2008, 2009, 2017; Moreno mental and climate change over the last millennia as well as et al., 2010a; Baldini et al., 2015, 2019; Smith et al., 2016; Rossi et al., 2018). For example, Baldini et al. (2019) modelled the sea- sonal distribution of temperature and rainfall from the Younger * Corresponding author. Dryas, while Smith et al. (2016) revealed quasi-cyclical events of nchez-Morales). E-mail address: marc.sanchez.morales@uab.cat (M. Sa https://doi.org/10.1016/j.quascirev.2022.107373 0277-3791/© 2022 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/). nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 their presence was detected at high altitudes of the northern Abbreviations Cantabrian range (Barandiara n et al., 2006). The expansion of hu- man groups continued during the Mesolithic with more than 250 LGP Last Glacial Period documented sites in Cantabria, 30 of which are in the Pas Valley, the GS Greenland Stadial same as for La Molina peat bog (Pe rez-Bartolome , 2019). Further on, GI Greenland Interstadial the onset of agricultural practices triggered the transition from PAR pollen accumulation rate hunter-gatherer to farmer societies (Arias, 1991) which took place CONISS constrained cluster analysis by the method of at between 7700 and 6800 cal yr BP along the Cantabrian range incremental sum of squares (Fano et al., 2015). The expansion of human groups was evident PAZ pollen assemblage zones across all the territory since approximately 1250 megaliths have CHAC charcoal concentration been documented from sea level to more than 1800 m a. s. l. (Arias, CHAR charcoal accumulation rate 2005). The need to create forest openings was accompanied by important fire events detected in La Molina (Pe rez-Obiol et al., 2016) and in higher altitude environments (Carracedo et al., 2018). Metal mining exploitation stands out as the most impor- wet-to-dry conditions during the Holocene in Cantabria. tant impact on landscape during the following cultural phases (e.g. The cultural heritage of Cantabria, located on the easternmost Manteco  n, 2000; Arias, 2005; Blas-Cortina, 2014), while human flank of the Cantabrian range, is rich in prehistoric anthropogenic society exhibited an increasing control of landscape until contem- evidence. Up to now El Castillo cave, at about 3e4 km from La porary dates. Molina peat bog, has been the site with the oldest documented Reconstruction of the plant dynamics has also been possible human presence, dated at 89,000 years BP (Bischoff et al., 1992). In thanks to the large number of palynological records made all over a more recent period, populations grew up across the Cantabrian the Cantabrian range (Fig. 2). The characterisation of the Last region (Gonza lez-Sainz, 1989; Gonza lez-Sainz and Gonza lez- Glacial Period (LGP) has otherwise only been possible above Urquijo, 2004; Chauvin, 2007; Chauvin et al., 2018; Fano et al., 1300 m a. s. l. From West to East, the sequences of Laguna de 2020) coinciding with the temperate climate of the Bølling- Lucenza (Aira-Rodríguez, 1986; Santos-Fidalgo et al., 1997; Mun ~ oz- Allerød chronozone (GI-1; Iversen, 1953; Rasmussen et al., 2014). Sobrino et al., 2001), Villaseca de Laciana (Jalut et al., 2010), La Mata Human groups expanded towards mid-mountain areas (0e700 m (Jalut et al., 2010), Lago de Ajo (Allen et al., 1996), Puerto de Tarna a. s. l.) along the Upper Magdalenian and the first stages of the (Ruíz-Zapata et al., 2000) and Puertos de Ríofrio (Mene ndez-Amor Azilian (Fig. 1), achieving an increasing control of landscape with and Florschütz, 1963) revealed an open area with a large presence important migration rates among sites (Fern andez-Tresguerres, of xerophytes during some intervals of the LGP. A few more records 2004; Gonz alez-Sainz and Gonza lez-Urquijo, 2004). During the from Burgos (north-eastern Castile and Leo n), such as La Piedra Azilian (Fig. 1), the Epipalaeolithic culture developed in several (Ramil-Rego et al., 1998), San Mame s de Abar (Iriarte-Chiapusso eastern sites from the northern Cantabrian range among other et al., 2001) and La Nava (Mene ndez-Amor, 1968), documented a European regions, human populations still based their economies higher presence of pines and birches, while the dominance of pines on hunter-gatherer activities (Ferna ndez-Tresguerres, 2004), and becomes more evident in the northern Iberian System as indicated Fig. 1. Cultural stages in Cantabria from 22,000 cal yr BP to the present and the climatic characterisation of the sequence. Boundaries for the cultural stages have been drawn according to Gonz alez-Sainz (1994), Rasilla-Vives and Straus (2004), Gonza lez-Sainz and Gonza lez-Urquijo (2004), Marín-Arroyo et al. (2018) and Straus (2018) for the Palaeolithic interval; to Gonzalez-Sainz and Gonzalez-Urquijo (2004), Alday-Ruiz (2009), Fano et al. (2015) and Straus (2018) for the Azilian, Mesolithic and Neolithic; Arias (1995), Ontan ~o n (2003) and Gonz alez-Rabanal et al. (2020) for the Chalcolithic; Marín-Suarez (2011) for the Bronze and Iron Ages; Costa-García (2018) for the onset of the Roman Period. Climate: Heinrich event 1 identified and discussed by Heinrich (1988), Bond et al. (1992) and Hemming (2004). The chronology for the Würm glaciation based on Greenland Stadials (GS) and Greenland Interstadials (GI) is shown (Rasmussen et al., 2014), and also the stages for the Holocene based on Walker et al. (2018, 2019). 2 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 Fig. 2. Top: geographical map of the northern Iberian Peninsula. Bottom: palaeoecological records studied in the western sector of the Cantabrian range. The location of La Molina peat bog is highlighted as a yellow dot. Legend: 1: Enol lake (Mene ndez-Amor, 1975; Lo  pez-Merino, 2009; Moreno et al., 2010b, 2011) 2: Ercina lake (Mene ndez-Amor, 1975) 3: Comella/Comeya (Ruiz-Zapata et al., 2001; Jime nez-Sanchez et al., 2003) 4: Esla (Sanchez-Hernando et al., 1999) 5: Puertos de Ríofrio (Mene ndez-Amor and Florschütz, 1963) 6: Vidiago (Mene ndez-Amor, 1950a, 1950b) 7: Llano de Mesa (Mene ndez-Amor, 1950a, 1950b) 8: Pendueles (Mene ndez-Amor, 1950a, 1950b; Lo  pez-Días et al., 2013a) 9: Buelna (Mene ndez-Amor, 1950a, 1950b; Mene ndez-Amor and Florschütz, 1961; Lo  pez-Días et al., 2013a) 10: Borbolla (García-Amorena et al., 2008; Lo  pez-Días et al., 2013a) 11: Llano Ron~ anzas (Mene ndez-Amor, 1950a, 1950b; Ortiz et al., 2008, 2010; Moreno et al., 2009; Gallego et al., 2013; Lo  pez-Días et al., 2013a, 2013b) 12: Las Arenas e Tina Mayor (Mary et al.,  n (Mary, 1990; García-Amorena et al., 2008) 17: Oyambre (Mary, 1990; 1975) 13: Río Bederna (Mary et al., 1975) 14: Jerra I (Mary et al., 1975) 15: Jerra II (Mary et al., 1975) 16: Mero García-Amorena et al., 2008) 18: Culazo  n (Gonza lez-Pellejero et al., 2014) 19: Pico Sertal (Mariscal, 1986; Carracedo et al., 2018) 20: El Cueto de la Avellanosa (Mariscal, 1983; Carracedo et al., 2018; Nun ~ ez, 2018) 21: La Molina (Pe rez-Obiol et al., 2016; Carracedo et al., 2018; this paper) 22: Pico Ano (Salas, 1993) 23: Alsa (Mariscal, 1993) 24: Santa Gadea (Iriarte-Chiapusso et al., 2003) 25: La Nava (Mene ndez-Amor, 1968) 26: El Cueto de la Espina (Rodríguez-Cotero n, unpublished PhD; Sa nchez-Morales, unpublished PhD) 27: Puerto del Escudo (Mun ~ oz-Sobrino, 2001) 28: Estacas de Trueba (Mariscal, 1987, 1989) 29: Sotombo (Pe rez-Díaz et al., 2016a) 30: Noja (García-Amorena et al., 2008) 31: Los Tornos (Pen~ alba, 1994; Mun ~ oz-Sobrino et al., 2005) 32: Zalama (Pe rez-Díaz et al., 2016b; Souto et al., 2016; Souto, 2018) 33: San Mame s de Abar (Iriarte-Chiapusso et al., 2001, 2016; Mun ~ oz-Sobrino, 2001) 34: La Piedra (Ramil-Rego et al., 1998) 35: Tubilla del Agua (Moreno-Amat et al., 2009; García-Amorena et al., 2011) 36: Huidobro (Iriarte-Chiapusso et al., 2003) 37: Arreo Lake (Corella et al., 2013). (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.) by the papers on Hoyos de Iregua (Gil-García et al., 2002), Laguna Tornos (Pen ~ alba, 1994; Mun ~ oz-Sobrino et al., 2005) and La Molina del Hornillo (Go  mez-Lobo et al., 1996), Laguna Negra (Von rez-Obiol et al., 2016). (Pe EngerIbrechten, 1998), Lago Las Pardillas (Sa nchez-Gon~ i and Among all the mentioned sites, however, only a few studies Hannon, 1999), Quintanar de la Sierra (Pen ~ alba, 1994; Pen~ alba considered the role of fire in the reconstruction of the landscape. et al., 1997) and Laguna Grande (Ruiz-Zapata et al., 2002). This may lead to partial interpretations, since fire is an important On the northern slope of the Cantabrian range, several palyno- factor in the complex scenario between vegetation, climate and logical records have been used to characterise the plant dynamics anthropic pressure (Harrison et al., 2010; Whitlock et al., 2010; throughout the Holocene (Fig. 2), mainly dominated by mixed de- Krawchuk and Moritz, 2011). On the one hand, biomass burnings ciduous formations. In Asturias, the papers on El Alto de la Espina seem to have been strongly influenced by climate variability during pez-Merino, 2009), Las Duen (Lo ~ as (Lo  pez-Merino et al., 2006), El certain glacial and postglacial intervals (Power et al., 2008; Daniau Monte Areo peat bogs (Lo pez-Merino, 2009; Lo  pez-Merino et al., et al., 2010, 2012; Marlon et al., 2016). In this line, several papers 2010) and the Enol lake (Lo  pez-Merino, 2009) characterised the have examined the influence of both the climate and anthropic landscape from the early Holocene to the present and stand out as factors on determining the frequency and intensity of the fire the most relevant. In Cantabria, located in the eastern sector of the regime over the Holocene, especially from the Neolithic period mountain system (Fig. 1), the main sequences, covering from the onwards (e.g., Vannie re et al., 2008; Rius et al., 2009; Mercuri and Mid-Holocene to the present, are the obtained from the peat bogs Sadori, 2014; Gil-Romera et al., 2014; Sadori et al., 2015; Beffa of Culazo  n (Gonza lez-Pellejero et al., 2014), Pico Sertal (Mariscal, et al., 2016). The sedimentary charcoal records performed in El 1986; Carracedo et al., 2018), El Cueto de la Avellanosa (Mariscal, Sertal, El Cueto de la Avellanosa (Carracedo et al., 2018) and La 1983; Nun ~ ez, 2018), Pico Ano (Salas, 1993), Alsa (Mariscal, 1993), Molina (Pe rez-Obiol et al., 2016), all located in the eastern sector of Puerto del Escudo (Mun ~ oz-Sobrino, 2001), Estacas de Trueba the Cantabrian range, indicated intense use of fire from c. (Mariscal, 1987, 1989), Sotombo (Pe rez-Díaz et al., 2016a), Los 7000 cal yr BP coinciding with the onset of agricultural practices 3 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 and the need to create forest openings, thus revealing an anthropic Brushwood groves also develop and cover large extensions on origin. Nevertheless, the lack of research on the role of fire has mid-mountain areas. They are favoured by openings generated by made it difficult to determine how long fire activity has been pre- anthropogenic activities such as grazing and recurring fires, which sent in the Cantabrian region. can be replicated over centuries or millennia (Pe rez-Obiol et al., Here we present a multiproxy approach performed in a sedi- 2016; Carracedo et al., 2018). The most recurrent species are mentary record from La Molina peat bog (Cantabria) from c. 6740 to Rhamnus cathartica, Prunus spinosa, Calluna vulgaris, Daboecia rez-Obiol et al. (2016) studied the most 17,550 cal yr BP. Since Pe cantabrica, Erica vagans, Erica mackaiana, Erica ciliaris, Ulex euro- recent 6740 years, we obtained palynological, charcoal and organic paeus and Ulex gallii. matter records spanning from the LGP to the present. In this In the mountain environment, birchwoods of Betula pubescens respect, the sequence of La Molina covers an unstudied interval of (¼ Betula alba) appear on nutrient-poor siliceous soils and steep the LGP in the northern Cantabrian range, from the GS-2.1a on- slopes. These communities hold a scattered presence of Sorbus aria, wards. The appropriate chronological resolution also provides Sorbus aucuparia, Quercus petraea, Fagus sylvatica, Taxus baccata and added value to the sequence. Thus, the main objective of this study Ilex aquifolium. Also, there is a great variety of grasslands in cry- was to characterise the vegetation dynamics and fire regime of the orotemperate and supra-orotemperate environments (Díaz- north-eastern Cantabrian range since c. 17,550 cal yr BP. We sought Gonza lez and Penas, 2017). to assess how the plant landscape changed during the different In contrast to the Atlantic façade, the southern slope presents climatic and cultural phases and also determine whether the colder winters, a water deficit in summer and less than 800 mm of northern slope of the mountain system was a glacial refuge for annual precipitation (Csb, according to the Ko €ppen-Geiger classi- certain plant species. We aimed to discern the source of landscape fication: Beck et al., 2018). Accordingly, the tree layer of the conti- changes by interpreting whether they are linked to climatic fea- nental slope is mainly composed of marcescent and evergreen oaks tures, anthropic pressure or to a combination of both. Besides, we (Penas et al., 2017). hoped to determine from when there have been fires in the study rez-Obiol et al. (2016) described the vegetation of the peat Pe area and from what point they have played a significant role in the bog. They identified Molinia caerulea, Rhynchospora alba, Erica tet- configuration of the plant landscape. ralix, Drosera rotundifolia and Eriophorum angustifolium among other hygrophilous taxa and peat mosses of the genus Sphagnum. 1.1. Study setting Besides, there are a few individuals of Betula pubescens and Salix spp. on the edge of the peat bog, while in the surrounding area the La Molina is an acidic peat bog in the municipality of Puente potential formations are composed of Quercus robur and Fraxinus Viesgo (Cantabria), approximately 25 km south of Santander excelsior (Rivas-Martínez et al., 1987). Although there are some (Latitude: 43.26 N; Longitude: 3.97 W; Altitude: 484 m a. s. l.). patches of deciduous communities, shrublands are the principal The site is in the northern foothills of the Cantabrian range in the formations, mainly composed of Ulex spp., Pteridium aquilinum and Pas valley close to the Pas-Besaya water-divide. The surrounding Erica spp. and are accompanied by several plantations of Pinus spp. terrain consists of sandstones, silts and clays, and there are some and Eucalyptus spp. (Fig. 3). Approximately 3 km north of the peat other minerotrophic peat bogs nearby which are smaller than La bog, El Monte Castillo (355 m a. s. l.) stands out as a limestone Molina. formation that hosts a forest of Quercus ilex subsp. ilex on the The orography of the Cantabrian range has important implica- southern slope while there is a plantation of Eucalyptus spp. in the tions in the climate configuration of the northern Iberian Peninsula. northern area (Fig. 3). Moisture-laden oceanic winds usually condense on the northern slope leading to annual precipitations ranging from 1000 to 2. Materials and methods 1800 mm (climate Cfb according to the Ko €ppen-Geiger classifica- tion: Beck et al., 2018), reaching over 2000 mm at the highest al- 2.1. Sampling titudes (Ancell and Ce lis-Díaz, 2012). Accordingly, the principal plant formations along the Cantabrian In 2013, a sampling campaign was conducted in La Molina peat low valleys and mid-mountain areas are subatlantic and sub- bog by Pe rez-Obiol et al. (2016). A continuous sedimentary record mediterranean broad leaved mixed forests (Costa-Tenorio et al., (MOL2) was retrieved from the top to 260 cm depth by using a PVC 2005). In particular Fagus sylvatica, Quercus robur and Quercus tube, 11 cm in diameter. Analyses on the organic matter, sedi- petraea, all of them with a Eurosiberian distribution, are the forest mentary charcoals (>150 mm), pollen and metal analyses (titanium species with the largest presence on the northern slope of the and aluminium) enabled the reconstruction of the environmental Cantabrian range. Fagus sylvatica is well adapted to both siliceous geohistory of the region for the last 6740 cal yr BP. Another record and calcareous lithology and has a greater presence on the eastern (MOL4 m1, from 367 to 467 cm depth) was also sampled in 2013 to side of the Cantabrian range. In terms of oak trees, Quercus robur is estimate the bottom age of the peat bog, with a basal age of c. the most common in the Cantabrian sector (Amigo et al., 2017) and 18,840 cal yr BP at 467 cm depth (Table 1). is dominant on the siliceous basal plains (Costa-Tenorio et al., Due to the long chronology and optimum preservation of the 2005). Quercus petraea, however, is less resistant to cold winters MOL2 sediment, a second sampling campaign was performed in and is therefore found at a higher altitude range to avoid thermal 2016 to sample the oldest sediments from La Molina. Ten sedi- inversion and also in some points on the southern mountain slope mentary cores were sampled using a Russian peat corer with a total (Costa-Tenorio et al., 2005). On the other hand, recent plantations length of 499 cm. Under laboratory conditions, the colour of the of Pinus radiata, P. sylvestris, P. pinaster, P. nigra and Eucalyptus spp. sediment was determined with a Munsell colour chart. The MOL6 constitute important components of the current landscape records were then kept in a fridge (4  C) until they were cut into a composition. total of 654 samples around 1 cm (Table 1). Communities dominated by Quercus ilex subsp. ilex and Laurus nobilis are a common formation on the Cantabrian littoral, while 2.2. Chronology Quercus ilex subsp. rotundifolia (¼ Quercus rotundifolia) develop at higher stages on poorly developed mountain soils (Costa-Tenorio rez-Obiol et al. (2016) established an age-depth chronological Pe et al., 2005; Sainz-Ollero et al., 2010; Amigo et al., 2017). model for the MOL2 record of La Molina peat bog using radiocarbon 4 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 ~a Fig. 3. Left: Potential vegetal map, modified from Sainz-Ollero et al. (2010) Right: Vegetation formations surrounding La Molina peat bog. Modified from Mapa forestal de Espan 1:25,000 (Miteco, 2014). Table 1 2.3. Organic matter Sampling year, code, depth and number of samples of each record from La Molina peat bog. Analyses of the organic matter content were carried out on 383 Sampling year Code of the record Depth (cm) Samples samples from MOL6 (499 - 250 cm depth; c. 17,550e6190 cal yr BP) 2013 MOL2 0e260 302 using 1 g of wet sample. The samples were put in an oven for 24 h at 2013 MOL4 m1 367e467 e 60  C and after this process dry weight was measured. Organic 2016 MOL6 m1 0e50 48 content was determined by the weight loss after ignition according MOL6 m2 50e100 50 to standard procedures (4 h, 550  C; modified from Dean, 1974; MOL6 m3 100e150 56 MOL6 m4 150e200 55 Heiri et al., 2001). A single organic matter sequence (MOL2/MOL6) MOL6 m5 200e250 62 was obtained by the combination of 287 samples from the top MOL6 m6 250e300 61 250 cm of MOL2, analysed in Pe rez-Obiol et al. (2016), and 383 MOL6 m7 300e350 63 samples of the interval from 250 to 499 cm of MOL6, obtaining a MOL6 m8 350e400 64 mean annual resolution of 26.19 years per sample. MOL6 m9 400e450 92 MOL6 m10 450e499 103 2.4. Pollen dates (n ¼ 5) and one documental datum related to Pinus spp. and A total of 251 samples from MOL6 (499 - 250 cm depth; c. Eucalyptus spp. plantations that occurred at 1950 years AD (at 17,550e6190 cal yr BP) were treated for palynological analyses 14 cm depth). For the chronological model used in this work, ten following standard chemical procedures. One gram of each sample 14 C AMS dates (Beta Analytic Inc.) from the MOL6 record were was weighed and sieved through a 300 mm mesh. A tablet of added to the initial model. Both the 14C ages from MOL2 and MOL6 Lycopodium spores was added to estimate the pollen concentration were calibrated using the IntCal20 Northern Hemisphere radio- (Stockmarr, 1971). The chemical procedure included washes with carbon age calibration curve (Reimer et al., 2020). In addition, the HCl (10%), HF (70%), KOH (10%) and acetolysis, followed by glycerol chronology from the most superficial sediments of MOL2 was mounting (Faegri and Iversen, 1989; Moore et al., 1991; Bennett and established by measuring the 210Pb specific activity of ten samples Willis, 2001). from 0 to 20 cm depth. Measurements of 210Pb were performed The counting and identification were carried out under an op- applying a constant rate of supply model (Appleby and Oldfield, tical microscope using reference collections from the Universitat 1978). The documental datum included in the chronological Auto noma de Barcelona, visual atlases (e.g., Reille, 1992, 1998; van model used in Pe rez-Obiol et al. (2016) was not considered since we Geel and Aptroot, 2006) and morphological identification keys (e.g., had the new 210Pb-based surface age model. A 14C date from MOL2 Faegri and Iversen, 1989; Moore et al., 1991). was also rejected because it did not fit into the 95% confidence Pollen percentages were calculated as a percentage of the total interval of the chronological model (Table 2). Considering this data terrestrial pollen, thus excluding hygrophytes, aquatic plants and set, the MOL2/MOL6 age-depth model was performed using the non-pollen palynomorphs. Also, the total pollen concentration of open-source environment RStudio for R (RStudio, 2020), the Clam the terrestrial pollen (arboreal and non-arboreal grains cm3) and package (Blaauw, 2010) and smoothing spline interpolation (Fig. 5). the pollen influx (palynomorph cm2 yr1; Bennet and Willis, 2001), also known as the pollen accumulation rate (PAR), were calculated and included in the diagrams. 5 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 Table 2 Chronological data and calibrated ages for La Molina cores (MOL2 and MOL6). Radiocarbon dates were calibrated using the IntCal20 Northern Hemisphere radiocarbon age calibration curve (Reimer et al., 2020). Core Laboratory code Depth (cm) Material Dating 13 C/12C (‰) Conventional radiocarbon Calibrated age (2s) Median probability age, used for dated technique age (yr BP) (cal yr BP) chronological model (cal yr BP) MOL2 Surface 0 peat e e e e 63 210 MOL2 e 1.02 peat Pb e e e 63 210 MOL2 e 3.07 peat Pb e e e 61 210 MOL2 e 5.11 peat Pb e e e 55 210 MOL2 e 7.16 peat Pb e e e 47 210 MOL2 e 9.2 peat Pb e e e 38 210 MOL2 e 11.25 peat Pb e e e 30 210 MOL2 e 13.29 peat Pb e e e 23 210 MOL2 e 15.34 peat Pb e e e 19 210 MOL2 e 17.38 peat Pb e e e 6 210 MOL2 e 20.71 peat Pb e e e 22 14 MOL2 Beta-371859 40 peat C 27.5 650 ± 30 555e605 600 14 MOL2 Beta-385973 68 peat C 27.3 3340 ± 30 e Rejected 14 MOL2 Beta-371860 113 peat C 25.9 3480 ± 30 3687e3836 3757 14 MOL2 Beta-371861 186 peat C 26.6 4130 ± 30 4567e4729 4676 14 MOL6 Beta-490941 252 peat C 27.4 5280 ± 30 5988e6122 6079 14 MOL2 Beta-360118 260 peat C 27.7 5910 ± 30 6661e6795 6726 14 MOL6 Beta-490942 299 peat C 28.1 8060 ± 40 8846e9032 8972 14 MOL6 Beta-480943 302 peat C 27.8 7840 ± 30 8543e8656 8616 14 MOL6 Beta-480942 348 peat C 29 9180 ± 30 10,244e10,420 10,329 14 MOL6 Beta-475549 352 peat C 29.8 9380 ± 30 10,510e10,691 10,611 14 MOL6 Beta-475548 399 peat C 28.5 11,940 ± 30 13,751e13,871 13,806 14 MOL6 Beta-475547 401 peat C 28.2 12,270 ± 30 14,076e14,324 14,178 14 MOL6 Beta-475546 449 peat C 27.3 13,800 ± 30 16,581e16,929 16,749 14 MOL6 Beta-468818 450 peat C 29.5 12,830 ± 60 15,137e15,552 15,323 14 MOL6 Beta-468819 499 peat C 27.1 14,400 ± 40 17,368e17,802 17,552 The palynological diagrams (MOL2/MOL6) included 144 pollen from the MOL2 record (from 0 to 250 cm depth) and 193 charcoal samples counted in the MOL2 record (0e250 cm depth, previously samples analysed from the MOL6 record (from 250 to 499 cm analysed in Pe rez-Obiol et al., 2016) and 251 pollen samples depth), obtaining a mean annual resolution of 36.5 years per counted in the MOL6 record (250e499 cm depth). The mean annual sample. Charcoal counts were divided by the sample volume and resolution of the MOL2/MOL6 record was of 44.4 years per sample. then by the sedimentation rate to obtain the charcoal concentration Pollen diagrams were constructed using the Tilia and Tiliagraph (CHAC, particles cm3) and the charcoal accumulation rate (CHAR, software (Grimm, 1991). Pollen assemblage zones (PAZ) were particles cm2 yr1), respectively. Also, charcoal peaks were determined through a constrained cluster analysis by the method determined by a Gaussian mixture model for each window of 500 of incremental sum of squares (CONISS; Grimm, 1987). PAZ were years, with a threshold of the 95% of modelled noise distribution. identified with La Molina acronym (MOL) followed by an identifi- The signal-to-noise index was calculated to quantify the suitability cation number and a letter in the case of the subzones (e.g., MOL/ of the charcoal record for peak analysis (Kelly et al., 2011). A peak 1a). analysis was performed to identify the main charcoal peaks and the peak's magnitude (pieces cm2 peak1). All the analyses were performed using the CharAnalysis v1.1 software (Higuera et al., 2.5. Sedimentary charcoals 2009). The fire regime interpreted from the charcoal record must have Sedimentary charcoals (>150 mm) were analysed on 193 sam- had a local origin (within the watershed) since charcoal particles ples from the MOL6 record (499 - 250 cm depth; c. larger than 150 mm are known to be deposited in an area close from 17,550e6190 cal yr BP). The chemical procedure broadly followed their source (Patterson et al., 1987; Whitlock and Larsen, 2001). the protocol described in Carcaillet et al. (2001, 2007). One cm3 of Additionally, the CHAR of sedimentary charcoals larger than sample was treated with a solution of NaClO (15%) and KOH (5%) 0.25 mm2 was also plotted to better distinguish the local fire signal (Finsinger et al., 2014). Samples were heated at 70  C during 90 min (Whitlock and Larsen, 2001; Finsinger et al., 2014), as was previ- with a magnet inside the beaker that moved following an electro- ously done by Perez-Obiol et al. (2016). magnetic field to facilitate the reaction. When cooled, samples were sieved through a 150 mm mesh under a water jet. The remaining charcoal particles (>150 mm) were counted using 3. Results a stereomicroscope at a magnification of 40x. The surface area of each charcoal was measured with an ocular grid of 100 squares, 3.1. Sedimentary description each 0.0625 mm2 (Rhodes, 1998; Carcaillet et al., 2001). The char- coal particles were classified into size-classes that increased Perez-Obiol et al. (2016) described three stratigraphic phases in exponentially. the MOL2 record for the interval between 0 and 260 cm (Fig. 4). The In a similar way to the organic matter and palynological se- shallowest layer (0e40 cm) was described as the acrotelm. It was quences, a single charcoal sequence (MOL2/MOL6) was obtained by composed of light brown material (10 YR e 3/6, according to the joining 288 charcoal samples analysed in Pe rez-Obiol et al. (2016) Munsell colour chart), containing living Sphagnum spp. in the upper 6 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 Fig. 4. Lithological structure of the MOL2 and MOL6 records and organic matter content of the MOL2/MOL6 record from La Molina peat bog. The MOL2/MOL6 record (on the right) was obtained by the combination of the top 250 cm of the MOL2 record, analysed in Pe rez-Obiol et al. (2016), and the interval from 250 to 499 cm of the MOL6 record. 5 cm. The organic values ranged from 55 to 87%. A deeper level 3.2. Chronological model (40e250 cm), identified as the catotelm, was composed of dark brown gyttja (10 YR e 2/2 and 10 YR e 2/1), yielding high organic The age-depth chronological model spans the last c. 17,550 cal yr values ranging from 78 to 95%. Within the deepest part of MOL2 BP and supports continuous sedimentation (Fig. 5). The 210Pb pro- (250e260 cm), a progressive decrease in organic matter was file indicated that the top 20 cm of the peat bog was not altered. The detected with a greater presence of silts, followed by an organic sedimentary accumulation rate, with a mean value of 0.028 cm pulse. This oscillation was also captured by the MOL6 record, which yr1, revealed two sectors with the highest chronological resolution was followed by a series of pulses until 373 cm depth. Peaks of (the 20th century and the period comprised between the Neolithic inorganic material were detected at 6230, 7090, 8210 and and the Chalcolithic, the latter registering a maximum at 9350 cal yr BP. In particular, from 342 to 369.5 cm depth (from 4590 cal yr BP), with peaks of 0.5 and 0.15 cm yr1, respectively. 10,130 to 11,760 cal yr BP) the organic content was notably lower and ranged between 76 and 55%. An organic increase was detected 3.3. Pollen from 369.5 to 373 cm (from 11,760 to 12,010 cal yr BP), prior to the transition towards a lacustrine material. Along this deepest phase, The diagrams of pollen percentages and pollen accumulation the colour progressively changed from dark brown (10 YR e 2/1) to rate (PAR) obtained from the MOL2/MOL6 record of La Molina peat grey (10 YR e 4/1) tones. bog (Figs. 6 and 7) permitted the landscape reconstruction of the At 373 cm depth (12,010 cal yr BP) an abrupt lithological change area for the last c. 17,550 cal yr BP. According to the CONISS, four in a short interval was detected. The organic content sharply pollen assemblage zones (PAZ) and seven subzones were identified. decreased and remained at low values until the bottom of the core, From 17,550 to 16,070 cal yr BP (MOL/1a), non-arboreal pollen even though some organic pulses were registered with values values moved around 85e90% (Fig. 6), while the PAR were minimal ranging from 4 to 36%. The organic peaks were detected at 13,410, (Figs. 7 and 9). Poaceae clearly dominated the landscape, regis- 14,870, 16,320 and 16,650 cal yr BP. The material was essentially tering a PAR between 300 and 700 grains cm2 yr1 and percent- made up of lacustrine grey clays (10 YR e 4/1) until the end of the ages around 50%. There was also presence of xerophytes such as record, exhibiting a more flexible and plastic structure than the Artemisia (7e18%), Amaranthaceae (<5%) and Ephedra (<5%), while peat material from shallower layers. other herbs and shrubs registered low values: Calluna vulgaris, 7 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 Fig. 5. Dates, smoothed age-depth chronological model and sedimentary accumulation rate of La Molina peat bog. The age-depth model included radicarbon dates (n ¼ 4) analysed in the MOL2 record by Pe rez-Obiol et al. (2016) and radiocarbon dates (n ¼ 10) analysed in the MOL6 record in this paper, all of them calibrated using the IntCal20 Northern Hemisphere radiocarbon age calibration curve (Reimer et al., 2020). Measurements of 210Pb were performed from 0 to 20 cm depth applying a constant rate of supply model (Appleby and Oldfield, 1978). Asteroideae, Cichorioideae, Plantago, Rubiaceae, Helianthemum and this line, Betula signal represented the greatest portion of the Centaurea. Pinus (5e10%), Ericaceae (<5%) and Juniperus (<5%) arboreal pollen percentages, which revealed an interval of sharp stood out over the woody species which appeared interruptedly. fluctuations along the Bølling-Allerød chronozone (GI-1) and the Concerning the local vegetation, there was a great presence of hy- Younger Dryas (GS-1). The great quantity of Betula also triggered drophytes, hygrophytes (Myriophyllum alterniflorum, Potamogeton, the first significant peak of pollen influx, detected at 13,260 cal yr Ranunculaceae, Cyperaceae, Apiaceae, Potentilla-type and Typha/ BP (Figs. 7 and 9). After this period, Betula progressively decreased Sparganium) and algae (Botryococcus). Ranunculaceae, probably an down to 12% during the last stages of the LGP (13,370e11,900 cal yr aquatic buttercup, were the best represented among the excluded BP), resulting in an increase of Pinus (20%) and non-arboreal per- taxa, with a 50e70% over the total taxa until 16,250 cal yr BP when centages (65%). Also particularly significant were the short- their PAR sharply decreased to be surpassed by Cyperaceae duration pulses of Poaceae (39%), Cichorioideae (28%), Aster- percentages. oideae (10%), Centaurea (8%) and Filipendula cf. F. ulmaria (10%) The period from 16,070 to 13,700 cal yr BP (MOL/1b) started registered prior to the onset of the Holocene. with a progressive Poaceae increase reaching a PAR greater than During the first millennia of the Holocene (MOL/2b), peaks of 1000 grains cm2 yr1 at 14,900 cal yr BP, representing 85% of the terrestrial PAR were detected at 11,600, 11,200, 10,130 and terrestrial pollen. Following the opposite trend, Artemisia decreased 9600 cal yr BP, the latter being the greatest of all the sequence (up from 20 to 15% to less than 5% and Juniperus values became mini- to 60,000 grains cm2 yr1; Fig. 9). The pollen influx of Pinus, Betula, mal. After 14,900 cal yr BP, a series of pollen fluctuations was evergreen Quercus, deciduous Quercus and Salix increased, and the detected. Poaceae oscillated in accordance with organic matter values of Betula (>32,000 grains cm2 yr1) and deciduous Quercus pulses while Juniperus recovered its minimal presence between (>14,000 grains cm2 yr1) were significantly relevant. Betula 14,700 and 13,960 cal yr BP. In addition, particularly noteworthy registered mean percentages between 50 and 70%, deciduous were also the peaks of Sanguisorba officinalis at 14,550 cal yr BP Quercus between 20 and 40% and Pinus around 5%, yielding arboreal (23%) and Ericaceae at 13,900 cal yr BP (51%). Overall, the arboreal pollen percentages above 90%. The terrestrial pollen concentration pollen was still low (10%), with similar values of Pinus and a slight was also the greatest detected in all the sequence, with peaks up to increase in deciduous Quercus. At a local scale, a Sordaria pulsation 2 $ 106 grains cm3. Pteridium-type also stood out during the first (25%) was registered between 16,070 and 14,600 cal yr BP. The stages of the Holocene, with values around 56% at 11,900 cal yr BP. presence of aquatic taxa decreased during MOL/1b. Later on, the beginning of the Corylus continuous signal was Between 13,700 and 11,900 cal yr BP (MOL/2a), corresponding to detected at 10,680 cal yr BP, which increased progressively to reach the end of the Würm glaciation, Betula percentages sharply more than 10% at 9720 cal yr BP. A decrease of Betula percentages in increased and represent 81% of pollen at 13,400 cal yr BP. The Salix favour of deciduous Quercus was also identified between 10,100 signal increased during this PAZ too, with values close to 10% and 9850 cal yr BP. around 13,000 cal yr BP. It is also worth stressing the increase of the PAZ MOL/3 (9300e4860 cal yr BP) started with a sharp decrease Pinus influx up to 1050 grains cm2 yr1, even though its per- of Betula at 9300 cal yr BP. Apart from a pulsation of 25% at centage did not vary due to the great quantity of Betula pollen. In 9100 cal yr BP, Betula stood at around 10% throughout the period. 8 Fig. 6. Diagram of pollen percentages of La Molina peat bog (MOL2/MOL6 record). Light silhouettes indicate x5 exaggeration. The organic matter content (%) is shown as a red line and the charcoal accumulation rate (CHAR, pieces cm2 yr1) of charcoal particles >150 mm is shown as a grey area. Pollen assemblage zones (PAZ) are also shown. (For inter- pretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.) 9 Fig. 7. Diagram of pollen accumulation rates (grains cm2 yr1) of La Molina peat bog (MOL2/MOL6 record). Light silhouettes indicate x5 exaggeration. The organic matter content (%) is shown as a red line and the charcoal accumulation rate (CHAR, pieces cm2 yr1) of charcoal particles >150 mm is shown as a grey area. Pollen assemblage zones (PAZ) are also shown. (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.) 10 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 Deciduous Quercus also registered a lower PAR than in MOL/2b, suggesting that charcoal peaks were confidently separated from despite not being reflected in the percentage diagram mainly due to noise (Kelly et al., 2011). From 8400 to 5700 cal yr BP, the charcoal Betula's significant decline. Both decreases, however, led to the frequency decreased with a mean CHAR value at around 0.1 pieces greatest percentages of Corylus along all the sequence, varying cm2 yr1. The CHAR value started increasing again at 5700 cal yr between 35 and 45%. Between 8330 and 8180 cal yr BP, an impor- BP and was followed by the greatest fire peak of all the sequence tant decrease in the Corylus and deciduous Quercus influx led to a registered at 5020 cal yr BP (834 pieces cm2 yr1). Subsequently, Pinus increase up to 27%. Also, the beginning of the Ulmus, Alnus and the CHAR value stood at up to 28 pieces cm2 yr1 between 4650 Fraxinus continuous curves was found at 8800, 7200 and and 4430 cal yr BP. Also, the charcoal influx of particles larger than 5000 cal yr BP, respectively. A low pulsation of Vitis was detected 0.25 mm2 reached 0.85 pieces cm2 yr1 within these peaks. After between 8180 and 8040 cal yr BP and the first appearances of the event of 4430 cal yr BP, however, the charcoal values and the Cerealia and Olea were estimated at around 6660 cal yr BP. Espe- peak magnitude started to decline, registering the last significant cially significant is the detection of a single instance of Fagus pollen fire episode at around 990 cal yr BP. The charcoal curves displayed a at 8620 cal yr BP and another of Carpinus at 8840 cal yr BP. Equally, low signal during the most recent period (MOL/4b, from 1900 the non-arboreal spectra expanded due to the Poaceae increase to 63 cal yr BP), with peaks below 65 pieces cm2 peak1 that (25e35%) as the main taxon of the local openings. Pteridium-type were not well separated from noise in agreement with the signal- and Gentiana pneumonanthe signal also increased mainly within to-noise index (Kelly et al., 2011). this PAZ. Between 5270 and 4100 cal yr BP, corresponding to the late 4. Discussion stages of MOL/3 and the first centuries of MOL/4a, the greatest values of arboreal pollen influx were registered, particularly for To date, there have been few studies examining what the plant Pinus, Betula, evergreen Quercus, deciduous Quercus, Corylus, Ulmus, landscape of the Cantabrian region was like during the Last Glacial Alnus, Fraxinus and Salix. As indicated by Pe rez-Obiol et al. (2016), Period (LGP). The multiproxy approach performed in La Molina during the time period encompassed by MOL/4a (4860e1900 cal yr (484 m a. s. l.) is the oldest chronology studied on the northern BP), Corylus and deciduous Quercus continued to express high slope of the Cantabrian range and made it possible to determine the values. However, the non-arboreal composition shifted as Poaceae succession of different landscape units over the last c. 17,550 cal yr decreased in benefit of Ericaceae. At the same time, the signal of BP. Pteridium-type also became minimal. Low values of Assulina, a testate amoeba, were registered from 4560 cal yr BP together with 4.1. An open landscape after the Last Glacial Maximum an intermittent but continuous signal of coprophilous fungi, mainly Sordaria and Sporormiella. A steppe-like formation with scarce tree cover is deduced from From 1900 to 63 cal yr BP (MOL/4b) a falling arboreal pollen the pollen spectra during the Greenland Stadial 2.1a (from 17,550 to trend was observed until the onset of the 20th century. Alnus and 14,700 cal yr BP; Rasmussen et al., 2014). This phase, which also Fagus stood out among the tree species, coinciding with the encompasses the Heinrich event 1 (Heinrich, 1988; Bond et al., decrease of Corylus and deciduous Quercus percentages. According 1992; Hemming, 2004), was characterised on the northern slope to the updated chronological model, Alnus reached the highest of the Cantabrian range by cold and dry climate features until percentages at 1820 cal yr BP, and the onset of the continuous signal 15,500 cal yr BP, inferred through isotopic analyses on a speleothem of Fagus is estimated at 1680 cal yr BP. from the nearby Asiul cave (Moreno et al., 2010a). Accordingly, the The decline of Corylus and deciduous Quercus coincided with plant landscape fully reflects the climatic severity of that time with high values of Poaceae and Ericaceae PAR, leading to increased non- limited water availability which did not allow either the tree or arboreal pollen percentages. An important peak of Pteridium-type shrub cover to develop. The dominance of Poaceae over Artemisia was also recorded coinciding with the greatest non-arboreal per- and other xerophytes is consistent with the plant composition centages. During the 20th century, however, the decreasing arbo- documented in the littoral and coastal mountain sites from the real trend shifted as a result of a significant Pinus influx increase. In north-western Iberian Peninsula in Portugal (Go mez-Orellana et al., addition, Olea, Platanus, Eucalyptus and Castanea reached a 2001, 2013), Galicia (Saa-Otero and Víaz-Fierros, 1988; Go  mez- continuous signal, yielding lower percentages of deciduous Quer- Orellana et al., 1998, 2001; Lo  pez-Merino et al., 2012; García- cus, Corylus, Fagus and Alnus while maintaining a similar pollen Moreiras et al., 2019) and Asturias (Lo  pez-Merino, 2009). From influx. The coprophilous signal was similar to the preceding PAZ, analyses of modern surface samples (mosses), it can be deduced now with pulsations of Sordaria and Podospora instead of Spor- that the open landscape could be equivalent to the current alpine ormiella. An increase in Botryococcus and Assulina was detected grasslands located above the treeline in the European Alps among the aquatic taxa from 500 cal yr BP onwards. (2360 m a. s. l.), which are communities that are certainly well- adapted to water scarcity and cold temperatures (Furlanetto et al., 3.4. Sedimentary charcoals (>150 mm) 2019). A greater proportion of Mediterranean xerophytes has been detected in Iberian sites with a greater Mediterranean influ- The interpretation of the fire dynamics was based on the same ence during the LGP (e.g., Pantale on-Cano et al., 2003; Camuera pollen assemblage zones identified within the pollen diagrams. The et al., 2019), suggesting drier climate features than in the Euro- charcoal signal displayed low values from 17,550 to 13,700 cal yr BP siberian biogeographical region. (MOL/1), with charcoal peaks with less than 10 pieces cm2 yr1 At 15,500 cal yr BP, a shift to warmer and wetter conditions is (Figs. 8 and 9). Between 13,700 and 9300 cal yr BP (MOL/2), the fire detected in the Cantabrian region (Moreno et al., 2010a). This evidence was still low, even though the first particles larger than climate change ties in well with the decrease of xerophyte taxa 0.25 mm2 were detected. The first significant charcoal peaks from 16,000 cal yr BP onwards. At a local scale, the loss of lacustrine occurred at 9300 cal yr BP (206 pieces cm2 peak1) and 8650 cal yr features and the transition to an early stage of a peat bog has also BP (128 pieces cm2 peak1). The local component of the two fire been noted, as indicated by the decrease of hydrophytes, hygro- events is supported by the presence of charcoal particles larger phytes and algae. Here the Sordaria pulsation detected between than 0.25 mm2. Additionally, most charcoal peaks coincide with a 16,070 and 14,600 cal yr BP might indicate a higher amount of singal-to-noise index above or close to 3 from that point onwards, decaying organic matter. 11 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 Fig. 8. Diagram of the charcoal concentration (CHAC), charcoal accumulation rate (CHAR), fire peaks as red dots, fire peak magnitude and charcoal accumulation rate of charcoal particles larger than 0.25 mm2 of La Molina peat bog (MOL2/MOL6 record). The organic matter is shown as a red line, the pollen accumulation rate (arboreal: AP þ non arboreal: NAP) as a green line and the signal-to-noise index (SNI) as a blue line. The cutoff value of SNI ¼ 3 (Kelly et al., 2011) is indicated as a blue dashed line. Pollen assemblage zones (PAZ) are also shown. (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.) At a regional scale, the landscape described on the northern trees, mainly Pinus and Betula, in La Piedra (Ramil-Rego et al., 1998) slope differs from what has been documented in other sectors of and San Mame s de Abar (Iriarte-Chiapusso et al., 2001), in what is the Cantabrian range, which seems to have played a significant role likely to be the last stages of the GS-2.1a (Rasmussen et al., 2014). in the geographical distribution of certain species. On the one hand, Unfortunately, the absence of a robust chronology did not allow glaciers expanded from 22,500 to 18,000 cal yr BP at higher altitude accurate identification the climate episodes, though it is possible to areas of the mountain system (Frochoso et al., 2013; Ruiz- recognise different behaviour from the Cantabrian oceanic slope. Fernandez et al., 2016). Accordingly, the vegetation community Moreover, the dominance of Pinus becomes more evident if we look within this altitudinal range appears to have been sensitive to more further south. It is the case of the northern Iberian System, where arid and colder conditions, since steppe-like taxa displayed greater several sites reveal vegetation communities with a great presence values than on the Atlantic façade from the Last Glacial Maximum of pines at certain stages of the LGP (Pen ~ alba, 1994; Gomez-Lobo (from 26,500 to 19,000 cal yr BP), both on the westernmost flank et al., 1996; Pen ~ alba et al., 1997; Von EngerIbrechten, 1998; (Aira-Rodríguez, 1986; Santos-Fidalgo et al., 1997; Mun ~ oz-Sobrino Sanchez-Gon ~ i and Hannon, 1999; Gil-García et al., 2002), reaching et al., 2001) and also in the central sector (Ruíz-Zapata et al., pollen values up to 70e90% in Laguna Grande (Ruiz-Zapata et al., 2000; Jime nez-Sanchez et al., 2003). The community composition 2002). These forest communities contrast with the open grass- did not change during the immediately preceding millennia in the lands from the Atlantic coastal region. It is likely that the conti- mountain environment as recorded, from West to East, in Laguna nental conditions of the southern slope, which implies higher de Lucenza (Aira-Rodríguez, 1986; Santos-Fidalgo et al., 1997; maximum temperatures or at least more water availability, may Mun ~ oz-Sobrino et al., 2001), Villaseca de Laciana (Jalut et al., 2010), have facilitated the development of the tree cover in the inland La Mata (Jalut et al., 2010), Lago de Ajo (Allen et al., 1996) and, closer region. On the northern slope, anthracological analyses performed to La Molina, in Puertos de Ríofrio (1700 m a. s. l.), which displays in caves reported high abundance of pine macroremains during the 30% of Artemisia during the Late Glacial (Mene ndez-Amor and Upper Pleistocene (Baena et al., 2005). However, there is no evi- Florschütz, 1963). In the latter site, although only a few pollen dence of large extensions of pinewoods in the region around La samples were analysed from the Late Glacial, the significant pres- Molina, at least from the Magdalenian to the present. The pine ence of Artemisia may indicate that the arid conditions were values are not only far below the documented percentages in maintained in the upper reaches of the mountain system for a southern areas, but also modern pollen rain calibration attributed longer period of time than in La Molina. more than 50e60% of Pinus to pinewoods in the Central System On the other hand, the situation was very different on the (Lo pez-S aez et al., 2013), which is substantially higher than the southern slope. In northern Burgos, there was a greater presence of Pinus signal documented in La Molina. The lack of large extensions 12 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 Fig. 9. Comparison of CHAR, Ice Rafted Debris (IRD) index: Stack of MC52-V29191 þ MC21-GGC22 (Bond et al., 2001), pollen accumulation rate (arboreal: AP þ non arboreal: NAP), organic matter and d18O from the NGRIP ice core, adapted from Rasmussen et al. (2014). At the top of the figure, the chronology for the Würm glaciation based on Greenland Stadials (GS) and Greenland Interstadials (GI) is shown (Rasmussen et al., 2014), and also the stages for the Holocene based on Walker et al. (2018, 2019). The Younger Dryas Stadial (YD) and the substages of the Greenland Interstadial 1 (aee) are indicated. of pines is also supported by the low pollen values recorded in the 2006; Iriarte-Chiapusso et al., 2006; Lopez-Merino et al., 2006, vast majority of Holocene pollen records along the Cantabrian 2010; Lopez-Merino, 2009; Gonza lez-Pellejero et al., 2014). ~ alba, 1994; García-Anto littoral (e.g., Mariscal, 1993; Pen n et al., Without ruling out the possible existence of scattered pine 13 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 individuals, the origin of the low but continuous Pinus signal level, but in both cases their presence reflects an environmental recorded in La Molina could be explained by the recognised large change from 13,900 cal yr BP onwards. pollen dispersal of this species (Poska and Pidek, 2010) and the long At the same time, Betula led the succession to a more forested transport from southern regions where Pinus populations were environment. Salix exhibited a similar behaviour, although its more consolidated. signal is substantially lower than that of Betula perhaps because it Concerning the role of fire, the lack of woody species went hand has a certain degree of zoophily (Lara-Ruiz, 2019) and its pollen in hand with the absence of significant fire events during the dispersal may be lower. In any case, the expansion of both taxa, Palaeolithic section of the record. The lack of charcoal particles which represented the first significant biomass pulsation of the larger than 0.25 mm2 during that period also reinforces the idea sequence, is explained by a gradual humidity increase that took that no significant fires occurred at a local level, since a larger size place in the region between 15,400 and 13,400 cal yr BP (Moreno would indicate a closer origin (Whitlock and Larsen, 2001; et al., 2010a). Betula and Salix started increasing during that inter- Finsinger et al., 2014). We need to look beyond the Cantabrian re- val and Betula displayed its maximum values at 13,370 cal yr BP, gion to compare La Molina's fire regime during the LGP, since the having likely benefited from increased water availability. From that available charcoal records only cover the period up to c. 7000 cal yr point onwards, Betula suffered a first decreasing pulse in what BP (Carracedo et al., 2018). Thus, the lack of fires registered in La might have been a response to the cold spell registered from 13,310 Molina before the Holocene is in line with other Iberian sites with to 13,100 cal yr BP in Greenland ice cores (GI-1b; Rasmussen et al., an Eurosiberian distribution, such as Charco da Candieira (Portugal; 2014). The decline of birches and willows was consolidated within Connor et al., 2012), El Portalet (Central Pyrenees; Gil-Romera et al., the Younger Dryas, a dry and cold phase of regional extent 2014) and also in the Mediterranean site of Siles (Andalucía; (Rasmussen et al., 2014) well characterised in north-western Iberia Carrio n, 2002) where the fire signal was weak during the LGP. The (Moreno et al., 2010a; Mun ~ oz-Sobrino et al., 2013; Baldini et al., low intensity of fires meant it was not possible to explore any 2015, 2019). In particular, at the onset of the Younger Dryas, a se- possible connexion with an environmental signal during that vere drop in temperatures of between 6 and 9  C was inferred from period (e.g., the d18O curve from Greenland ice cores: Dansgaard La Garma cave (Baldini et al., 2015, 2019) at about 40 km from La et al., 1982; Johnsen et al., 1992; Rasmussen et al., 2014; Seierstad Molina. The study also provided climatic models that revealed et al., 2014). especially cold and dry winters for that period. The increasing Turning to the relationship between humans and the Cantabrian influx of non-arboreal species, then dominated by Poaceae and landscape, anthropic pressure does not seem to have been signifi- accompanied by Asteroideae, Cichorioideae and Centaurea, is a cant during the LGP. On the contrary, the plant formations could good reflection of the tough conditions prior to the Holocene. Pinus, have affected the distribution of some animal species, the prey of on the other hand, does not seem to have been significantly affected the hunter-gatherer human groups that inhabited the Cantabrian by the climatic fluctuations, exhibiting less variability than Betula. range (e.g., Solo rzano et al., 1999; Gonzalez-Sainz and Gonza lez- Betula's fluctuations were not an isolated dynamic detected in La Urquijo, 2004) and other hunting animals. In particular, the Ibe- Molina peat bog, as they were also captured in the western deposits rian ibex (Capra pyrenaica) was the principal prey documented in of Laguna de Lucenza (Aira-Rodríguez, 1986; Santos-Fidalgo et al., north-Iberian archaeological sites far from the current shoreline 1997; Mun ~ oz-Sobrino et al., 2001), Villaseca de Laciana and La (>9 km) and above 200 m from the current sea level, as is the case Mata (Jalut et al., 2010), indicating a climatic response at a regional of La Molina. On the other hand, remains of red deer (Cervus ela- scale. Similarly, these oscillations have also been detected in the phus) were the most abundant fossils found close to the shoreline Pyrenees during the same period, although with a greater presence (Portero et al., 2019). Thus, the non-forested landscape inferred of Pinus (Reille and Lowe, 1993; Cunill et al., 2013; Gil-Romera et al., from La Molina's pollen spectra seems to have been a suitable 2014). habitat for the Iberian ibex, which is currently found in moun- The lack of taxonomical resolution does not allow interpreting tainous areas and has a diet mainly made up of herbaceous plants which birch species undertook the woody colonisation. At present, (Granados et al., 2001). The steppe-like formation was otherwise Betula pubescens is the birch species present in the mountainous probably a limiting factor in the distribution of red deer, whose area, while Betula pendula is not so frequent in the Cantabrian range principal habitat is woodlands and open areas close to woody for- but has a wide distribution in north-eastern Iberian mountains mations as they are often used as a refuge (Carranza, 2017). The red (Pyrenees) and Europe. We cannot rule out these species having led deer distribution might therefore suggest a more forested envi- the colonisation since they are recognised for their ability to colo- ronment close to the coastline, which perhaps also occupied a nise open areas, such as heathlands (Shaw et al., 2014a; Stritch wider area since the sea level was more than 100 m lower than at et al., 2014). In this line, a significant exploitation of Betula's present (Lambeck et al., 2014; Spratt and Lisiecki, 2016). However, wood has been documented in the littoral cave of La Pila during the for now there is a lack of palynological records covering this period late Magdalenian and early Azilian: from c. 12,500 to 11,700 yr BP on the Cantabrian littoral plain to support this hypothesis. (Bernaldo de Quiro s et al., 1992; Uzquiano, 2014). Such a large amount of wood might suggest the presence of arboreal Betula 4.2. The spread of woody species over the Bølling-Allerød species. Alternatively, shrubby birches such as Betula nana or Betula chronozone and their decline during the Younger Dryas humilis, both widely distributed across Europe at present (Beck et al., 2016), could have also developed in the Cantabrian area. The period from 14,690 to 12,900 cal yr BP (GI-1) is charac- Considering their habitat, Betula nana is an arctic or high mountain terised by being an overall mild phase with a certain degree of plant of immature or peaty soils, found in alpine or subalpine en- climate variability (Rasmussen et al., 2014). Our results indicate vironments (Stritch, 2014). Betula humilis can be found as part of several vegetation fluctuations for that period, very different from different communities from the hill to the montane zone, both in the little-disturbed landscape that remained after the LGM. A first acidic and wet forest soils and also close to wetlands (Shaw et al., vegetation shift occurred at the beginning of the Bølling-Allerød 2014b). Thus, both the arboreal and shrub birch species Interstadial in the form of Juniperus and Ericaceae pulses accom- mentioned above could fit well into the pre-Holocene Cantabrian panied by a decrease of Poaceae in what can be interpreted as a landscape. In the case of Salix, it would be reasonable to think of weak vegetation response to the warmer climate features. Unfor- low shrub or creeping dwarf willows. Future analyses on the tunately, Juniperus and Ericaceae could not be identified to species vegetation macroremains or ancient DNA would help to identify 14 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 Ericaceae, Salix and Betula to a more specific level, thus shedding afterwards due to an aridity increase detected in the region from more light on how this succession occurred. 8.5 to 8.0 ka onwards (Rossi et al., 2018). This environmental shift, together with the local fire episodes registered during the 9.3 and 4.3. The establishment of mixed deciduous formations during the 8.2 ka events, may have favoured Corylus expansion, thus coin- early Holocene ciding with the hypothesis of Huntley (1993) and Finsinger et al. (2006). The early Holocene was characterised by a general warming and The pollen record also revealed a very occasional presence of humidity increase in the north-western Iberian Peninsula (Smith hornbeam and beech during the early Holocene. In regard to Car- et al., 2016; Rossi et al., 2018; Morello n et al., 2018; Baldini et al., pinus betulus, an isolated pollen grain was detected at 8840 cal yr 2019), coinciding with the greatest Holocene values of summer BP. Such minimal presence has also been documented during the insolation in the Northern Hemisphere (Berger and Loutre, 1991). early and mid-Holocene in nearby littoral sites (Mary et al., 1975; Although most rainfall was concentrated during the winter months Iriarte-Chiapusso et al., 2006, 2016) and mountainous areas in the Cantabrian range (Baldini et al., 2019), the humidity increase (Mene ndez-Amor and Florschütz, 1963; Pen ~ alba, 1989), while was enough to support the rapid expansion of forests at the onset of slightly more developed populations were settled along the west- the Holocene. Additionally, the presence of woody species during ern Galician coast (Mun ~ oz-Sobrino et al., 2018). In several sites in the Younger Dryas, mainly Betula, would have favoured soil northern Iberia, the scattered presence of hornbeam trees repre- development and is another important factor in explaining how sented a vestige from a larger distribution achieved during the fast the arboreal succession occurred. The sharp biomass increase Würm glaciation (e.g., S anchez-Gon~ i et al., 2005, 2008; Go  mez- suggests a different temperature threshold from higher altitude Orellana et al., 2007, 2013). However, C. betulus was not detected areas at the alpine Pyrenees, where colder temperatures postponed in the glacial interval of La Molina, probably due to the cold and arid the increase of the PAR by a few millennia (Leunda et al., 2017). conditions deduced from the steppe-like landscape, very far from During the Lateglacial-Holocene transition, Azilian human the ecological requirements of this mesophyte species. In the groups were settled at high altitudes of the Cantabrian range aforementioned sites where it occupied a wider extension, the rise (Barandiar an et al., 2006). The Cantabrian landscape, however, of the sea level, which flooded coastal regions where it was seems to have remained little affected by anthropic pressure during established, together with the post-glacial expansion of more that time. Plant biomass, mainly represented by tree species competitive arboreal species in inland sites reduced its distribution (>95%), exhibited the maximum values of the entire sequence, area and relegated it to a very marginal role (Mun ~ oz-Sobrino et al., whereas human signals were not documented in the palynological 2018). Besides, landscape anthropisation did not benefit its pop- record. Conversely, plant production seems to have been sensitive ulations which still have a very scattered presence in the Iberian to a series of century-to-millennial scale environmental pulses Peninsula at present (Costa-Tenorio et al., 2005). triggered by changes in the North Atlantic Ocean circulation which In regard to Fagus, an isolated pollen grain was detected at have been well recorded in isotopic records on ice cores (Bond et al., 8624 cal yr BP. This finding, although it cannot indicate the estab- 2001; Rasmussen et al., 2007, 2014). In the Cantabrian range, such lishment of beech populations at a local level, is consistent with the climate variability entailed cyclical shifts of wetting and drying discontinuous and low pollen curves detected in other sites from conditions, with a periodicity of 1290 years during the early Ho- the eastern Cantabrian range and the Basque mountains during the locene (Smith et al., 2016). The frequency of biomass declines in La early Holocene, such as La Nava (870 m a. s. l.; Mene ndez-Amor, Molina was very similar to that of humidity shifts during that 1968), La Piedra (950 m a. s. l.; Ramil-Rego et al., 1998), Belate period, revealing a close relationship between climate conditions (825 m a. s. l.; Pen~ alba, 1994), Santa Gadea (837 m a. s. l.; Iriarte- and vegetation growth with reduced productivity during the driest Chiapusso et al., 2003), Puerto del Escudo (940 m a. s. l.; Mun ~ oz- intervals. Sobrino, 2001) and Puerto de los Tornos (920 m a. s. l.; Pen ~ alba, Concerning the vegetation communities, the Cantabrian forests 1994; Mun ~ oz-Sobrino et al., 2005), all of them located at a higher were mainly composed of Betula, deciduous Quercus and a scattered altitude than La Molina (484 m a. s. l.). Thus, a plausible hypothesis presence of Salix, Pinus and evergreen Quercus. The signal of de- for the origin of the isolated beech pollen found in La Molina would ciduous Quercus might mainly correspond to Quercus robur, since it be the long transport from higher altitude regions, where there was is the most common oak in the inner valleys of the current land- greater evidence of Fagus. Equally, it might also be held that small scape. Corylus, on the other hand, started to expand some centuries glacial refuges from the Cantabrian range (Magri et al., 2006) could later, in line with other montane and subalpine locations of the have contributed to this pollen presence. The existence of these Cantabrian range (Mene ndez-Amor and Florschütz, 1963; ancient refuges in the region is supported by the fossil evidence Mene ndez-Amor, 1968; Allen et al., 1996; Lo pez-Merino, 2009; during the glacial period, which has been found in the form of Jalut et al., 2010; Mun ~ oz-Sobrino et al., 2012). This is also consistent pollen (Mene ndez-Amor, 1968; Pen ~ alba, 1989, 1994; Ramil-Rego with the hazel's dynamics in south European regions, where its et al., 1998; Iriarte-Chiapusso et al., 2016) and macroremains expansion occurred later than the spread of other deciduous taxa (Uzquiano, 1992). The sequence of La Molina, however, does not (Finsinger et al., 2006; Giesecke et al., 2011), contrary to what provide evidence of beech before the Holocene, since in the same happened in northern and central Europe (e.g., Huntley, 1993; way as with hornbeam the climatic conditions do not seem to have Tinner and Lotter, 2001; Tallantire, 2002). Huntley (1993) suggests favoured Fagus development in the coastal mountains of the Can- that high seasonality was an important factor in the development tabrian range. In a more recent period, Fagus sylvatica appeared in of hazel populations at high latitude regions, probably also linked to the area near La Molina at around c. 2800 cal yr BP. Pe rez-Obiol increased fire activity. Indeed, Corylus is a resprouting taxon well et al. (2016) point out that fire could have facilitated its expan- adapted to fire events (Delarze et al., 1992; Tinner et al., 1999). sion along the Cantabrian range during the last millennium. Finsinger et al. (2006) underlined similar reasons for the late spread of Corylus in the Southern Alps. In the Cantabrian region, the warm 4.4. Implications of the 9.3 and 8.2 ka events on the Cantabrian and humid climate conditions during the early Holocene (Smith landscape et al., 2016; Rossi et al., 2018; Morello n et al., 2018; Baldini et al., 2019) and the lack of fire activity during that period may have Apart from the decline in biomass production, the early Holo- prevented an earlier hazel expansion. The situation changed shortly cene climate pulses (Smith et al., 2016) were not accompanied by 15 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 other significant ecological implications until the 9.3 and 8.2 ka aforementioned early Holocene climate pulses of wetting and events, when two fire episodes of high magnitude occurred. Both drying conditions. Pines were the least affected species, leading to a events are known to have had a regional scope (Rasmussen et al., percentage increase especially well-recorded within the 8.2 ka 2007, 2014), coinciding closely with the Bond event 6 and the event. Its lesser impact may be due to the fact that its pollen signal first peak of ice-rafted debris within the Bond event 5, respectively could have come from a regional area given its large pollen (Fig. 9; Bond et al., 2001). In the Cantabrian range, these climatic dispersion (Poska and Pidek, 2010), and thus being less affected by phases have been recorded as isotopic oscillations in cave speleo- the local fires detected in La Molina. Betula, on the other hand, was thems (Domínguez-Villar et al., 2009; Moreno et al., 2010a; Smith seriously affected, and its presence became minimal after the 9.3 ka et al., 2016; Rossi et al., 2018; Baldini et al., 2019). In a similar fire event. The fire episode created forest clearings dominated by way, the organic matter record of La Molina, which has proved to be grasses and Pteridium, and in this scenario hazels could colonise an efficient proxy to reflect environmental changes (Meyers and and develop well to become important components of mixed de- Lallier-Verges, 1999), also captured these intervals in the form of ciduous formations which have dominated the landscape until the falls in organic content. present. Although Corylus is an important element in the secondary The climate interpretation performed in Cantabria suggests a succession of many temperate forests (Hegi, 1981; Carreras et al., relatively humid phase between 9900 and 9700 cal yr BP (Rossi 2016), it should be noted that its expansion during the mid- et al., 2018). Accordingly, the PAR exhibited the highest values of Holocene did not merely represent a secondary colonisation after all the record, just before the first fire episode. During the 9.3 ka fire episodes, but rather the species was part of a stable community event, temperatures rose and summers became drier in the Can- that remained in equilibrium for several millennia. tabrian range (Baldini et al., 2019). The 8.2 ka event was also Although the species succession appears to have been logical, characterised as a dry interval close to La Molina (Domínguez-Villar the arboreal biomass did not regain previous values during the et al., 2009; Smith et al., 2016; Rossi et al., 2018), even though it was Mesolithic or increase following climatic wetter pulses (Smith et al., one of the coldest pulses of the entire Holocene (Baldini et al., 2016) as it did during the early Holocene. This situation leads us to 2019). It has been shown that climate changes of this magnitude believe that vegetation was no longer solely influenced by climate can cause major fires in a brief period of time (Daniau et al., 2019) features, but rather the anthropic factor could have begun to since both temperature and humidity are acknowledged to be contribute significantly to the configuration of the landscape dur- important factors in controlling biomass burning on a regional scale ing this period. Several Mesolithic groups were settled in the (Daniau et al., 2012). Additionally, Power et al. (2008) also point out Cantabrian littoral and mid-mountain areas up to 750 m a. s. l., that increased seasonality during the early Holocene could have some of them documented in La Molina valley (Pe rez-Bartolome , regulated the fire regime in the Northern Hemisphere. Therefore, 2019). It is known that Corylus was exploited by these Mesolithic the climate pulses recorded during the 9.3 and 8.2 ka events in the groups, who used its wood for manufacturing tools and as fuel and Cantabrian range, together with the large fuel accumulation hazelnuts as a source of food (Uzquiano, 2018). What is clear, resulting from increased productivity, seem to have been key to the however, is that fire was not employed to maintain this vegetation occurrence of the first major Holocene fire episodes. community, as indicated by the macrocharcoal curve. Also, the At a regional scale, increased fire incidence has also been detection of coprophilous Cercophora might also indicate an in- recorded around Europe during the early Holocene (e.g., Power crease in animal presence in La Molina's surroundings, which most et al., 2008; Daniau et al., 2012; Marlon et al., 2016). In the Ibe- likely benefited from the forest openings and may also have rian Peninsula, although not all charcoal records have detected contributed to its maintenance by eating herbs and tree fresh buds. particularly localised fire intensity during this interval (Carrio n and It is assumed that the coprophilous signal probably indicates an Van Geel, 1999; Carrio  n et al., 2007; Miras et al., 2007; Anderson increase of wild animals rather than farming activities, since the et al., 2011; Rius et al., 2011, 2012; Connor et al., 2012; Morales- oldest domesticated fauna remains date approximately from 6900 Molino et al., 2013; Schneider et al., 2016; Garce s-Pastor et al., to 6500 cal yr BP in the Cantabrian region (Cubas and Fano, 2011). 2017; Leunda et al., 2020), there are some sites where fire activity Taking everything into account, increased human pressure and was notable. For example, Carrion (2002) detected high intensity of animal presence in the Cantabrian region could have regulated the fires between the onset of the Holocene and c. 8000 cal yr BP in the vegetation biomass, thus explaining the absence of pollen influx Siles lake, while Davis and Stevenson (2007) clearly identified peaks from the 8.2 ka event onwards. macrocharcoal peaks in Hoya del Castillo at around 9500 cal yr BP. Fires specifically located during the 8.2 ka event have provided 4.5. The creation and maintenance of open areas through the use of more evidence, in particular the charcoal peaks detected in El fire since the onset of agricultural practices Portalet (Gil-Romera et al., 2014), Bassa de la Mora (Pe rez-Sanz et al., 2013; Leunda et al., 2020) and Laguna Guallar (Davis and rez-Obiol et al. (2016) attribute an age of 6735 - 6495 cal yr BP Pe Stevenson, 2007). Therefore, the link between climate and fires to the oldest cereal pollen. The study of deeper sediment in La seems to be confirmed. In any case, it should be considered that Molina has not yielded older cereal evidence. Thus, according to the these fire episodes represented an opportunity to create open areas updated chronological model, an age of c. 6660 cal yr BP is esti- exploited by various Mesolithic groups. Furthermore, Mesolithic mated for the oldest cereal pollen in line with other locations from hunter-gatherers could have taken advantage of the arid conditions the northern Iberian Peninsula (Zapata, 2002; Zapata et al., 2004; to start the fires, which would represent a shared strategy with Iriarte-Chiapusso et al., 2005; Pen ~ a-Chocarro et al., 2005; Iriarte- other European regions. Chiapusso, 2009; Lo  pez-Merino et al., 2010). Likewise, the Whether the origin of fires was climatic, as seems to be the case, domestication of animals for human consumption went hand in anthropogenic, or resulted from the combination of both factors, hand with the beginning of agriculture, with an estimated origin the 9.3 and 8.2 ka fire episodes brought about a profund change in age between 6900 and 6500 cal yr BP (Cubas and Fano, 2011). In the Cantabrian landscape, representing a turning point towards a this regard, significant percentages of Cercophora, followed by more open environment. Both episodes triggered one-off biomass Sporormiella, have been detected since the onset of the Neolithic, declines affecting the vast majority of plant taxa, similar to the providing evidence of this grazing pressure. Agricultural practices 16 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 brought with them the need to create forest openings and the use Cantabrian range since the LGP and determining the role played of fire became frequent from c. 5700 onwards, as indicated by the by both the climate and anthropic factors: macrocharcoal record. In this case, the increase in the PAR detected between 5000 and 4000 cal yr BP, just before the beginning of fire 1) A weak local fire signal was detected from 17,550 to 13,700 cal yr activity, may not reflect a biomass increase as it did during the early BP that did not cause any significant change on the plant land- Holocene climate pulses. An increase in the peat bog sedimentary scape. The landscape was mainly composed of non-arboreal accumulation rate (Fig. 5) is probably responsible for such high species with a dominance of Poaceae accompanied by a lower values of PAR, which therefore do not reflect any climate pulse. proportion of other xerophyte taxa. The signal of arboreal pol- Although the fire signal increased from 5700 cal yr BP onwards, len, consisting of Pinus, evergreen and deciduous Quercus, it was not until 5000 cal yr BP when fires started exhibiting a more Betula, Corylus and the occasional presence of other meso- prominent magnitude. It seems that the maintenance of the philous taxa, did not reach 20%. The onset of the Bølling-Allerød continuous fire signal is likely due to anthropogenic causes, such as chronozone came together with Juniperus and Ericaceae pulses the need to preserve open spaces, yet a climatic relationship could and a decrease of Poaceae values. The low signal of Pinus found be attributed to the main fire peaks linked to Bond cycles (Bond through all the pollen spectra is indicative that the region has et al., 2001) as discussed in Pe rez-Obiol et al. (2016). In this line, never had extensive pine forests in the last 17,550 years. intervals of variable rainfall amount were detected in the Canta- 2) The woody colonisation was followed by the spread of Betula brian range during the late Holocene (Domínguez-Villar et al., and Salix at 13,700 cal yr BP, coinciding with increased water 2008). The palaeoclimate reconstruction performed in La Garma availability in the Cantabrian region. Betula and Salix decreased cave, at about 40 km from La Molina, determined that summers in favour of Poaceae and other herbs with the appearance of became progressively drier from 5900 cal yr BP onwards to reach tougher climate phases, first during the GI-1b and later within pronounced summer aridity between 4500 and 4200 cal yr BP the Younger Dryas Stadial (GS-1). (Baldini et al., 2019), coinciding with Bond event 3 (Bond et al., 1997, 3) A sharp increase in organic matter was detected at c. 2001). Baldini et al. (2019) reported Mediterranean-like conditions 12,000 cal yr BP, indicating an important change on climatic in the northern Iberian Peninsula at that time. Consistent with this, conditions. Shortly afterwards, a mixed deciduous formation the fire magnitude increased with local charcoal maximums with Betula and deciduous Quercus (>95% of arboreal pollen) around Bond event 3 (Fig. 9) which revealed a certain correlation dominated the landscape from 11,900 cal yr BP onwards. As a with climate. Another significant fire episode is detected around whole, a series of biomass fluctuations were detected at c. Bond event 2 (c. 2800 cal yr BP). In this respect, it is plausible that 11,600, 11,200, 10,130 and 9600 cal yr BP, probably related to the fire magnitude may have been particularly intense following century-to-millennial scale shifts of wetting and drying climatic pulses. conditions. Although the use of fire created forest openings, the arboreal 4) The arid conditions triggered by the 9.3 and 8.2 ka events composition does not seem to have been strongly influenced. It is together with the fuel accumulation produced during the early observed that some trees, such as Alnus or Fagus, started to expand Holocene were most likely responsible for the first significant thereafter, as already discussed in Pe rez-Obiol et al. (2016). On the fire episodes detected in the charcoal sequence. The fire epi- other hand, the most notable change is found in the composition of sodes created forest openings, as indicated by the higher pres- non-arboreal species. The open areas were henceforth dominated ence of Poaceae and Pteridium. Betula was severely affected and by Calluna vulgaris and other Ericaceae instead of grasses and its signal sharply decreased, exhibiting minimal values from this Pteridium. The germination of some heaths is stimulated by point onwards. On the other hand, the establishment of Corylus increased temperature, while most Erica spp. can resprout well was likely favoured by the fire episodes and a climate shift. after fire events (Iglesia-Rodríguez, 2010). Corylus became an important component of the mixed decidu- The vegetation dynamics from the Neolithic (c. 6740 cal yr BP) ous formations together with deciduous Quercus from that point onwards in La Molina are extensively described in Pe rez-Obiol et al. onwards. The plant landscape underwent no further significant (2016). They showed the importance of human influence on the changes until the Neolithic without the presence of biomass landscape related to fire activity during the main cultural phases fluctuations following climate pulses, contrary to what studied: the Neolithic, Bronze Age, Iron Age, Roman period, and happened during the early Holocene. It cannot be ruled out that Middle Ages. anthropic pressure favoured these landscapes during the Mesolithic, although it is clear that Mesolithic groups did not 5. Conclusions use fire to maintain them. The presence of coprophilous fungi likely indicates more animal presence, probably favoured by the The multiproxy approach performed in La Molina peat bog increase in open areas. (484 m a. s. l. Puente Viesgo, Cantabria) represents the oldest 5) The most recent 6600 years were characterised by a major continuous chronology studied from the northern Cantabrian anthropisation of the landscape. The updated chronological range, spanning the last 17,550 years. model indicated an age of c. 6657 cal yr BP for the first cereal The consideration of the pollen accumulation rate together with pollen grain. Coprophilous fungi maintained stable values the pollen percentages has allowed a better interpretation of the probably linked to grazing activites. The need to create open dynamics of each plant species, as well as the detection of biomass spaces went hand in hand with increased fire activity, which pulses that may go unnoticed when only pollen percentages are could no longer be easily related to the climate influence given available. the multitude of fire peaks. Some peaks of particular intensity Concerning the role of fire, the study of sedimentary charcoals could have been fostered by drier climate intervals, particularly (>150 mm) has proved to be an efficient tool to detect local fire during the Bond events 3 and 2 (c. 4200 and 2800 cal yr BP, episodes and helped to understand some landscape changes during respectively). The maintenance of fires involved a change in the the early Holocene, while the organic matter fluctuations served to non-arboreal composition, since Calluna vulgaris and other Eri- identify and delimit the main climate phases of a regional scope. caceae, well-adapted to fire, colonised the open areas where The multiproxy approach was a key to characterise the different Poaceae was dominant. plant landscape units and the fire regime of the north-eastern 17 nchez-Morales, A. Pelachs, J.C. García-Codron et al. M. Sa Quaternary Science Reviews 278 (2022) 107373 Funding Iberian Peninsula. Springer, Cham, pp. 191e250. https://doi.org/10.1007/978-3- 319-54867-8. Ancell, R., Ce lis-Díaz, R., 2012. T. In: ermopluviometría de Cantabria durante el This work has been supported by coordinated project grants periodo 1981-2010. Agencia Estatal de Meteorología (AEMET), nota te cnica nº from Spain's Ministry of Economics and Competitiveness (MEC): 10. NIPO: 281-12-014-0, vol. 20. 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Baena, J., Carrio  n, E., Ruiz-Zapata, B., Ellwood, B., Sese , C., Yravedra, J., Jorda, J., Uzquiano, P., Vela zquez, R., Manzano, I., Sanchez-Marco, A., Hern andez, F., 2005. Author contributions Paleoecología y comportamiento humano durante el Pleistoceno Superior en la comarca de Lie bana: La secuencia de la Cueva de El Esquilleu (Occidente de All the authors have made substantial contributions to the Cantabria, Espan ~ a). Baldini, L.M., McDermott, F., Baldini, J.U., Arias, P., Cueto, M., Fairchild, I.J., submission. In particular, Marc Sanchez Morales (Conceptualiza- ~ on, R., Garcia - Hoffmann, D.L., Mattey, D.P., Müller, W., Nita, D.C., Ontan tion; Methodology; Investigation; Writing - Original Draft; Visu- Monco  , C., Richards, D.A., 2015. Regional temperature, atmospheric circulation, alization), Albert lachs Pe Man~ osa (Conceptualization; and sea-ice variability within the Younger Dryas Event constrained using a Methodology; Resources; Writing - Review & Editing; Supervision; speleothem from northern Iberia. Earth Planet Sci. Lett. 419, 101e110. https:// doi.org/10.1016/j.quascirev.2019.105998. Project administration; Funding acquisition); Juan Carlos García Baldini, L.M., Baldini, J.U., McDermott, F., Arias, P., Cueto, M., Fairchild, I.J., Codron (Conceptualization; Writing - Review & Editing), Virginia Hoffmann, D.L., Mattey, D.P., Müller, W., Nita, D.C., Ontan ~ on, R., Garcia - Monco  , C., Richards, D.A., 2019. North iberian temperature and rainfall sea- Carracedo (Conceptualization; Writing - Review & Editing) and sonality over the younger Dryas and Holocene. Quat. Sci. Rev. 226. https:// Ramon Pe rez Obiol (Conceptualization; Methodology; Resources; doi.org/10.1016/j.quascirev.2019.105998, 105-998. 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