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ÔØ Å ÒÙ× Ö ÔØ
Landscape dynamics of Abies and Fagus in the southern Pyrenees during the
last 2200 years as a result of ant...
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1    LANDSCAPE DYNAMICS OF ABIES AND FAGUS IN THE SOUTHERN

2    PYRENEES DURING THE LAST 2200 YEARS ...
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15   LANDSCAPE DYNAMICS OF ABIES AND FAGUS IN THE SOUTHERN

16   PYRENEES DURING THE LAST 2200 YEARS ...
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40   its evolution (Galop and Jalut, 1994; Esteban et al., 2003; Riera et al., 2004; Beaulieu et

41 ...
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65   al., 2009). The hypotheses about distribution from the glacial refuges based on

66   isoenzyme ...
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90    Pérez-Obiol and Sadori, 2007) seem to indicate that Fagus had refuge in some concave

91    are...
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114   Similarly, much farther west of the Pyrenees, Montserrat (1992) explains that, although

115   ...
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139   Present day distribution of Fagus and Abies in the Iberian Peninsula related to

140   anthropo...
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164   the Iberian Peninsula could have been stopped by human interference in the Pyrenees.

165   Nev...
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189   winter to a mean minimum temperature of around -3.8 ºC. Potential evapotranspiration

190   (co...
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214   level, characterized by a granite conglomerate with very compacted gravel and some

215   pebbl...
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239   sites. In addition, plots that were absence sites for the species we considered were

240   avo...
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264   Pollen diagram from the València d’Àneu peat bog (B)

265   The pollen diagram from the Valènci...
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289   The intervention of the prior phase on the Abies alba forest opens up land that is

290   occup...
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314   VAL-III / III (1300-650 cal. yr BP; AD 650 – AD 1300): the explosion of human

315   activities...
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338   This zone is characterized by the notable presence of Alnus, which together with the

339   dyn...
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363   drastic change in the forest landscape is also evidenced by the decline in Alnus, Abies

364   ...
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388

389   VAL-III / VI (150 cal. yr BP--present; AD 1800 – present): the preamble to the

390   curr...
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413   chronologically similar and has three stages of impact, each of which is followed by a

414   d...
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438   by silver fir; this was the product of accumulating circumstances where climate change

439   a...
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463   think in terms of intertaxonomic differences if we are able to properly interpret pollen

464  ...
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488   that required limiting the diversity of resources available to peasants, who had to

489   spec...
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513   Abies recovers more readily when topoclimatic factors outweigh anthropic ones. From

514   the ...
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538   Silver fir shows a decline in this area due to factors much more related to human

539   interv...
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563   and also of COPCISA, which authorized access under the supervision of María Álvarez,

564   to ...
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587   Burjachs, F., Julià, R., 1994. Abrupt climatic changes during the last glaciation based on

588...
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611   Eisenhut, G., 1961. Untersuchungen iiber die Morphologie und Okologie der

612      Pollenkorne...
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636   Goeury, C., Beaulieu, JL de., 1979. A propos de la concentration du pollen a l’aide de la

637 ...
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660   Kenla, J.V., Jalut, G., 1979. Déterminisme anthropique du développement du hêtre dans

661     ...
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683   López- Merino, L., López-Sáez, J.A., Ruiz, M.B., Gil, M.J., 2008. Reconstructing the

684     h...
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707      Néolithique ancien, d’après l’analyse pollinique de la tourbière de Bosc dels

708      Esta...
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730   Ninyerola, M., Pons, X., Roure, JM., 2007b. Objective air temperature mapping for the

731     ...
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754   Pérez-Obiol, R., Sadori, L., 2007. Similarities and dissimilarities, synchronisms and

755     ...
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779   Talma, A. S, Vogel, J. C., 1993. Radiocarbon 35 (2), 317-322.

780   Tantinyà, M., 2007. L’acci...
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794   Fig. 1. Location of the València d’Àneu (VAL-III) peatbog (star) and fir forest (in grey)

795 ...
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      Sample     Laboratory     Material    Conventional   Dating calibrated to Intercept calibr...
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                  MX_HOT MT_AN MN_COL PR_WIN PR_SPR PR_SUM PR_AUT RAD_WIN RAD_SPR RAD_SUM RAD_AUT...
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Landscape dynamics of Abies and Fagus in the southern Pyrenees during the last 2200 years as a result of anthropogenic impacts - Albert Pelachs

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Landscape dynamics of Abies and Fagus in the southern Pyrenees during the last 2200 years as a result of anthropogenic impacts - Albert Pelachs

  1. 1. ÔØ Å ÒÙ× Ö ÔØ Landscape dynamics of Abies and Fagus in the southern Pyrenees during the last 2200 years as a result of anthropogenic impacts Albert P` lachs, Ramon P´ rez-Obiol, Miquel Ninyerola, Jordi Nadal e e PII: S0034-6667(09)00048-7 DOI: doi: 10.1016/j.revpalbo.2009.04.005 Reference: PALBO 3026 To appear in: Review of Palaeobotany and Palynology Received date: 26 September 2008 Revised date: 24 March 2009 Accepted date: 1 April 2009 Please cite this article as: P`lachs, Albert, P´rez-Obiol, Ramon, Ninyerola, Miquel, e e Nadal, Jordi, Landscape dynamics of Abies and Fagus in the southern Pyrenees during the last 2200 years as a result of anthropogenic impacts, Review of Palaeobotany and Palynology (2009), doi: 10.1016/j.revpalbo.2009.04.005 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
  2. 2. ACCEPTED MANUSCRIPT 1 LANDSCAPE DYNAMICS OF ABIES AND FAGUS IN THE SOUTHERN 2 PYRENEES DURING THE LAST 2200 YEARS AS A RESULT OF 3 ANTHROPOGENIC IMPACTS T 4 5 Albert Pèlachs a,*, Ramon Pérez-Obiol b, Miquel Ninyerola b, Jordi Nadal a P 6 RI 7 a GRAMP, Departament de Geografia, Universitat Autònoma de Barcelona. 08193 Bellaterra 8 (Cerdanyola del Vallès). Spain. 9 SC 10 b Unitatde Botànica, Facultat de Biociències, Universitat Autònoma de Barcelona, 08193 11 Bellaterra (Cerdanyola del Vallès). Spain. 12 NU 13 * correspondingauthor. Tel.: +34 93 5868057; fax: +34 93 5812001. 14 E-mail address: albert.pelachs@uab.cat (A. Pèlachs). MA ED PT CE AC
  3. 3. ACCEPTED MANUSCRIPT 15 LANDSCAPE DYNAMICS OF ABIES AND FAGUS IN THE SOUTHERN 16 PYRENEES DURING THE LAST 2200 YEARS AS A RESULT OF 17 ANTHROPOGENIC IMPACTS T 18 P 19 Abstract RI 20 The vegetation landscape dynamic is derived from the relationship established between SC 21 a society and its environment through time, and the current landscape has never been 22 seen in the previous 2000 years. The pollen study of a core from a peat bog in València 23 24 NU d'Àneu (Lleida, NE Iberian Peninsula) shows a maximum extension of Abies alba forest MA about 2200-2000 cal. yr BP. Later on, there is evidence of selective actions affecting 25 this forest and the expansion of Fagus sylvatica at about 2000-1300 cal. yr BP. 26 Beginning in 1300 cal. yr BP, deforestation due to agricultural activities expanded and ED 27 beech definitively disappeared at 800 cal. yr BP. Natural and human disturbances PT 28 affected the dynamics of Abies alba and Fagus sylvatica from their first appearance to 29 the current vegetation landscape. Human impact on the silver fir forest, which reached CE 30 its maximum in the last millennium, favoured the beech population. Pollen data from AC 31 this region support our finding that human impact, not climate, is the most important 32 influential factor in the development of beech forests. 33 34 Keywords: Pyrenees, Holocene, palynology, GIS suitability mapping, Abies alba, 35 Fagus sylvatica. 36 37 Introduction (A) 38 The current discussion concerning the dynamics of the vegetation landscape is rooted in 39 the reasons for change over time and in the weighting of natural and human factors in
  4. 4. ACCEPTED MANUSCRIPT 40 its evolution (Galop and Jalut, 1994; Esteban et al., 2003; Riera et al., 2004; Beaulieu et 41 al., 2005; Riera et al., 2006; Pèlachs et al., 2007). Although climatic factors have a very 42 important role in the development of vegetation, palaeobotanic studies have T 43 demonstrated the importance of taking into account the role played by human society. P 44 Therefore, the primary objective of this study is to determine the extent to which the RI 45 human imprint has affected the current vegetation landscape, focussing on the dynamics SC 46 of Abies and Fagus forests in the Pyrenees. 47 48 49 NU Colonization of Abies alba and Fagus sylvatica: the current state of affairs (B) MA An explanation of the plant colonization of the Pyrenees from the beginning of the 50 Holocene can be undertaken on the basis of pollen analyses available from the Pyrenees 51 mountain range (Jalut et al., 1998). It is impossible to interpret which factors affect this ED 52 evolution without taking into account at least three variables: the location of refuge PT 53 zones, the development of climatic factors and the edaphic dynamics of the soils 54 (Pèlachs, 2005). CE 55 In recent years, the study of Abies alba dynamics in Europe (Terhürne-Berson et al., AC 56 2004; Liepelt et al., 2009) has been associated with other species, such as Fagus 57 sylvatica (Tinner and Lotter, 2006). This area of study has developed from a series of 58 interpretations based on the study of climate change, migratory change, unequal growth 59 of species, and the effects of human disturbances and forest fires (Tinner and Lotter, 60 2006). 61 In this sense, phylogenetic studies reveal how the Abies populations in the Pyrenees 62 were isolated from the rest of Europe (Konnert and Bergmann, 1995). This argument 63 was definitive in defending the proximity of the Pyrenees to Abies alba refuge zones, 64 based on plant macroremains and pollen data (Terhürne-Berson et al., 2004; Liepelt et
  5. 5. ACCEPTED MANUSCRIPT 65 al., 2009). The hypotheses about distribution from the glacial refuges based on 66 isoenzyme studies and other genetic markers (El Mousadik and Petit, 1996) seem to 67 substantiate the existence of five areas of Abies alba refuge and recolonization: the T 68 Pyrenees, central and eastern France, central Italy and the southern Balkans. Pollen and P 69 genetic data indicate clearly that the Abies alba and Fagus sylvatica refuges in the RI 70 Pyrenees have suffered the “bottleneck” phenomenon during their history and that SC 71 recolonization was not produced exclusively from refuge populations. This theory is 72 well supported because of the low allelic levels, which can be correlated to the current 73 74 NU distribution of silver fir in the Pyrenees, with populations that are not extensive in MA comparison with the rest of Europe. 75 Palaeobotanical and genetic data for Fagus sylvatica (Magri et al., 2006) have been 76 used to evaluate the genetic consequences in Europe of long-term survival in refuge ED 77 areas and postglacial spread. The largely complementary palaeobotanical and genetic PT 78 data indicate that Fagus sylvatica survived the last glacial period in multiple refuge 79 areas. The central European refuges were separated from the Mediterranean refuges, CE 80 which did not contribute to the colonization of central and northern Europe. Likewise, AC 81 some populations expanded considerably during the postglacial period (Magri, 2008), 82 while others experienced only limited expansion. According to Ninyerola et al. (2007a), 83 inferences from more than a few studies lend credibility to the presence in the 84 Mediterranean of deciduous taxa such as Fagus during the early and mid-Holocene. The 85 climatic suitability of Fagus during the early Holocene has been shown by Lozano et al. 86 (2002), who identified Fagus and dated it at c. 17,895 cal. yr BP in Urdaibai (Basque 87 County) or López-Merino et al. (2008) in Sierra de Neila at c. 15,600-13,700 cal. yr BP. 88 This led them to suggest the northern Iberian Peninsula as a possible refuge zone 89 (Hewitt, 1999). In the Balearic Islands, the available data (Ninyerola et al., 2007a;
  6. 6. ACCEPTED MANUSCRIPT 90 Pérez-Obiol and Sadori, 2007) seem to indicate that Fagus had refuge in some concave 91 areas during the upper Pleistocene and the Holocene. The presence of small stands of 92 Fagus in Majorca, before the colonization from the Pyrenees took place, makes this a T 93 credible hypothesis. Similarly, examining the Iberian Peninsula, Pott (2000) indicates P 94 that over the last 9000 years Fagus has colonized northern areas from diverse RI 95 Pleistocene Mediterranean refuges. SC 96 In the Iberian Peninsula, evidence exists (Costa et al., 1998) of the presence of Fagus 97 sylvatica in the Basque Country (Saldropo) and Tramacastilla more than 4000 and 7000 98 99 NU years ago, respectively, which would confirm the presence of various refuge zones in MA the southern slope of the Pyrenees (Montserrat, 1992). This pattern of colonization is 100 supported by pollen records from the northeast Iberian Peninsula (Pérez-Obiol, 1988), 101 showing that Fagus colonization began between 8800 and 7850 cal. yr BP. ED 102 The difficulty comes from site differences that enormously complicate the interpretation PT 103 of local pollen and charcoal records, as at Burg Lake in the Pyrenees, close to the study 104 CE area, where Fagus sylvatica does not appear until 3000 cal. yr BP (1050 BC) (Pèlachs, 105 2005). AC 106 On the other hand, regional data support the introduction of Fagus sylvatica at Redó 107 Lake at about 4900 cal. yr BP (Catalan et al., 2001), and a little later at Redon Lake 108 (Catalan and Pla, 1998), where it arrives in about 4500 cal. yr BP, probably as a 109 consequence of the difference in altitude (Esteban et al., 2003). Miras et al. (2007) 110 implicate both anthropic participation and onset of new climate conditions (lower 111 summer temperatures and higher annual precipitation) in the timing of the first regular 112 observations of Fagus sylvatica in the Andorran valley of Madriu, at about 4800 cal. yr 113 BP.
  7. 7. ACCEPTED MANUSCRIPT 114 Similarly, much farther west of the Pyrenees, Montserrat (1992) explains that, although 115 beech appears intermittently at Ibon de Tramacastilla after 7859 cal. yr BP, its curve 116 does not become continuous until c. 5760 – 4476 cal. yr BP, making its appearance T 117 contemporaneous with the other Pyrenees sites. P 118 No one disputes that the Abies alba dynamics in the Pyrenees during the Holocene RI 119 indicate colonization followed by expansion from east to west (Jalut et al., 1998; SC 120 Esteban et al., 2003; Pèlachs, 2005; Le Flao, 2005), which would confirm the existence 121 NU of refuges located in the Mediterranean basin. In fact, analysis of the current western 122 boundaries of Abies alba in the Pyrenees shows a progressive lag between the western MA 123 and eastern half of the mountain chain, which could be attributed to the progressive 124 distancing of this conifer from its refuge areas (Reille and Andrieu, 1991). Similarly, ED 125 some authors have reported that this species first developed on the north slope of the 126 Mediterranean Pyrenees at 11,224 cal. yr BP, specifically in the area of Nohèdes (Jalut, PT 127 1974; Reille and Lowe, 1993); this is consistent with the very first appearance of Abies 128 CE alba in the Garrotxa at about 10,204 cal. yr BP (Pérez-Obiol, 1988). Other registries of 129 long-term silver fir presence in the eastern Mediterranean also concur, e.g., Pla de AC 130 l’Estany (Burjachs, 1994), Banyoles (Pérez-Obiol and Julià, 1994), and Abric Romaní 131 (Burjachs and Julià, 1994), confirming the presence of refuges in coastal zones and in 132 intramountain valleys of the Iberian Peninsula (Carrión-García et al., 2000). Therefore, 133 colonization of Abies alba and Fagus sylvatica in the meridional slope of the Pyrenees 134 could be due, in part, to refuge zones located to the south and east of the Pyrenees (Fig. 135 1). 136 137 [FIGURE 1] 138
  8. 8. ACCEPTED MANUSCRIPT 139 Present day distribution of Fagus and Abies in the Iberian Peninsula related to 140 anthropogenically forced landscape changes (B) 141 Although climate has been regarded as the determining factor in the development of T 142 Fagus sylvatica forests at c. 4500 cal. yr BP (Jalut, 1974; Giesecke et al., 2007), it has P 143 also been demonstrated that human influence may be responsible for its strong RI 144 expansion at that time (Kenla and Jalut 1979; Jalut 1984; López-Merino et al., 2008). SC 145 According to Tinner and Lotter (2006), beech survived human pressure, while other 146 deciduous trees (e.g. Tilia, Ulmus, Fraxinus excelsior) and silver fir (Abies alba) were 147 148 NU strongly disadvantaged. The authors hypothesize that in the absence of human impact, MA silver fir would have expanded to areas in Europe where the species is absent today. 149 According to Peñalba (1994), the western and southernmost parts of the peninsula have 150 not been colonized by Fagus. The absence of Fagus in northwestern Spain is striking, ED 151 given the importance of this genus in similar climatic conditions in the other Cantabrian PT 152 regions. It is unlikely that the spread of Fagus was stopped in Galicia by natural causes 153 at 1390 cal. yr BP. At that time, humans exerted strong influence on the vegetation in CE 154 this region; their presence there is recorded since 5760 cal. yr BP. Anthropogenic AC 155 disturbance has proved responsible for the final, abrupt decline of Fagus populations in 156 the Cantabrian region. It is likely that severe anthropic pressure on populations of Fagus 157 at their range limit stopped the spread to the west. A similar situation could be inferred 158 for Abies, confined today to the eastern part of the Pyrenees although it had a wider 159 distribution in the Iberian Peninsula during previous interglacial periods. Two facts 160 must be considered: first, man favoured Fagus to the detriment of Abies at the 161 beginning of its extension to the northern side of the Pyrenees (Jalut 1984), and second, 162 Abies grows today in Italy under climatic conditions also found in Spain (Terhürne- 163 Berson et al., 2004; Liepelt et al., 2009), suggesting that the spread of the species into
  9. 9. ACCEPTED MANUSCRIPT 164 the Iberian Peninsula could have been stopped by human interference in the Pyrenees. 165 Nevertheless, climate forcing in the Post-Bronze Iberian Roman Humid Period (2600- 166 1600 cal. yr BP) could be a consideration, as proposed by Martín-Puertas et al. (2008). T 167 Fig. 2 shows the clear decline of Abies alba beginning in the medieval period. When the P 168 human impact was too strong, silver fir totally disappeared (Pérez and Roure, 1990; RI 169 Pèlachs, 2005; Tantinyà, 2007). SC 170 The potential distribution of Abies alba in the northwest Iberian Peninsula proposed by NU 171 Rivas (1987) would result in a much larger region with a much more suitable surface if 172 we consider numerous biotic and abiotic factors that exist at present. To enhance the MA 173 potential distribution of these two taxa, a combined spatial suitability surface has been 174 developed through GIS and multivariate statistical methods. This map allows us to ED 175 understand the spatial behaviour of Fagus and Abies at regional scale, complementing 176 the palaeopalynological results. PT 177 CE 178 [FIGURE 2] 179 AC 180 Study Area (A) 181 The Prats de Vila peat bog (longitude 1° 6’ 13” E and latitude 42° 38’ 17” N) is found at 182 1,150 masl and has an estimated area of 2.8 hectares. The lithological substrate 183 corresponds to Cambro-Ordovician slates, even though during the fieldwork we found 184 important granite deposits of glacial remains. 185 The climatic conditions surrounding the peat bog (within a 1 km radius) are humid 186 (Thornthwaite humidity index) with an Autumn-Spring-Summer-Winter precipitation 187 pattern and mean annual values ranging between 652 mm and 887 mm (=736 mm). The 188 mean annual temperature ranges between 6.5 ºC and 10.5 ºC (=8.9 ºC), decreasing in
  10. 10. ACCEPTED MANUSCRIPT 189 winter to a mean minimum temperature of around -3.8 ºC. Potential evapotranspiration 190 (computed following the Hargreaves method) shows annual values ranging between 574 191 mm and 850 mm (=716). These values are close to precipitation values, meaning that T 192 this area is free of hydric stress. All the climate data have been extracted from the P 193 Digital Climatic Atlas of the Iberian Peninsula (Ninyerola et al., 2007b and 2007c). RI 194 The present vegetation on the peat bog is Subalpine-Montane mesophilous and siliceous SC 195 meadows with Agrostis capillaris, Festuca nigrescens, Anthoxanthum odoratum, 196 Galium verum, and Genistella sagittalis. Vegetation surrounding the peat bog in shady 197 198 NU places includes some deciduous Quercus together with Corylus, Betula and Pinus, MA which in many cases occupy formerly cultivated fields and give way to the most 199 extensive Abies alba stands of the Pyrenees: la Mata de València d’Àneu. In northern 200 Spain, distribution of Quercus petraea (the dominant oak near the study zone) is ED 201 typically fragmented. Taking into account its minimal presence in the pollen diagram, it PT 202 appears that its distribution area in the study zone has not been of great importance 203 during the last millennia. At the same time, in sunny places, the deciduous Quercus CE 204 share their protagonism with Q. Ilex subsp rotundifolia. AC 205 206 Materials and Methods (A) 207 The study methodology was based on a combination of pollen data extracted from a 208 peat bog in València d’Àneu (Axial Pyrenees) and fieldwork to identify the main plant 209 communities in the zone. 210 Three core samples were taken with a mechanical sampler and the one with the most 211 consolidated peat was selected for analysis. Two large, clearly differentiated 212 sedimentary units have been described in the register of the peat bog studied (Fig. 3): 213 the upper unit, characterized by the abundance and continuity of the bog, and the lower
  11. 11. ACCEPTED MANUSCRIPT 214 level, characterized by a granite conglomerate with very compacted gravel and some 215 pebbles at the transition between the two units. Two samples were selected for dating 216 using 14C-AMS (Beta Analytic Inc.), based on a piece of wood at 59-60 cm depth and a T 217 peat fragment at 165-166 cm depth (Table 1). The resulting sedimentation rate for the P 218 peat section was 0.72 mm/year for the first 60 cm and 0.83 mm/year for the rest. Age RI 219 was calibrated to calendar age using the INTCAL04 program (Talma and Vogel, 1993). SC 220 For the pollen analysis, we selected only the first two meters of peat from one of the 221 cores (named VAL-III), down to the transition to gravel conglomerate. Chemical 222 223 NU treatment of the samples was carried out according to the protocol described by Goeury MA and Beaulieu (1979). 224 [FIGURE 3] 225 [TABLE 1] ED 226 PT 227 Suitability mapping (B) 228 The suitability vegetation maps for Fagus and Abies were developed using presence- CE 229 absence models adjusted with logistic linking in a General Linear Model (GLM). AC 230 Presence data were obtained by choosing plots where these species are dominant from 231 the third National Forest Inventory (a project administered by the Spanish state). The 232 resulting distribution of both species is shown in figures 7-8. This forest inventory 233 regularly samples the territory with a grid density of 1 km. This type of sampling is 234 very interesting because it covers a large area but especially because regular sampling 235 avoids the sampling bias that exists in many other types of chorological data. We would 236 also note that we have access to plots in which the absence of the species studied is 237 ensured, mitigating the problem of pseudo-absences (Chefaoui and Lobo, 2008). To 238 obtain an absence sample, we randomly chose a number of sites equal to the presence
  12. 12. ACCEPTED MANUSCRIPT 239 sites. In addition, plots that were absence sites for the species we considered were 240 avoided if they were within a 5 km radius of the presence plots and therefore had very 241 similar topoclimatic conditions. T 242 With respect to predictor variables, we incorporated geoclimatic variables obtained by P 243 spatial interpolation methods (Ninyerola et al., 2000), based on a Digital Elevation RI 244 Model with 200-m resolution and data from Spain’s National Institute of Meteorology SC 245 (INM), which provided readings from 1346 temperature stations and 2519 for 246 precipitation. We would emphasize here that having access to a plot that was 247 248 NU georeferenced with a high level of precision allowed us to capture the climatology at MA toposcale, minimizing methodological errors. Five variables were analysed: maximum 249 mean temperature for the warmest month, mean annual temperature, minimum mean 250 temperature for the coldest month, accumulated precipitation by season and potential ED 251 solar radiation by season. Table 2 shows the ranges for Abies alba and Fagus sylvatica. PT 252 We then enriched the databases using vector point files (presence-absence distribution) 253 with the corresponding values from the geoclimatic variables. This enriched database CE 254 was submitted to statistical analysis using a GLM with logistic linking, as in other AC 255 studies (Felicísimo et al., 2002) of the suitability of forest species. For the process of 256 adjusting the model we used 60% of the plots and saved 40% for validation and to be 257 able to quantify in this way the quality of the resulting maps. 258 Finally, mapping algebra was used to obtain the suitability maps by species using the 259 completed analysis. The regression equations, adjusted by statistical analysis, were 260 reproduced using GIS, replacing each variable with the corresponding topoclimatic 261 map. 262 [TABLE 2] 263 Results and discussion (A)
  13. 13. ACCEPTED MANUSCRIPT 264 Pollen diagram from the València d’Àneu peat bog (B) 265 The pollen diagram from the València d’Àneu peat bog permitted us to reconstruct the 266 vegetation changes in the studied zone over the last two millennia (Fig. 4). The diagram T 267 is described using pollen assemblage zones (PAZ). P 268 RI 269 [FIGURE 4] SC 270 271 VAL-III / I (2200-2000 cal. yr BP; 250 BC –50 BC): the decline of the 272 273 “original”Abies alba forest (C) NU MA At the beginning of this time period, Abies frequency of more than 10% with a peak at 274 approximately 22% can only be explained by the Abies alba in situ occupying a much 275 larger land area than at present. The drop in Abies at the end of this period may be due ED 276 to selective human intervention with respect to this species, favouring other species PT 277 such as Corylus, which would colonize the space left by silver fir. We must take into 278 account the fact that wood forms part of the Roman social and economic system and is CE 279 an indispensable element (Conedera et al., 2004). AC 280 Mining is another sector related to exploitation of forest resources. We noted an 281 increase in lead in the sediment of Redon Lake (also in the axial Pyrenees) during the 282 Roman era and a high point in about AD 600 (Catalan and Pla, 1998). The dating of five 283 charcoal kiln sites between the 3rd and 4th VI centuries AD and the identification of 284 charcoals (Pinus and Abies) allows us to relate this first metallurgy with selective acts 285 related to the forest (Pèlachs and Soriano, 2003). 286 287 VAL-III / II (2000-1300 cal. yr BP; 50 BC – AD 650): Abies alba forest with Fagus 288 sylvatica (C)
  14. 14. ACCEPTED MANUSCRIPT 289 The intervention of the prior phase on the Abies alba forest opens up land that is 290 occupied first by some species that are typical of meadows and clearings (Poaceae, 291 Plantago sp., Asteraceae, etc.) and permit the expansion of plant populations that T 292 compete with the silver fir for space, such as Corylus in the lowest areas and Fagus, P 293 Pinus and Betula in the same zones as the Abies alba. RI 294 The occasional presence of Juglans, Juniperus and Artemisia and the start of Cerealia SC 295 and Castanea curves denote human management of the landscape, mostly related to 296 NU grazing and agricultural activities. Pseudoschizaea (an algal remain indicative of 297 erosive processes) appears. The first occurrences of Juglans are well dated at Ariege MA 298 (2000±107 cal. yr BP in Jalut el al. 1982; 1792±59 cal. yr BP, Galop, unpublished). 299 There are regular records from the 10th to the 13th centuries, though the dates may vary ED 300 with area and altitude (1048±79 cal. yr BP and 706±28 cal. yr BP, Galop, unpublished; 301 near 643±61 cal. yr BP, Planchais 1985). This cultivated tree is an excellent marker of PT 302 the Greco-Roman times. It was introduced in western Mediterranean regions as early as 303 CE c. 1952 cal. yr BP (Bottema, 1980) by Greek and Roman settlers. According to the 304 curve of pollen concentration (pollen grains/g), the arboreal biomass does not suffer a AC 305 significant decline (Fig. 5). However, forest activities are evident. 306 In any case, the plant dynamics indicate a human pressure that shifts the permanent 307 character of the land. Without technical resources to minimize labour expenditure, mid- 308 slope soils are preferred for agricultural uses (Esteban et al., 2003). This is the reason 309 for disturbances of mid and lower slopes of the forest that affect the dynamics of the 310 silver fir-beech forest. 311 312 [FIGURE 5] 313
  15. 15. ACCEPTED MANUSCRIPT 314 VAL-III / III (1300-650 cal. yr BP; AD 650 – AD 1300): the explosion of human 315 activities (C) 316 This entire phase is characterized by a declining AP percentage and an absolute increase T 317 of herbaceous plants (Fig. 5). The massive forest clearance during this period is shown P 318 by the fall in AP values and the greater Poaceae abundance in the studied area. The RI 319 decline of Abies and the noticeable extension of Fagus are probable evidence of this SC 320 deforestation. 321 NU The pollen diagram shows certain peculiarities that led to splitting the zone into three 322 subzones: MA 323 324 VAL-III / IIIa (1300-1100 cal. yr BP; AD 650 – AD 850): global disturbance (C) ED 325 At the same time that Pinus recedes below 20% and Abies falls below 5%, Fagus, 326 Betula and Corylus take advantage of this by increasing their presence even though, PT 327 later on, they will decline just as the rest of the tree population did. CE 328 The large increase in Artemisia, Poaceae, Rumex and Polygonum can be explained by 329 the increase in grazing. The strong increment of Cerealia (mostly Secale) and Fabaceae AC 330 also indicate the implementation of agricultural practices. This evidence permits us to 331 assume that opening up the landscape led to the arrival of Olea pollen. In that era, olive 332 tree cultivation is documented in the domains of a nearby monastery (Esteban et al., 333 2003). These facts are clearly evidenced by the drop in pollen concentration. The impact 334 of human disturbance is more noticeable from the Late Medieval period onward. 335 336 VAL-III / IIIb (1100-800 cal. yr BP; AD 850 – AD 1150): management of the peat 337 bog (C)
  16. 16. ACCEPTED MANUSCRIPT 338 This zone is characterized by the notable presence of Alnus, which together with the 339 dynamics of Cyperaceae and Typha-Sparganium pollen type allows us to connect this 340 period with an increase in the groundwater level of the peat bog and possibly with its T 341 expansion. P 342 Sparganium sp. has great colonizing abilities and may cause a rapid silting in shallow RI 343 waters. At this time, its development coincides with the establishment of Alnus. Before SC 344 this colonization, Pediastrum was already present, indicating a rise in water level. These 345 percentage increases in taxa are related to a major sedimentary stability (Andrieu et al., 346 347 NU 2000). Late Medieval period documents explain that during this period it was common MA to plant crops in muddy zones along river banks, which flooded periodically and were 348 called “insules” (Esteban et al., 2003); consequently, it would seem reasonable that a 349 hygrophilous environment was favoured, controlling the flow and the hydric resources ED 350 of the area. PT 351 The rapidly invading Abies would out-compete Fagus, or substantially slow down its 352 recruitment rate until canopy disturbance created light openings large enough for CE 353 successful establishment and growth. According to Doležal et al. (2004), the higher AC 354 mortality of Fagus in denser Abies patches and the resulting spatial segregation of the 355 species reflect asymmetric interspecies competition. 356 357 VAL-III / IIIc (800-650 cal. yr BP; AD 1150 – AD 1300): disappearance of the Abies 358 alba-Fagus sylvatica forest (C) 359 The beginning of this phase is characterized by high percentages of Poaceae and 360 Cerealia and the disappearance of Fagus from the study area, a disappearance attributed 361 to the strong human impact on the landscape. From this point on, there will never again 362 be a beech forest or a small mixed Abies alba-Fagus sylvatica forest in the zone. This
  17. 17. ACCEPTED MANUSCRIPT 363 drastic change in the forest landscape is also evidenced by the decline in Alnus, Abies 364 and Pinus, which at the end of the sequence permits the return of Corylus. 365 T 366 VAL-III / IV (650-350 cal. yr BP; AD 1300 – AD 1600): recovery of the Abies alba P 367 forest (C) RI 368 Since 650 cal. yr BP (AD 1300) we have observed a certain recovery of the arboreal SC 369 cover, led by the presence of three primary species of trees that are distributed and 370 combined in various stages and habitats: Corylus, Abies and Pinus; Betula is added to 371 372 NU the list at the end of this time period. Human pressure on the environment is moderate. MA Therefore, it doesn’t seem that the repercussions of the Little Ice Age were sufficiently 373 important to affect the economic activities of the dominant classes, primarily herders. 374 All the same, documents report major declines in the Pyrenees in some of the species ED 375 grown (such as grapevines), which leads us to assume the existence of local differences. PT 376 377 VAL-III / V (350-150 cal. yr BP; AD 1600 – AD 1800): a new increase in human CE 378 pressure (C) AC 379 The slight percentage oscillations in various tree taxa, such as Abies, Corylus, Betula 380 and Pinus, are accompanied by a large increase in Poaceae and Juglans; this denotes a 381 new and different landscape management with the existence of pastures and plantations 382 of trees. It is worth noting that oil was extracted from the walnut trees and had a high 383 food and therapeutic value, equal or superior to that of olive oil, and therefore at 384 particular times could have offered an alternative to the cultivation of olive trees 385 (Esteban et al., 2003). In addition, the appearance of Ericaceae could indicate an 386 increase in ruderal species, given the use of a road network, and that of Glomus would 387 explain the more edaphic conditions of the peat bog.
  18. 18. ACCEPTED MANUSCRIPT 388 389 VAL-III / VI (150 cal. yr BP--present; AD 1800 – present): the preamble to the 390 current landscape (C) T 391 The final episode puts the vegetation landscape at the doorstep of the current landscape, P 392 with the percentages of Abies at about 5%, while Pinus recedes significantly and RI 393 heliophilous colonizers increase progressively in formerly cultivated zones and open SC 394 forest areas, including Corylus –especially at the end of the sequence – or plastic 395 species such as Betula. This occurred in other areas as well. 396 397 NU This denotes a decrease in the groundwater of the peat bog as indicated by the curve for MA Cyperaceae and Glomus and suggests the definitive disappearance of Alnus around the 398 bog studied here. Chlamydospores of Glomus cf. fasciculatum would be evidence of 399 erosive phenomena (Van Geel et al., 1989) related to anthropogenic activity and drought ED 400 (López-Sáez et al., 2000). PT 401 402 [FIGURE 5] CE 403 AC 404 Vegetation dynamics and suitability (B) 405 It is clear from the palynological data presented here that human impact became 406 stronger and reaches its maximum in this last millennium. This stage of the Pyrenean 407 forest history saw the final shaping of the present-day landscape (Kenla and Jalut 1979; 408 Galop, 1998). 409 The pollen diagram is comparable to numerous diagrams of the southern and central 410 Alps, central France and the Pyrenees themselves (Beaulieu, 1978; Clerc, 1988; David, 411 1993; Nakagawa, 1998; Tinner et al., 2005; Finsinger and Tinner, 2006; Pèlachs et al., 412 2007). In the central Alps, Nakagawa et al. (2000) found a sequence that is
  19. 19. ACCEPTED MANUSCRIPT 413 chronologically similar and has three stages of impact, each of which is followed by a 414 different pattern of forest restoration. The first deforestation occurs at about 2060 cal. yr 415 BP, during the Roman era, and a selective exploitation of Abies alba forest is evidenced. T 416 The silver fir forests formed part of a very active economy near the Rhine river. Küster P 417 (1994) compiles various pollen diagrams for the Rhine, Elbe, and Danube and RI 418 demonstrates that the use during Roman times was not totally destructive. Various SC 419 zones of silver fir forest were left untouched. The author concludes that the concept and 420 practice of “forest management” was common in Roman times. The second 421 422 NU deforestation, around 1520 cal. yr BP (or during the 5th and 6th centuries), denotes MA substantial evidence of agricultural activity. The third, around 810 cal. yr BP or right in 423 the middle of the 12th century, is similar to its predecessor but much longer and not at 424 all selective, so that the forest had no chance to recover. ED 425 These facts coincide quite well with the changes in percentages and pollen PT 426 concentration for Abies (Fig. 5). This dynamic also coincides with those found in other 427 localities close to the studied zone (Esteban et al., 2003; Pèlachs et al., 2007). The peat CE 428 bog studied demonstrates much more clearly a possible selective action involving Abies AC 429 alba forests during the Roman era and confirms the indices that explain how some 430 dynamics began in the medieval period, continued during the Modern Age and the 20th 431 century, and brought us to the current landscape. 432 In other areas of the Pyrenees, Abies alba was the primary species of trees between 433 6200 and 2800 cal. yr BP (4250-850 BC), a time when the stable Abies alba presence 434 gave way to red pine forest in the subalpine stage. At the same time this was happening, 435 a mixture of oak (Quercus sp., Tilia sp., Ulmus sp. etc.) also experienced a sharp 436 decrease. This strong disturbance of the subalpine and mountain area would permit the 437 pine forest to expand as a rapid colonizer and populate the space that had been occupied
  20. 20. ACCEPTED MANUSCRIPT 438 by silver fir; this was the product of accumulating circumstances where climate change 439 and human actions intersected (Pèlachs et al., 2007). In the studied zone this didn’t 440 happen and the silver fir, despite the strong disturbance they suffered, recovered again T 441 and again, even though the population would never reach the levels of 2000 years P 442 earlier (Fig. 4). Abies forests remained important during a large part of the Holocene, RI 443 which could be explained by the topography of the valley and slopes. SC 444 The current pollen spectrum had never been seen in the previous 2000 years. This fact 445 NU led us to deduce that models such as Modern Analogue Technique (MAT) could be 446 difficult to apply in this zone of the Pyrenees, at least during the last 2000 years. MA 447 Establishing detailed comparisons, we observe notable differences between two data 448 groups of interest: pollen and vegetation cover; this means that we must explore models ED 449 that work for mountain regions in particular. Calibrating the mountain vegetation and 450 pollen spectra is key to this type of research if we are to understand certain evolutionary PT 451 patterns. The hypotheses of authors such as Muller et al. (2005), which postulate that 452 CE there is an increase in regional and distant pollen in sediment at high altitude, is only 453 valid for certain taxa. In sedimentary samples of lake surfaces, we see that the presence AC 454 in the pollen spectra of taxa such as Tilia, Abies, Ulmus and Fagus almost always 455 represents a local or nearby presence in mountain regions. Many calibrations have used 456 correction factors or R-values (the ratio between the pollen group and the vegetation 457 community it represents). At present, different models are grouped within the Extended 458 R-value (ERV). With respect to Abies, a taxon that has had a strong impact on the 459 evolution of the vegetation landscape in this zone during the last 2000 years, it must be 460 said that it is very sensitive to the described method of weighing distance. For example, 461 according to Eisenhut (1961) Abies alba presents a falling speed of 0.12m.s-1, while 462 other similar plants such as Pinus sylvestris have values of 0.056 m.s-1. We must always
  21. 21. ACCEPTED MANUSCRIPT 463 think in terms of intertaxonomic differences if we are able to properly interpret pollen 464 dispersal and deposition patterns. 465 The pollen analysis presented advocates the possibility of an anthropogenic trigger for T 466 Fagus sylvatica expansion. Many other studies suggest that human disturbance P 467 facilitated the expansion of this tree where climatic conditions were favourable (Küster, RI 468 1997). This hypothesis has its origin in northern and north-western Europe (e.g. SC 469 Andersen, 1973; Iversen, 1973), where Fagus sylvatica expanded only after the 470 beginning of the Neolithic (Lang, 1994). According to Tinner and Lotter (2006: 541): 471 472 NU “human activities as one (if not the most important) cause for the invasion of Fagus MA sylvatica into Central Europe (e.g., Küster, 1997, 1999; Ralska-Jasiewiczowa et al., 473 2003) has repeatedly been questioned and is still debated (e.g., Huntley et al., 1989; 474 Lang, 1994; Huntley, 1996; Gardner and Willis, 1999; Pott, 2000)”. In locations where ED 475 Fagus is found forming monospecific communities, it is because in the middle of its PT 476 distribution area young beech has behaved like an eurioic species with a broad 477 ecological valence, capable of shaping itself to edaphic and climatic conditions that are CE 478 relatively diverse (Costa et al., 1998), which gives a certain advantage in confronting AC 479 Abies alba and other colonizers. From this point on it seems logical to think that Fagus 480 sylvatica was occupying the lower part of the Abies alba forest, exactly in the place that 481 was cut and burned to convert the land to cultivated fields. For this reason it did not 482 repopulate and was replaced by hazelnut. This process could only begin in the Middle 483 Ages, with the availability of technologies to occupy the valley floor, the experience 484 necessary to manage the drainage of peat bogs, and the consolidation of fluvial 485 boundaries, in addition to the political capacity to carry out the appropriation of these 486 spaces. In this moment in the history of “slash and burn” agriculture, which means that 487 itinerant agriculture was replaced by the permanent roturation of valley floors, a fact
  22. 22. ACCEPTED MANUSCRIPT 488 that required limiting the diversity of resources available to peasants, who had to 489 specialize in specific products selected not for their productivity but rather for their 490 adaptability to feudal uses (Esteban et al., 2003). T 491 According to the ecological literature, Tinner and Lotter (2006) affirm that Fagus P 492 sylvatica and Abies alba have similar environmental requirements. These authors have RI 493 put on record that 1) today, Abies alba is more competitive than Fagus sylvatica where SC 494 summer precipitation is higher and temperature is lower (Ellenberg, 1996) and 2) 495 palaeobotanical evidence suggests that high summer precipitation is more important for 496 497 NU Abies alba than low temperatures. If we analyse the distribution of Abies alba in the MA Spanish National Forest Inventory, we see how the silver fir on the Iberian Peninsula 498 today live with a mean annual precipitation of about 1100 mm/year and an estimated 499 mean annual temperature between 3.5ºC and 10.5ºC (Ninyerola, 2001). In the Iberian ED 500 Peninsula, young beech stands are found in zones in which the monthly average PT 501 temperatures fluctuate very little between the coldest and warmest month. Normally this 502 change does not exceed 15 ºC, although it might reach 25 ºC in the middle of the CE 503 peninsula. Young beech has great resistance to cold during the fallow times. AC 504 The present-day suitability maps of Abies, Fagus and Abies-Fagus mixed forest can be 505 observed in Fig. 6 and Fig. 7. If we focus on the area closest to the studied peat bog, we 506 find low (<0.3) and intermediate (0.3-0.7) Fagus suitability values. The closest nucleus 507 with high suitability (>0.7) is found about 10 km east of the bog. In contrast, with 508 respect to Abies we can see that cells with intermediate values dominate and, most of 509 all, less than 2 km away we find abundant areas that are highly appropriate for this 510 species. This makes one think that the studied area, and nearby zones, have topoclimatic 511 characteristics that are more favourable to the development of Abies. This situation is in 512 accord with the interpretation of the pollen diagram (Fig. 5), which makes us think that
  23. 23. ACCEPTED MANUSCRIPT 513 Abies recovers more readily when topoclimatic factors outweigh anthropic ones. From 514 the point of view of plant suitability, we can consider the València d’Àneu peat bog as 515 located in an area where the influence of ideal zones for Abies is clearly higher than for T 516 Fagus. Statistical details of the model (adjustment and validation) underlying this P 517 cartography can be found in table 3. RI 518 [FIGURE 6] SC 519 [FIGURE 7] 520 [TABLE 3] 521 522 NU MA 5. Conclusions (A) 523 The València d’Àneu peat bog has been shown to be a good palaeoenvironmental 524 record, giving us an image of the short-term changes that make possible a study of the ED 525 abrupt anthropic effects. The pollen analysis has made evident, in no uncertain terms, a PT 526 possible selective action affecting Abies alba forest in the Roman period and confirmed 527 the indicators that explain how during the medieval period some dynamics began that CE 528 would evolve during the Modern Age and the 20th century to produce the current AC 529 landscape in this area. The current vegetation landscape of this region of the Pyrenees 530 has never before existed over the course of the last 2000 years and the climatic frame is 531 not well represented due to human disturbance of the landscape during this period. 532 The surroundings of the peat bog provided good conditions for human settlement and 533 pastures by removing forest. The palynological data support that human impact became 534 stronger and reached its maximum in the last millennium. 535 A direct climatic inference cannot be made. It is not possible to isolate the human 536 presence from the plant dynamics and therefore there can be no clear correlation during 537 this period between climate and original vegetation.
  24. 24. ACCEPTED MANUSCRIPT 538 Silver fir shows a decline in this area due to factors much more related to human 539 intervention than to climate. Likewise, Abies recovers with a certain ease, in contrast to 540 what happens in other parts of the Pyrenees and pre-Pyrenees; a higher suitability with T 541 respect to its current habitat is evidenced. P 542 During the first millennium of our era, we note the presence of beech woods, most RI 543 likely coexisting with Abies alba as a product of continual and selective actions in the SC 544 forest. Fagus sylvatica acts as a colonizer of open space and can be? directly related 545 with human activity, especially since the Middle Ages, provoking a change in the 546 547 NU altitude limits of forest and other ecotonic zones. In the same way, the maximum levels MA of Corylus avellana currently present are due to the colonization of humid lowlands 548 previously used for crops and pasture. 549 In summary, then, plant succession over the past two millennia in the studied area can ED 550 be described as a maximum extension of Abies alba forest (2200-2000 cal. yr BP); PT 551 selective actions affecting the silver fir forest and arrival of beech (2000-1300 cal. yr 552 BP); deforestation as the agricultural zone expanded, with a reduction in the upper CE 553 altitude limit of the forest and definitive disappearance of Fagus sylvatica (1300-800 AC 554 cal. yr BP); total Abies alba deforestation (800-650 cal. yr BP) and the recovery of 555 silver fir forest (without Fagus sylvatica presence) that, with various fluctuations, 556 persists into the present. 557 558 Acknowledgments (A) 559 This research would not have been possible without the support received from those 560 responsible for the High Pyrenees Natural Park; we especially want to acknowledge 561 Agustí Esteban Amat for his sensitivity to environmental research and his knowledge of 562 the area. Sampling of the peat bog was possible thanks to the efforts of Aureli Carnicer
  25. 25. ACCEPTED MANUSCRIPT 563 and also of COPCISA, which authorized access under the supervision of María Álvarez, 564 to whom we are especially grateful for the assistance she provided. We also wish to 565 gratefully acknowledge the unselfish collaboration in the field that we received from T 566 Riker Yll and Jordi Llorens. Finally, the authors thank Elaine Lilly, Ph.D., of Writer’s P 567 First Aid for English translation and revision. RI 568 SC 569 References (A) 570 Andersen, S.T., 1973. The differential pollen productivity of trees and its significance 571 572 NU for the interpretation of a pollen diagram from a forested region. In: Birks, H.J.B., West, R. G., (Eds). Quaternary Plant Ecology Blackwell, Oxford, 109-115. MA 573 Andrieu, V., Ponel, P., Jul, A.J.T., Beaulieu, J.L. de., Bruneton, H., Leveau, P., 2000. ED 574 Towards the reconstruction of the Holocene vegetation history of lower Provence: 575 two new pollen profiles from Marais des Baux. Vegetation History and PT 576 Archaebotany 9, 71-84. CE 577 Beaulieu, JL de., 1978. Contribution pollenanalytique à l'histoire tardiglaciaire et 578 holocène de la végétation des Alpes méridionales françaises. Ph D Thesis. Université AC 579 Aix-Marseille 3. 580 Beaulieu, JL de, Miras, Y., Andrieu-Ponel, V., Guiter, F., 2005. Vegetation dynamics in 581 north-western Mediterranean regions. Instability of the Mediterranean bioclimate. 582 Plant Biosystems 139(2), 114-126. 583 Bottema, S., 1980. On the history of the walnut (Juglans regia L.) in southeastern 584 Europe. Acta Bot. Neerl. 29, 343-349. 585 Burjachs, F., 1994. Palynology of the upper Pleistocene and Holocene of the north-east 586 Iberian Peninsula: Pla de l'Estany (Catalonia). Historical Biology 9, 17-33.
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  28. 28. ACCEPTED MANUSCRIPT 636 Goeury, C., Beaulieu, JL de., 1979. A propos de la concentration du pollen a l’aide de la 637 liqueur de Thoulet dans les sédiments minéraux. Pollen et Spores 21, 239-251. 638 Hewitt, G.M., 1999. Post-glacial re-colonization of European biota. Biological Journal T 639 of the Linnean Society 68, 87-112. P 640 Huntley, B., 1996. Quaternary paleoecology and ecology. Quaternary Science Reviews RI 641 15, 591–606. SC 642 Huntley, B., Bartlein, P.J., Prentice, I.C., 1989. Climatic control of the distribution and 643 NU abundance of beech (Fagus L.) in Europe and North America. Journal of 644 Biogeography 16, 551–560. MA 645 Iversen, J., 1973. The development of Denmark’s Nature since the Last Glacial. 646 Danmarks Geologiske Underso gelse 7C, 1–126. ED 647 Jalut, G., 1974. Évolution de la végétation et variations climatiques durant les quinze 648 derniers millénaires dans l'extrémité orientales des Pyrénées. Ph D Thesis, Université PT 649 Paul Sabatier. CE 650 Jalut, G., 1984. L'action de I'homme sur la forêt montagnarde des Pyrénées ariégeoise et 651 orientales depuis 4000 B.P. d'après I'analyse pollinique. Actes du 106e Congrès AC 652 national des sociétés savantes, 163-172. 653 Jalut, G., Delibrias, G., Dagnac, J., Mardones, M., Bouhours, M., 1982. A 654 palaeoecological approach to the last 21,000 years in the Pyrenees: the peat bog of 655 Freychinede (Alt. 1350 m., Ariege, South France). Palaeogeography, 656 Palaeoclimatology, Palaeoecology 40: 321-359. 657 Jalut, G., Galop, D., Belet, J.M., Aubert, S., Esteban, A., Bouchette, A., Dedoubat, J.J.I., 658 Fontugne, M., 1998. Histoire des forêts du versant nord des Pyrénées au cours des 659 30000 dernières années. J. Bot. Soc. Bot. Fr. 5, 73-84.
  29. 29. ACCEPTED MANUSCRIPT 660 Kenla, J.V., Jalut, G., 1979. Déterminisme anthropique du développement du hêtre dans 661 la sapinière du Couserans (Pyrénées ariègeoises, France) durant le subatlantique. 662 Geobios 12 (5), 735–738. T 663 Konnert, M., Bergmann, F., 1995. The geographical distribution of genetic variation of P 664 silver fir (Abies alba, Pinaceae) in relation to its migration history. Plant Systematics RI 665 and Evolution 196, 19-30. SC 666 Küster, H., 1994. The economic use of Abies wood as timber in central Europe during 667 Roman times. Vegetation History and Archaeobotany 3 (1), 25-32. 668 NU Küster, H., 1997. The role of farming in the postglacial expansion of beech and MA 669 hornbeam in the oak woodlands of central Europe. The Holocene 7 (2), 239–242. 670 Küster, H., 1999. Prehistoric farming and the postglacial expansion of beech and ED 671 hornbeam: a reply to Gardner and Willis. The Holocene 9 (1), 121–122. 672 Lang, G., 1994. Quartäre Vegetationsgeschichte Europas. Methoden und Ergebnisse. PT 673 Gustav Fischer, Jena (462 pp). CE 674 Le Flao, A., 2005. Apports des reconstitutions cartographiques à l’histoire de 675 AC l’environnement. Essai de spatialisation de données palynologiques pyrénéennes. 676 Diplôme d’Etudes Approfondies. Environnenment et Paysage. Université Toulouse- 677 Le Mirail – Institut Daniel Faucher. 678 Liepelt, S., Cheddadi, R., Beaulieu, J.L. de, Fady, B., Gömöry, D., Hussendörfer, E., 679 Konnert, M., Litt, T., Longauer, R., Terhürne-Berson, R., Ziegenhagen, B. 2009. 680 Postglacial range expansion and its generic imprints in Abies alba (Mill.) – A 681 synthesis from palaeobotanic and genetic data. Review of Palaeobotany and 682 Palynology 153, 139-149.
  30. 30. ACCEPTED MANUSCRIPT 683 López- Merino, L., López-Sáez, J.A., Ruiz, M.B., Gil, M.J., 2008. Reconstructing the 684 history of beech (Fagus sylvatica L.) in the north-western Iberian Range (Spain): 685 From Late-Glacial refugia to the Holocene anthropic-induced forests. Review of T 686 Palaeobotany and Palynology 152, 58-68. P 687 López-Sáez, J.A., Van Geel, B., Martínez-Sánchez, M., 2000. Aplicación de los RI 688 microfósiles no polínicos en Palinología Arqueológica. In: Oliveira Jorge, V., (Ed.), SC 689 Contributos das Ciências e das Technologias para a Arqueologia da Península 690 Ibérica. Actas 3º Congresso de Arqueologia Peninsular, vol. IX, Vila-Real, Portugal, 691 Adecap, Porto, 11-20. NU MA 692 Lozano, P., Meaza, G., Cadiñanos, J.A., 2002. Paleobiogeografía cultural de la reserva 693 de la biosfera de Urdaibai (Vizcaya). Boletín de la AGE. Geografía cultural 34, 193- ED 694 211. 695 Magri, D., Vendramin, G.G., Comps, B., Dupaunloup, I., Geburek, T. , Gömöry, D., PT 696 Latalowa, M., Litt, T., Paule, L., Roure, J.M., Tantau, I., Van der Knapp, W.O., Petit, 697 CE R.J., De Beaulieu, J.-L., Jackson, S.T., 2006. A new scenario for the Quaternary 698 history of European beech populations. palaeobotanical evidence and genetic AC 699 consequences. New Phytologist 171 (1), 199-221. 700 Magri, D., 2008. Patterns of post-glacial spread and the extent of glacial refugia of 701 European beech (Fagus sylvatica). Journal of Biogeography 35 (3), 450-463. 702 Martín-Puertas, Valero-Garcés, B., González-Sampériz, P., Bao, R., Moreno, A., 703 Stefanova, V., 2008. Arid and humid phases in southern Spain during the last 4000 704 years: the Zoñar Lake record, Cordoba. The Holocene 18 (6), 907–921. 705 Miras, Y., Ejarque, A., Riera, S., Palet, J.M., Orengo, H., Euba, I., 2007. Dynamique 706 holocène de la végétation et occupation des Pyrénées andorranes depuis le
  31. 31. ACCEPTED MANUSCRIPT 707 Néolithique ancien, d’après l’analyse pollinique de la tourbière de Bosc dels 708 Estanyons (2180 m, Vall del Madriu, Andorre). C.R. Palevol 6, 291-300. 709 Montserrat, J.M., 1992. Evolución glaciar y postglaciar del clima y la vegetación en la T 710 vertiente sur del Pirineo. Estudio Palinológico. Zaragoza. Instituto Pirenaico de P 711 Ecología - C.S.I.C. RI 712 Muller, U.C, Klotz, S., Geyh, M. A., Pross, J., Bond, G.C., 2005. Cyclic climate SC 713 fluctuations during the last interglacial in central Europe. Geology 33, 449-452. 714 Nakagawa, T., 1998. Etudes palynologiques dans les Alpes françaises centrales et 715 716 NU méridionales. Histoire de la végétation tardiglaciaire et holocene. Ph D Thesis. MA Université d'Aix-Marseille 3. 717 Nakagawa, T., Beaulieu, J.L. De, Kitagawa, H., 2000. Pollen-derived history of timber 718 exploitation from the Roman period onwards in the Romanche valley, central French ED 719 Alps. Vegetation History and Archaeobotany 9, 85-89. PT 720 Ninyerola, M., 2001. Modelització climàtica mitjançant tècniques SIG i la seva 721 aplicació a l’anàlisi quantitativa de la distribució d’espècies vegetals a l’Espanya CE 722 Peninsular. Ph D. Thesis. Universitat Autònoma de Barcelona, AC 723 http.//www.tesisenxarxa.net/TDX-0618101-111736. 724 Ninyerola, M., Pons, X., Roure, J.M., 2000. A methodological approach of 725 climatological modelling of air temperature and precipitation through GIS 726 techniques. International Journal of Climatology 20, 1823-1841. 727 Ninyerola, M., Sáez, L., Pérez-Obiol, R., 2007a. Relating postglacial relict plants and 728 Holocene vegetation dynamics in the Balearic Islands through field surveys, pollen 729 analysis and GIS modelling. Plant Biosystems 141 (3), 292-304.
  32. 32. ACCEPTED MANUSCRIPT 730 Ninyerola, M., Pons, X., Roure, JM., 2007b. Objective air temperature mapping for the 731 Iberian Peninsula using spatial interpolation and GIS. International Journal of 732 Climatology 27: 1231-1242. T 733 Ninyerola M, Pons X and Roure JM., 2007c. Monthly precipitation mapping of the P 734 Iberian Peninsula using spatial interpolation tools implemented in a Geographic RI 735 Information System. Theoretical and Applied Climatology 89: 195-209. DOI: SC 736 10.1007/s00704-006-0264-2. NU 737 Pèlachs, A., 2005. Deu mil anys de geohistòria ambiental al Pirineu Central Català. 738 Aplicació de tècniques paleogeogràfiques per a l’estudi del territori i el paisatge a la MA 739 Coma de Burg i a la Vallferrera. Ph D Thesis. Universitat Autònoma de Barcelona. 740 http.//www.tdx.cesca.es/TDX-0119105-162806/#documents. ED 741 Pèlachs, A., Soriano, J.M., 2003. Las fuentes paleobotánicas y la historia forestal. el 742 ejemplo de los valles de la Coma de Burg y Vallferrera (Pallars Sobirà-Lleida). PT 743 Cuadernos de la Sociedad Española de Ciencias Forestales 16, 155-160. CE 744 Pèlachs, A., Soriano, J.M., Nadal, J., Esteban, A., 2007. Holocene environmental 745 history and human impact in the Pyrenees. Contributions to Science 3 (3), 423–431. AC 746 Peñalba, M. C., 1994. The history of the Holocene vegetation in northern Spain from 747 pollen analysis. Journal of Ecology 82, 815-832. 748 Pérez-Obiol, R., 1988. Histoire Tardiglaciaire et Holocène de la végétation de la région 749 volcanique d’Olot (NE Péninsule Ibérique). Pollen et Spores 30 (2), 189-202. 750 Pérez-Obiol, R., Roure, J.M., 1990. Evidència de la regressió recent de les avetoses a 751 partir de les anàlisis pol·líniques. Orsis 5, 5-11. 752 Pérez-Obiol, R., Julià, R., 1994. Climatic change on the Iberian Peninsula recorded in a 753 30,000-yr pollen record from Lake Banyoles. Quaternary Research 41, 91-98.
  33. 33. ACCEPTED MANUSCRIPT 754 Pérez-Obiol, R., Sadori, L., 2007. Similarities and dissimilarities, synchronisms and 755 diachronisms in the Holocene vegetation history of the Balearic Islands and Sicily. 756 Vegetation History and Archaeobotany 16 (4), 259-265. T 757 Planchais, N., 1985. Analyse pollinique du remplissage holocène de la lagune de Canet P 758 (plaine du Roussillon, département des Pyrénées-orientales). Ecologia Mediterranea RI 759 11, 117–127. SC 760 Pott, R., 2000. Palaeoclimate and vegetation—long-term vegetation dynamics in central 761 Europe with particular reference to beech. Phytocoenologia 30, 285–333. 762 NU Ralska-Jasiewiczowa, M., Nalepka, D., Goslar, T., 2003. Some problems of forest MA 763 transformation at the transition to the oligocratic/Homo sapiens phase of the 764 Holocene interglacial in northern lowlands of central Europe. Vegetation History and ED 765 Archaeobotany 12 (4), 233–247. 766 Reille, M., Andrieu, V., 1991. Données nouvelles sur l’histoire postglaciaire de la PT 767 végétation des Pyrénées occidentales (France). C.R. Acad. Sci. Paris 312, 97-103. CE 768 Reille, M., Loewe, J. J., 1993. A re-evaluation of the vegetation history of the eastern 769 Pyrénées (France), from the end of the last Glacial to present. Quaternary Science AC 770 Reviews 12, 47-77. 771 Riera, S., Wansard, G., Julià, R., 2004. 2000-year environmental history of a karstic 772 lake in the Mediterranean Pre-Pyrenees. the Estanya lakes (Spain). Catena 55, 293- 773 324. 774 Riera, S., López-Sáez, J. A, Julià, R., 2006. Lake responses to historical land use 775 changes in northern Spain. The contribution of non-pollen palynomorphs in a 776 multiproxy study. Review of Palaeobotany and Palynology 141, 127-137. 777 Rivas, S., 1987. Memoria del mapa de series de vegetación de España. ICONA. 778 Madrid.
  34. 34. ACCEPTED MANUSCRIPT 779 Talma, A. S, Vogel, J. C., 1993. Radiocarbon 35 (2), 317-322. 780 Tantinyà, M., 2007. L’acció antrópica i la dinàmica climàtica en l’evolució del paisatge 781 vegetal de la serra del Catllaràs. Master’s degree research project. Universitat T 782 Autònoma de Barcelona. P 783 Terhürne-Berson, R., Litt, T., Cheddadi, R., 2004. The spread of Abies throughout RI 784 Europe since the last glacial period. combined macrofossil and pollen data. SC 785 Vegetation History and Archaeobotany 13, 257-268. 786 Tinner, W., Condedera, M., Ammann, B., Lotter, A.F. 2005. Fire ecology north and 787 788 NU south of the Alps since the last ice age. Holocene 15, 1214-1226. MA Tinner, W., Lotter, AF., 2006. Holocene expansions of Fagus sylvatica and Abies alba 789 in Central Europe: where are we after eight decades of debate? Quaternary Science 790 Reviews 25, 526-549. ED 791 Van Geel, B., Coope, G.R., Van der Hammen, T., 1989. Palaeoecology and stratigraphy PT 792 of the lateglacial type section at Usselo (The Netherlands). Review of Palaeobotany 793 and Palynology 60, 25-129. CE AC
  35. 35. ACCEPTED MANUSCRIPT 794 Fig. 1. Location of the València d’Àneu (VAL-III) peatbog (star) and fir forest (in grey) 795 in the Pyrenees. 796 T 797 Fig. 2. Current distribution of Abies alba (white dots) in the Pyrenees and first P 798 occurrences and dynamics during the Holocene (Pérez-Obiol, 1988; Pèlachs et al., RI 799 2007). SC 800 Fig. 3. Lithologic column and sediment structure of the peat bog. 801 802 NU Fig. 4. Main taxa pollen diagram and calibrated dates from the València d’Àneu (VAL- MA 803 III). 804 ED 805 Fig. 5. 806 Left: Non Arboreal Pollen Concentration vs. Arboreal Pollen Concentration (pol/g). PT 807 Right: Arboreal Pollen Concentration of Abies alba and Fagus sylvatica. Peat bog of 808 CE València d’Àneu (VAL-III) 809 AC 810 811 Fig. 6. Suitability maps of Abies alba (a) and Fagus sylvatica (b). Dots represent the 812 present observed distribution (National Forest Inventory). High, medium and low 813 suitability are denoted by black, grey and white tones, respectively. 814 815 Fig. 7. Suitability map of mixed Abies-Fagus. This map is based on the layered 816 combination of suitability maps of each species. Black colours represent areas where 817 both species have high suitability, grey tones where only one has high suitability and 818 white colour where there is no suitability.
  36. 36. ACCEPTED MANUSCRIPT 819 Fig 1 TP RI SC NU MA ED 820 PT CE AC
  37. 37. ACCEPTED MANUSCRIPT 821 Fig 2 TP RI SC NU MA ED PT CE AC 822
  38. 38. ACCEPTED MANUSCRIPT 823 Fig 3 TP RI SC NU MA ED PT CE 824 AC
  39. 39. ACCEPTED MANUSCRIPT 825 Fig 4 TP RI SC NU MA ED PT CE AC 826
  40. 40. ACCEPTED MANUSCRIPT 827 Fig 5 TP RI SC NU MA 828 ED PT CE AC
  41. 41. ACCEPTED MANUSCRIPT 829 Fig 6 TP RI SC 830 NU MA ED PT CE AC
  42. 42. ACCEPTED MANUSCRIPT 831 Fig 7 TP RI SC NU MA ED PT CE 832 AC
  43. 43. ACCEPTED MANUSCRIPT 833 Sample Laboratory Material Conventional Dating calibrated to Intercept calibration (cm) Code dating BP 2σ (95% curve probability) 59-60 Beta- Wood 780±40 cal BP (730-680) cal BP 690 T 240388 165- Beta- Peat 1990±50 cal BP (1990-1880) cal BP 1940 P 166 240387 RI 834 835 Table 1. 14C dating of peat bog VAL-III SC NU MA ED PT CE AC
  44. 44. ACCEPTED MANUSCRIPT 836 MX_HOT MT_AN MN_COL PR_WIN PR_SPR PR_SUM PR_AUT RAD_WIN RAD_SPR RAD_SUM RAD_AUT N Fagus 20.6-27.6 6.0-12.9 -5.9-3.1 121-509 166-449 94-330 139-436 338-1310 1978-2804 2656-3137 909-1935 3681 sylvatica Abies 18.2- 26.4 3.5-10.4 -8.1, -1.1 139-385 212-395 186- 369 201-380 159-1345 1734-2799 2457-3133 652- 1968 614 alba 837 T 838 Table 2. Ranging values from the predictors used in the GLM suitability models. The values P 839 presented avoid the lowest and highest 2.5% of values. 840 RI SC NU MA ED PT CE AC
  45. 45. ACCEPTED MANUSCRIPT 841 R2 Cut-off Predicted CCR fpos fneg Species Nagelkerke point Observed absence presence (a+d)/N b/(b+d) c((a+c) Fagus absence 669 (a) 65 (b) 0.30 sylvatica 0.84 presence 53 (c) 685 (d) 92% 0.09% 0.07% 0.91 absence 105 (a) 12 (b) Abies alba 0.55 presence 7 (c) 121 (d) 92% 0.09% 0.06% T 842 P 843 Table 3. Fitting and validation results from the Abies and Fagus suitability GLM models. CCR RI 844 (Correct classification rate) represents the general performance of the model. Fpos (False SC 845 positive rate) shows the percentage of suitability areas that do not match with present-day 846 distribution. This cannot be considered an error because exists suitability for both species. Fneg NU 847 (false negative rate) is the measure that can be considered as the error. MA ED PT CE AC

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