Early Holocene Vegetation in the Ayllón Massif (Central System Range, Spain) Based on Macroremains

Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822

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Palaeogeography, Palaeoclimatology, Palaeoecology

journal homepage: www.elsevier.com/locate/palaeo

Early Holocene vegetation in the Ayllón Massif (Central System Range, Spain) based on macroremains. A paleoecological approach

Mar Génova ⁎, Fernando Gómez-Manzaneque, Felipe Martínez-García, José Mª. Postigo-Mijarra

Departamento de Sistemas y Recursos Naturales, Escuela de Ingeniería de Montes, Forestal y del Medio Natural, Universidad Politécnica de Madrid, 28040 Madrid, Spain article info abstract

Article history: We present a palaeoecological study of well-preserved vegetal macroremains in Spain, which age range (10,025– Received 25 May 2015 5371 cal yr BP) confers it an outstanding interest, because for the first half of the Holocene, palaeobotanical data Received in revised form 13 October 2015 are extremely scarce in the Iberian Central System Range. We found, for the first time in the easternmost zone of Accepted 16 October 2015 the Central System Range, macroremains belonging to the genus Pinus; these have specifically been identified as Available online 24 October 2015 Pinus cf. sylvestris and Pinus gr. sylvestris. These findings provide valuable information on the role of the genus

Keywords: Pinus, regarding the natural character of these pine forests in this relatively unknown region. Palaeobiogeography In the larger wood samples we measured tree rings and cross-dated growth series, which represent a 75% increase in Macroremains the number of floating fossil chronologies and increase the time span by almost 1750 years, in the centre of the Dendrochronology Iberian Peninsula. Remarkably, this is the firsttimeinSpainandinsouthernEuropethatsomeofthefloating chro- Spain nologies have been successfully crossdated, creating four composite chronologies. Furthermore, we discuss some Pinus cf. sylvestris palaeoclimatic inferences comparing with different sites in southern Europe and provide new data for best knowing Bos primigenius the palaeoecological characteristics of the first half of the Holocene in Spain. Likewise, we found two bones belonging to aurochs (Bos primigenius), one of these also dating from the first half of the Holocene, with Valdojos constituting the only site with this taxon for this period in the Central System Range and surrounding areas. © 2015 Elsevier B.V. All rights reserved.

1. Introduction et al., 2012). Interestingly, all this research provides only information on the recent past (the last 4000 yr BP), and no study addressing the Inthelastfewdecades,therelevanceofthegenusPinus and the role it Ayllón Massif gives any information on the palaeovegetation existing played in the Iberian ecosystems throughout the Holocene has aroused in the area for the ﬁrst half of the Holocene. The scarce palaeobotanical intense debate and given rise to abundant research (e.g. Carrión et al., information has contributed to generate different interpretations on 2000a, 2010; Franco-Múgica et al., 2000, 2001a; Rubiales et al., 2010; the natural vegetation, especially with regard to the role played by García-Antón et al., 2011). In areas presenting notable anthropic activity, pine forests in this region (Martínez-García and Costa Tenorio, 2001; such as the Central System Range, palaeobotanical research is necessary Martínez-García, 2002). In fact, besides the mentioned palynological in- to understand the nature of changes in the anthropogenic landscapes. formation, different authors have discarded the presence of natural for- Thus, although abundant palynological studies have referred to the Cen- mations of these species in the area, being interpreted the current pine tral System Range, fundamentally in the Estrela, Béjar, Francia, Gredos forest in the region only as a result of recent afforestations (De la and Guadarrama mountains (López-Sáez et al., 2013) the eastern sector Fuente, 1985; Monje-Arenas, 1987; Peinado-Lorca and Martínez Parras, of this large mountain range, known as the Ayllón Massif, remains rela- 1987; Rivas-Martínez, 1987; Rivas-Martínez et al., 2011). On the other tively unexplored in the palaeobotanical sense. hand, any particular species belonging to the genus Pinus has been de- To date, the scarce available information on these mountains consists termined by means the palynological studies. Macroremains studies in of a few palynological studies presenting no accurate chronology the eastern area of this Range may signiﬁcantly help to know which spe- (Hernández-Vera and Ruiz-Zapata, 1984; Jiménez-Ballesta et al., 1985; cies occurred in these pre-anthropic forests. The only studies conducted Ruiz del Castillo, 1993) and some others dated by radiocarbon to date addressing vegetal macroremains in the Central Range refer to (Gil-García et al., 1994; Franco-Múgica et al., 2001a, 2001b; Currás the western sector — the Gredos mountains, Fig. 1 — (Rubiales et al., 2007; Génova et al., 2009; Rubiales and Génova, 2015). Our study of the Valdojos site, in the eastern sector, therefore greatly broadens the ⁎ Corresponding author at: Departamento de Sistemas y Recursos Naturales, Universidad geographical range of research on vegetal macroremains in the centre Politécnica de Madrid, 28040 Madrid, Spain. Tel.:+34 91 3367669. of the Iberian Peninsula. Additionally, the only dendrochronological E-mail addresses: [email protected] (M. Génova), [email protected] (F. Gómez-Manzaneque), [email protected] (F. Martínez-García), studies on macroremains in Spain also involved the above mentioned [email protected] (J.M.ª Postigo-Mijarra). research for the Gredos mountains. Although these constitute a very

Valdojos site

Estrela Francia Ayllón Somosierra Guadarrama

Gata Béjar Gredos

Fig. 1. Situation of the Central and Iberian System Ranges inland on the Iberian Peninsula. The enlarged box on the right shows the different mountains making up the Central System Range, and the location of the site studied is indicated in red. important milestone for the creation of multimillenial chronologies in forests of Quercus pyrenaica Willd., which comprise a belt of deciduous Spain, they involve mainly short floating sequences (Rubiales and trees between 1200 and 1500 m asl. Around this elevation, the oaks Génova, 2015). give way to forests of Scots pine (P. sylvestris L.), which form forests up Consequently, we have pursued the following aims in this work: to approximately 2000 m asl. These pine formations are currently well represented in the more continental sections of the Range (Guadarrama), – To identify the taxa of the remains found and to evaluate their becoming scarcer towards the east (Ayllón) and west (Gredos). Above biogeographical role (spatial and temporal) in the eastern Central 2000 m asl the woody vegetation is represented by a bushy stratum of System Range. Leguminosae (Cytisus, Genista, Echinospartum)orEricaceae(Erica, Calluna, – To study the palaeoecological information provided by the tree ring Arctostaphylos), which at higher altitudes gives way to alpine grassland growth sequences from the dendrochronological perspective. (Luceño and Vargas, 1991; Costa et al., 1997). The E/W and N/S gradients – To analyse the data we have obtained, comparing them with other give rise to local variations in the distribution of the plant communities, pre-existing palaeobotanical information in order to contribute to such as the presence of relict forests. This occurs, for instance, in the designing a vegetation evolution model for the region. Ayllón Massif, with the beech forests of Fagus sylvatica L. The Ayllón Massif makes contact with the Iberian System Range, a mountain range running through the east of the peninsula in the NW– 2. Regional setting SE direction. From the climatic perspective, the Ayllón Massif represents the least Mediterranean zone of the Central System Range, The Central System Range of the Iberian Peninsula constitutes a large it provides refuge for numerous boreal and boreo alpine species mountain range over 500 km long (one of the longest in the Mediterra- (Luceño and Vargas, 1991; Martínez-García, 2001; Ruiz-Labourdette nean Basin), running from West to East. It comprises a succession of et al., 2010) and also explain the presence of P. sylvestris forests and mountain ranges originating in central Portugal (Estrela mountains) the relict forests of beech (F. sylvatica)(Costa et al., 1997). and continuing in Spain: the Gata, Francia, Béjar, Gredos (presenting The Valdojos site (41° 14′ N, 3° 11′ W, 1320 m asl) is located in the the highest point — Almanzor peak, 2592 m asl–), Guadarrama, Ayllón Massif, within the municipality of Campisábalos (Guadalajara Somosierra and Ayllón ranges (Fig. 1). This mountain chain is so long, Province) (Fig. 1) and it comprises a marshy system covering an area with such a broad altitudinal gradient, that it ranges from 300 to over of approximately 5.5 ha. This site is located at the bottom of a valley 2500 m asl; it is highly heterogeneous from the climatic point of view filled with alluvial materials from the Holocene. It is located within a (Ruiz-Labourdette et al., 2010). To simplify, it could be said that the geological context of stratified limestone and dolomites of cretaceous more Mediterranean environments are located on the southern slope origin. In the surrounding area, these carbonated materials form small of the western part of the mountains, whereas the more Eurosiberian gorges by means of dissolution processes. Among the deposits of the ones correspond to the eastern end (Ayllón Massif). On the contrary, fluvial surface formations making up the valley bottom are limestone the more continental environments correspond to the central part boulders and dolomites. The matrix is clayey–sandy of a brown or (Guadarrama mountains). reddish-brown colour. This layer of Holocene alluvial materials is Fromageologicalpointofview,intheCentralSystemRangethereisa quite undeveloped, and does not exceed 4\\5m(Hernáiz et al., 2005). predominance of granites and metamorphic materials, with much more The current vegetation in the immediate vicinity comprises pine localised cretaceous limestone enclaves (Muñoz and Sanz-Herráiz, forests of Pinus sylvestris, with dispersed stands of Juniperus communis 1995). The orography, together with the climatic and geographic varia- L. subsp. hemisphaerica (C. Presl) Nyman, accompanied by typically tion, has provided the Central System Range with a high floristic diversity calcicole species such as Teucrium capitatum L., Thymus leptophyllus (Castro et al., 1996) and vegetation (Martínez-García and Costa Tenorio, Lange subsp.izcoi(Rivas Mart., A Molina & G. Navarro) R. Morales, 2001). The basal forests basically comprise evergreen sclerophyllous for- Satureja intricata Lange, Potentilla cinerea Chaix ex Vill., Sideritis hirsuta mations of Holm oak (Quercus ilex subsp. ballota (Desf.) Samp.), generally L., Teucrium chamaedrys L., Santolina chamaecyparissus L. or Aphyllanthes below 1000–1200 m asl. These are substituted at higher altitudes by oak monspeliensis L. M. Génova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822 813

3. Material and methods Taphonomic distortion can also be observed in some samples, but in general terms, this does not significantly prevent observation of the ana- 3.1. Sampling tomical features. As a result, the wood can be classified as duripartically preserved (Schopf, 1975) or preserved in a basically unaltered state The mechanical work involved in the Valdojos afforestation, conduct- (Spicer, 1991). The displacements described for continental environ- ed in the 1980s (Fernández Muñoz, 2002), uncovered numerous tree ments of this type of macroremains (MacGinitie, 1969; Rich, 1989; remains that have remained on the surface up to the present. We per- Spicer, 1991) point to a native character of these macrofossils for the formed three samplings at the site where these trunks appeared. In the basin (Fernández López, 1990). The presence of disarticulation processes first two of these, in September 2011 and December 2012, we only col- should not be ruled out, especially on the surface of the site, where differ- lected samples that had risen to the surface. In the third one we used a ent afforestation works have been conducted in the recent past. backhoe to make 7 test holes at different points. The firsttesthole(0), in the centre of the valley, provided no samples. The remaining ones, 3.2.1. Radiocarbon dating and calibration test holes 1–6, were made in the lateral zones, where remains had risen We dated a total of 22 wood samples, both from the surface and from to the surface (Fig. 2). The test holes, approximately 3 m wide, reached the 6 test holes made. The ones obtained from the test holes came from a depth ranging from 3.1 m in hole no. 1 to 4.2 m in hole no. 4; in most very different depths, ranging from the maximum of 4.2 m to samples cases, at these depths the transition between the more organic sediment collected on the surface. The radiocarbon dates were obtained from sam- and the underlying mother rock occurred. ples taken from the outer rings of the logs with AMS (Queen's University, We collected all the wood we found from each test hole: a total of 26 Belfast) and were calibrated using the Radiocarbon Calibration Program samples (Fig. 2). If one also counts the remains found on the surface, there (Reimer et al., 2009). Calibrated ages are expressed as the median of the is a total of 46 samples (Table 1). Most of the large-sized samples were higher probability of 2 σ cal yr BP, as suggested by Telford et al. (2004) found on the deposit surface or buried at shallow depths (not over (Table 1). 1.5 m), and at greater depths only small remains were uncovered. More- over, in two of the test holes, no. 2 and no. 3, we found two bones in a 3.2.2. Wood anatomical analysis good state of conservation. We analysed all the wood pieces, both those found on the surface and the ones removed from the test holes (a total of 46 samples), 3.2. Vegetal macroremains using traditional micrographic techniques (Schweingrüber, 1990; García et al., 2002). Fragments measuring approximately 1 × 1 × 2 cm Taking into account the size of the samples and their ring type, were processed with a microtome to provide thin section slides approx- shape and width, they mainly correspond to small and large pieces imately 15–25 μ thick. These slides were stained with safranin and then of trunk. However, branches of different sizes and likely small pieces washed with distilled water and alcohol. Some samples were sub- of roots also are present, all of which were analysed. Although these merged in xylol and fixed with Euquit. The samples were observed by pieces of wood present some evidence of biodegradation, carboniza- means of transmission light microscopy with an Olympus B × 50 series tion and carbonification processes, they were all studied without any optical microscope equipped with UIS lenses (magnification × 50, special treatments. These pieces of wood were found in different ×100, ×200 and ×500). We identified the samples by referring to the positions/orientations in the sediment. No upright trunks were observed. classic wood anatomy studies of Greguss (1955), Jacquiot (1955),

Fig. 2. General view of the site and of the test holes. The image also shows one of the biggest remains collected. The scale of the test holes, trunks and backhoe in relation to the landscape is 1:400. Author: Laura Toro Bermejo. 814 M. Génova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822

Table 1 Number of samples showing its location (test hole or superﬁcial –S–) and depth, conventional and calibrated age and identiﬁcation. *: Calibrated age expressed as the median of the higher probability of 2 σ cal years BP.

Macroremain type ID Test hole/Depth (m) Lab reference Conventional 14C Age (yr BP) Cal Age* Identiﬁcation

1 Wood fragments Val10 S –– –P. gr. sylvestris 2 Wood fragments Val12 S UBA-19753 138 ± 42 225 P. gr. sylvestris 3 Wood fragments Val13 S –– –P. gr. sylvestris 4 Wood fragments Val14 S UBA-19752 105 ± 24 83 P. gr. sylvestris 5 Trunk and wood fragments Val08 S UBA-23910 4712 ± 29 5371 P. cf. sylvestris 6 Trunk Val06 S UBA-23908 5103 ± 39 5791 P. cf. sylvestris 7 Trunk Val03 S –– –P. gr. sylvestris 8 Wood fragment Val21 1/0 –– –P. cf. sylvestris 9 Wood fragment Val20 1/0.7 –– –P. gr. sylvestris 10 Branch Val22 1/1.3 UBA-23911 5445 ± 30 6250 P. gr. sylvestris 11 Trunk Val23 1/1.5 UBA-22236 5556 ± 46 6349 P. cf. sylvestris 12 Wood fragment Val24 1/1.9 –– –P. gr. sylvestris 13 Wood fragment Val26 1/3 –– –P. cf. sylvestris 14 Trunk and wood fragments Val28 1/1 UBA-22238 5574 ± 53 6370 P. gr. sylvestris 15 Trunk Val04 S UBA-19751 5777 ± 33 6577 P. cf. sylvestris 16 Trunk Val30 S –– –P. cf. sylvestris 17 Wood fragments Val31 S –– –P. gr. sylvestris 18 Branch Val32 S UBA-22239 5780 ± 36 6580 P.cf. sylvestris 19 Trunk and wood fragments Val09 S UBA-22247 5832 ± 58 6625 P. gr. sylvestris 20 Trunk and wood fragments Val05 S UBA-22235 5821 ± 37 6632 P. gr. sylvestris 21 Trunk Val07 S UBA-23909 5927 ± 33 6735 P. cf. sylvestris 22 Trunk and wood fragments Val02 S UBA-23907 6175 ± 32 7074 P. cf. sylvestris 23 Trunk and wood fragments Val01 S UBA-22234 6547 ± 44 7470 P. cf. sylvestris 24 Wood fragments Val90 S –– –P. gr. sylvestris 25 Trunk Val80 S –– –P. cf.sylvestris 26 Trunk Val81 6/0.5 UBA-23913 7230 ± 34 8024 P. cf.sylvestris 27 Wood fragments Val82 6/2 –– –P. gr. sylvestris 28 Wood fragments Val83 6/2.3 –– –P. gr. sylvestris 29 Wood fragments Val84 6/2.9 –– –P. gr. sylvestris 30 Wood fragments Val60 4/0.6 –– –P. gr. sylvestris 31 Wood fragments Val61 4/0.9 –– –P. cf.sylvestris 32 Wood fragments Val62 4/1.2 –– –P. cf.sylvestris 33 Wood fragments Val63 4/1.8 –– –P. cf.sylvestris 34 Branch Val64 4/2,1 UBA-23912 7277 ± 42 8093 P. gr. sylvestris 35 Wood fragments Val65 4/3.5 –– –P. gr. sylvestris 36 Wood fragments Val66 4/4 UBA-22242 7384 ± 44 8246 P. gr. sylvestris 37 Wood fragments Val84 6/2.9 UBA-22244 7446 ± 44 8273 P. gr. sylvestris 38 Vertebra B1 3/2.2 UBA-23915 5638 ± 37 6406 Bos primigenius 39 Bark Val50 3/2,2 –– –Pinus sp. 40 Wood fragments Val51 3/3.6 UBA-22241 7529 ± 40 8355 Pinus sp. 41 Wood fragments Val27 1/3.1 UBA-22237 7756 ± 44 8515 P. gr. sylvestris 42 Wood fragments Val40 2/1.8 –– –P. gr. sylvestris 43 Wood fragments Val41 2/2.4 –– –P. gr. sylvestris 44 Metacarpal B2 2/2.4 –– –Bos primigenius 45 Wood fragments Val42 2/2.6 –– –P. gr. sylvestris 46 Wood fragments Val43 2/3.6 –– –P. gr. sylvestris 47 Wood fragments Val44 2/4.2 UBA-22240 7998 ± 48 8855 P. gr. sylvestris 48 Wood fragments Val70 5/3.7 UBA-22243 8890 ± 55 10025 P. cf. sylvestris

Peraza (1964) and Schweingrüber (1990). The samples were also com- 3.2.3. Dendrochronological analysis pared to those in reference collections of extant and fossil wood belong- For the dendrochronological analysis we selected the macroremains ing to the Universidad Politécnica de Madrid (Spain). Diagnosis was that had an appropriate number of rings (14 trunks and two branches) performed by means of a combination of qualitative and quantitative and cut a section in each of them. After surface preparation (sanding), features. We calculated means for features such as height, shape and the ring widths were measured on several radii of the sample (3 to 9, arrangement of the dentations present in the ray tracheids of the depending on shape and size); more radii were measured in irregularly cross-field pits after taking 20 measurements in different parts of the shaped or greater sections. Ring-width measurements were taken to an wood. All anatomically studied samples are housed in the Universidad accuracy of 1/100 mm with a digital LINTAB positioning table connected Politécnica de Madrid (Spain). to a stereomicroscope and TSAPWin software (Rinn, 2003). We employed Some authors indicate that identification at the species level of wood COFECHA (Holmes, 1992)andTSAPWinsoftwareto cross-date and corre- from mountain Iberian pines (including Pinus nigra Arnold, P. sylvestris L. late the chronologies. The series of each radius and section were cross- and P. uncinata Ramond ex DC.)isdifficult and sometimes impossible dated and averaged in an individual chronology corresponding to each (Schweingrüber, 1990; Carcaillet and Vernet, 2001). Nonetheless, other macroremain (Fig. 3). researchers (Greguss, 1955; Jacquiot, 1955; Peraza, 1964)indicatethe A total of 13 of the macroremains dendrochronologically analysed existence of diagnostic features that can be used to this end, especially had previously been radiocarbon dated and the dating had been cali- when young wood is avoided (which can exhibit a great deal of intraspe- brated. For each of the macroremains we applied this calibrated date cific variation) and the macroremains are well preserved (Mutz et al., to the final datum of the individual tree ring growth sequence (the clos- 2004; Figueiral and Carcaillet, 2005). The excellent state of preservation est one to the bark) and we successively dated the rest of the values. of our samples and the fact they were very thick enabled identifications Once we had dated all the individual sequences, we studied the correla- to be made. tions between those of the closest radiocarbon age to adjust the dating M. Génova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822 815

0 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 180 190 200 210 220 230 240 250 260 270 280 290 300 300 300 280 280 260 260 240 Pith 240 220 220

200 7 6 200 180 8 5 180 4 160 9 160 3

140 1 2 140 120 120 100 100 80 80 60 60 Tree ring width ring width (mm Tree x 100) 40 Outer ring 40 20 20 0 Val5. 9 rays measured, maximum radius:19 cm, 0 -20 258 tree rings, mean width ring: 0,72 mm -20 -40 -40

0 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 180 190 200 210 220 230 240 250 260 270 280 290 300 Tree ring number

Fig. 3. An example of the macroremains dendrochronologically analysed (VAL05), in which 9 radii were measured and the growth ring sequences measured were synchronised in order to create an average sequence.

to the sequences selected as a reference for each period and to obtain rest of the buried wood macroremains (36%) are older, up to a maxi- the dendrochronological age (Bradley, 1999; Nicolussi et al., 2009). mum of just over 10,000 cal year BP. Likewise, we analysed the individual mean chronologies that had not been radiocarbon dated, studying the possible temporal range of each 4.2. Wood identification one. When the dendrochronological statistical indicators were sufficiently reliable, we designed composite chronologies. The microscopical analysis performed on 46 samples exhibited the following anatomical features: 3.3. Bones Description. Transverse section: homoxylous wood with easily fi We analysed two bones from a bovid, one metacarpal and one verte- identi able growth rings (Fig. 4B). Polygonal tracheids in cross sec- bra. The metacarpal was found in test hole no. 2 at a depth of 2.4 m. The tion; mean diameter 20 μ. These tracheids are occasionally found vertebra was found in test hole no. 3 at a depth of 2.2 m. Both of them highly compressed as a result of taphonomic distortion processes were in a good state of conservation. Due to the low collagen content of (Fig. 4A). Physiological longitudinal resiniferous channels preferen- the metacarpal, we only dated the vertebra, using AMS (Queen's Univer- tially located in the summer wood and transition zones. Thin-walled sity, Belfast) calibrating it with the Radiocarbon Calibration Program epithelial cells that come away during preparation of thin slides. Radi- fi CALIB 7.0.0 (Stuiver and Reimer, 1993). The samples were identi ed at al section: cross-fields from parenchyma cells to tracheids with one, or the Archaezoology laboratory of the Universidad Autónoma de Madrid rarely two, fenestriform pits (Fig. 4C). Ray tracheids generally with no by the palaeontologist Arturo Morales Muñiz. isolated dentations, normally acute (Figs. 4Ca, 4 Da); length of dentate walls varies from 6 to 10 μ; these tooth-like wall thickenings are occa- 4. Results sionally found connected with other dentations of the opposite side in 4.1. Dating the cell lumen. Tangential section: areolate tracheid pits in uniseriate rows (Fig. 4E). Heterogeneous woody rays uniseriate or pluriseriate, Twenty-two subfossil wood samples were radiocarbon dated; one the latter showing physiological resiniferous canals. Average height half of them was collected from the surface and the remaining ones of rays 5 to 9 cells (Fig. 4F). from the test holes (Fig. 2). In Table 1 the macroremains were ordered Identification. These anatomical features are characteristic of the chronologically according to calibrated age, which ranges from 83 to P. sylvestris group, including P. sylvestris L., P. nigra Arnold and 10,025 cal yr BP. This table also includes the values corresponding to P. uncinata Ramond ex DC. (Greguss, 1955; Schweingrüber, 1990). the bone dated. Although two of the samples are very recent (225 to The set of characteristics such as the lack of remains of epithelial 83 cal yr BP, respectively), most of them date approximately from cells after the preparations and specially, the tooth-like wall thicken- 10,000 to 5000 cal yr BP: the first half of the Holocene, from the Boreal ings, which are pointed, not isolated and occasionally leaving colum- period to the Atlantic one. However, there is not an even distribution of samples, for example the 48% of the samples date between 7500– nar formations, let us to identify our samples as P. sylvestris L. 6000 cal yr BP, including both the samples appearing on the surface (Greguss, 1955; Jacquiot, 1955; Peraza, 1964). Nonetheless, the afore- and those found at shallow depths (between 1 and 1.5 m). Additionally, mentioned discrepancies regarding the reliability of the diagnosis the vertebra (located at a depth of 2.2 m) is also in that age range. The within these species put forward by others authors (Schweingrüber, 816 M. Génova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822

Fig. 4. A, Val-23. Transversal section. Poligonal tracheids compressed due to lithostatic pressure. Scale bar: 20 μ. B, Val-23. Transversal section. Well-deﬁned growth rings with resiniferous canals in the late wood. Scale bar: 250 μ.C,Val-1.Radialsection.Cross-ﬁelds with one or two fenestriform pits. Scale bar: 80 μ. D, Val-1. Radial section. Conspicuous and non isolated tooth- like wall thickenings. Scale bar 40 μ. E, Val-1. Tangential section. Areolate tracheid pits in uniseriate rows. Scale bar 40 μ. F, Val-28. Tangential section. Uniseriate ray. Scale bar: 10 μ.

1990; Carcaillet and Vernet, 2001) suggest to assign tentatively our sequences of the 3 macroremains not radiocarbon dated. Contrasting samples to the taxon P. cf. sylvestris. the radiocarbon calibrated age and the dendrochronological age, the maximum difference obtained was 226 years, which falls within the 4.3. Tree-ring chronologies range of ages calibrated by 2 σ, and we therefore consider it to be quite reliable. The 16 macroremains selected for dendrochronological analysis With the concordant series we designed 4 composite chronologies come from tree trunks, with the exception of two of them, involving (Table 2). We designed composite chronology 1 by successfully branches (Table 1). We measured 18,542 tree rings grouped into 93 synchronising 2 individual chronologies; Composite chronology 2 cor- growth series. The values of intercorrelation among the different responds to samples located in test hole no. 2, which were reliably sequences from each sample are high, and we obtained 16 individual synchronised among each other. Furthermore, other individual chronol- average ﬂoating chronologies. The maximum number of rings from the ogies, based upon macroremains found on the surface, design Compos- individual chronologies ranged from 95 to 429, and 73% had over 200 ite chronology 3. Composite chronology 4 was likely obtained from one tree rings. The vast majority of these remains does not present sapwood single trunk, as a result of disarticulation processes; both sequences pre- or bark, and are therefore most likely from older trees. sented a high level of correlation between growth values, although the In many of the samples, average ring width is very low, ranging from samples do exhibit different state of preservation. The remaining indi- 0.19 to 1.09 mm. Nonetheless, in all cases they correspond to heartwood vidual chronologies are extended over other temporal ranges and/or rings, because only one of the samples presents bark (VAL80), but with were not synchronised well with other chronologies, and these were such deformed sapwood rings that they were impossible to measure. therefore kept as individual ﬂoating chronologies (Table 2 and Fig. 5). The general growth patterns and statistical indicators are habitual for The chronologies cover almost 3000 years (approximately between P. sylvestris in this geographical area (Richter et al., 1991; Génova 8200 and 5800 cal yr BP). In the period 6978–6349 cal yr BP (630 et al., 1993; Génova, 2000, 2012), exceptions being the chronologies of years) there is greater replication, with over 9 individual chronolo- VAL09, corresponding to a sample with two piths, VAL23 with notewor- gies dated (60% of the dendrodated samples), corresponding to the thy release in the central zone, and VAL22, a very old branch, with larger-sized and better-conserved remains and which were mostly decrease followed by release in the central zone. grouped within composite chronologies 2 and 3. However, these We tested the synchronisation of the individual chronologies of sim- composite chronologies were not reliably correlated between each ilar calibrated age, taking as a reference the longest chronology in each other, or with chronologies VAL32 and VAL07, despite being situated group. We also tested the synchronisation of the individual average within coinciding temporal ranges. M. Génova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822 817

Table 2 Samples analysed with dendrochronological methods. *: macroremains found buried between 0.5 and 1.5 m. #CC: identiﬁcation of composite chronologies, IT: intercorrelation, MN tree-rings: maximum number of rings.

ID laboratory Max radius MN Mean tree-ring Std dev Cal BP Age+ Dendrochronological Cal BP age–dendro #CC/IT Floating No. of years reference (cm) tree-rings width Age age chron. BP

VAL08 11.5 364 0.34 0.13 5371 5734–5371 – 1/0.64 5752–5371 382 UBA-23910 VAL03 13 350 0.38 0.18 – 5752–5402 (Ref VAL08) – VAL06 8 304 0.26 0.08 5791 6094–5791 ––6094–5791 304 UBA-23908 *VAL28 9 240 0.42 0.21 6370 6589–6349 (Ref VAL23) 21 2/0.41 6781–6349 433 UBA-22238 *VAL23 12 429 0.27 0.17 6349 6781–6349 – UBA-22236 *VAL22 7 347 0.19 0.09 6250 6719–6367 (Ref VAL23) −117 UBA-23911 VAL32 9 375 0.25 0.17 6580 6954–6580 ––6954–6580 375 UBA-22239 VAL04 17 254 0.89 0.45 6577 6924–6671 (Ref VAL05) −94 3/0.73 6984–6632 347 UBA-19751 VAL05 19 282 0.79 0.34 6632 6913–6632 – UBA-22235 VAL30 10.5 95 1.09 0.35 – 6906–6812 (Ref VAL05) – VAL09 10 128 0.77 0.26 6625 6984–6851 (Ref VAL05) −226 UBA-22247 VAL07 9 128 0.88 0.28 6735 6862–6735 ––6862–6735 128 UBA-23909 VAL02 11.5 237 0.52 0.20 7074 7310–7074 ––7310–7074 237 UBA-23907 VAL01 9.5 180 0.54 0.25 7470 7649–7470 – 4/0.70 7649–7399 251 UBA-22234 VAL80 11.5 245 0.45 0.22 – 7643–7399 (Ref VAL01) – *VAL81 8 133 0.63 0.23 8024 8156–8024 ––8156–8024 133 UBA-23913

4.4. Identiﬁcation of bones 5. Discussion

The good state of conservation of the 2 bones found (one metacarpal 5.1. Relevance of Valdojos site in the Central System Range and geobotanical and one vertebra) enabled us to be attributed to the genus Bos. Anatomic implications analysis of the metacarpal indicates that it is a right-hand metacarpal whose surface presents various striations that would appear to corre- 5.1.1. Occurrence of Pinus sylvestris in the Ayllón Massif in the first half of spond to dragging marks from the diagenetic phase. This bone is likely the Holocene from a sub-adult male. Its morphometric data are the following: maxi- Despite the numerous palaeobotanical studies addressing the Central mum length: 23.8 cm; proximal width: 7.55 cm; minimum diaphysis System Range over the last 50 years, well over one hundred (López-Sáez width: 4.1 cm and distal width: 7.7 cm. The anatomical study conducted et al., 2013), there is still little palaeobotanical information for the first half with the reference collection from the Archaezoology laboratory of of the Holocene and for some areas of the mountain range. the Universidad Autónoma de Madrid and the Consejo Superior de This is the case of the Ayllón Massif. For the first time in the eastern Investigaciones Científicas, reveals that these two remains correspond zone of the Central System Range, the Valdojos site provides vegetation to the taxon Bos primigenius Boj. (Arturo Morales, pers. com.) (Fig. 6). data for the time interval 10,025–5371 cal yr BP. This fact therefore

Tree A Ring width (mm)

B 1 2 1 9 1 2

8200 7800 7400 7000 6600 6200 5800 5400

Fig. 5. Dendrochronologies obtained at the Valdojos site. a) Raw chronologies: individual chronologies in a lighter colour and composite chronologies in a darker colour. b) Temporal ranges of the ﬂoating chronologies, with an indication of the number of average sequences in each range. 818 M. Génova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822

Fig. 6. Left-hand side, right metacarpal in anterior view. Right-hand side: lumbar vertebra in anterior view. demonstrates the continuous presence of forests of P. cf. sylvestris at an pine forests in the zone had been demonstrated by pollen cores in altitude of over 1320 m asl. These data, which are completely new with these zones. Thus, in the above mentioned Pelagallinas sequence, regard to age and taxonomic determination, cannot be compared to any Pinus dominates ca 4000–2400 years BP, reaching percentages of 50% other, as all other research addressing the area presents chronologies and accompanied by up to 30% of Betula (Franco-Múgica et al., 2001b). that are generally much more recent and taxonomically less accurate. Similarly, in the study conducted in the Galve de Sorbe peat bog, close Indeed, the oldest data for the zone to date are from the pollen study to the Valdojos site (see Fig. 7), the authors highlight the fact that the pol- of the Pelagallinas peat bog (sierra de Alto Rey, 1340 m asl; Fig. 7), with len percentages of Pinus are higher than those of other tree taxa through- two datings (3980 ± 90 and 2400 ± 40 years BP) reflecting the vegeta- out the whole profile (Hernández-Vera and Ruiz-Zapata, 1984). Likewise, tion in the last four millennia (Franco-Múgica et al., 2001b). Other stud- Somolinos sequence exhibits high percentages of pine forests (N60%) ca ies exist that provide more recent palaeobotanical information, such as 870–540 cal years BC. the Somolinos (2410 ± 401385 ± 30 years BP; Currás et al., 2012)and Pico del Lobo (1170 ± 80 years BP; Gil-García et al., 1994), whereas 5.1.2. Is Pinus sylvestris native to the Eastern Central System Range? other studies lack absolute datings and correspond to short sequences The Iberian Peninsula has been subjected to intense anthropic action that are also probably very recent (Ruiz del Castillo, 1993; Gil-García for the last few millennia, which has given rise to profound changes in et al., 1994). Our research therefore provides new data on the the vegetal landscape, as an intense deforestation over vast areas. The palaeovegetation for the first half of the Holocene, the whole of Boreal Central System Range has not escaped this process and has also suffered and Atlantic periods, totally unknown to date for the Ayllón Massif. the impact of human activity. Thus, there has been a great reduction of Although only one taxon was found, the presence of P. cf. sylvestris in the area occupied by trees (pine, oak, beech forests, etc.) and conse- the area is relevant, because it helps to resolve the controversy regarding quently the extension of bush and herbaceous cover. Moreover, in the pre-anthropic vegetation in Ayllón, particularly that of the Alto Rey and second half of the XX century, intense afforestation was conducted in Pela mountains and the vicinity of Campisábalos. The presence of ancient Spain (Groome, 1990; Gómez Mendoza, 1992; Carrión et al., 2000b,

Fig. 7. A. Sites cited in the text with palaeobotanical information for the ﬁrst half of the Holocene. 1. Charco da Candieira (Estrela); 2. El Maíllo (Francia); 3. Garganta del Trampal (Béjar). 4. Garganta del Villar (Gredos); 5. Cuerpo de Hombre (Béjar); 6. Hoyos del Espino (Gredos) 7. Baterna (Gredos); 8. Hoyocasero (Gredos); 9. Rascafría (Guadarrama); 10. Valdojos (the present paper). B. Location of the sites cited in the text in the Ayllón Massif. A. Pelagallinas; B. Laguna de Somolinos; C. Galve de Sorbe; D. Pico del Lobo; E. Valdojos (the present paper). M. Génova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822 819

2010; Valbuena-Carabaña et al., 2010). One of the most commonly used and in most cases they analyse pollen sequences; very little information species was P. sylvestris, and 600,000 ha of this taxon were planted be- exists on macroremains (Fig. 7, Table 3). tween 1940 and 1993 (Valbuena-Carabaña et al., 2010). One of the ter- Thus, in the Western sector of the Central System (Estrela, Francia ritories afforested with a large amount of this species is the Central and Béjar mountains), the presence of Pinus may be detected since the System Range. In the Alto Sorbe basin, where the Valdojos site is located, Tardiglacial period, ca. 14,800 cal yr BP, showing the sequences of this afforestation programme was also implemented as from 1948, and Charco de Candieira, Maillo and Garganta del Trampal, a constant de- involved P. sylvestris and Pinus pinaster (Fernández-Muñoz, 2002). crease and relatively low values of Pinus during the firsthalfoftheHo- Due to these deforestation and afforestation processes, the current locene (Fig. 7; Ruiz-Zapata et al., 1989; Atienza et al., 1990; Van der landscape of the Central System Range comprises two types of forest: Knaap and Van Leuwen, 1997; Connor et al., 2012; Morales-Molino those resulting from recent afforestation and others that existed previ- et al., 2013). Regarding macroremains, one needle and stomata collect- ous to these implementations and for which historical documentation ed at Charco de Candieira have been attributed to P. cf. sylvestris and a exists (Montoya Ramírez, 1992; Calvo Sánchez, 2003). number of charcoals from the Maillo peat bog have been related with In parallel to this mass afforestation process, in the final third of the XX P. gr. sylvestris/nigra and P. pinaster/pinea (Table 3; Van der Knaap and century the idea spread among Spanish botanists and forestry engineers Van Leuwen, 1997; Morales-Molino et al., 2013). On the other hand, a that many of the formations dominated by species of Pinus on the Iberian more consolidated pine forest, with a dating close to the base of Peninsula were of anthropic origin. Among these, most of P. sylvestris 5160 ± 40 years BP can be detected in the sequence of Cuerpo de Hom- forests in the Central System Range, which were interpreted bre (Ruiz-Zapata et al., 2011). resulting from older afforestation in the potential ranges of other species The Gredos and Guadarrama mountains also have very few palyno- (Martínez-García and Costa Tenorio, 2001; Martínez-García, 2002). logical sequences to provide information for the first half of the Holo- Within the scientific scope, the idea that most of the current Iberian cene. In this region, the presence of pollen of Pinus sylvestris-type has pine forests are artificial formations has meant that the vast majority of been highlighted in the Garganta del Villar sequence and the Baterna pine forests have been ignored or underestimated in studies on the real core (Dorado-Valiño, 1993; López-Sáez et al., 2009, 2013). These sites, or potential vegetation in the territory. This, for instance, has occurred including the Hoyos del Espino peat bog (Franco-Múgica, 2009), in in the eastern zone of this mountain range where the Valdojos site is which a trunk attributed to P. gr. sylvestris has been found (Génova located (De la Fuente, 1985; Monje-Arenas, 1987; Peinado-Lorca and et al., 2009), reveal a greater predominance of Pinus than the previous Martínez-Parras, 1987; Rivas-Martínez, 1987). We should mention, how- ones in the western sector. Likewise, the Rascafría pollen core (see ever, that some phytosociological studies on the upper zones of Pela Fig. 7), (ca. 8410 ± 250 years BP to present day) shows the presence mountains have recently recognised communities of Scots pine on litho- of P. cf. sylvestris throughout practically the whole Holocene, the tree sols or on very steep shady slopes. According to these studies the decidu- pollen exceeding 90% in the interval between 8410 and 3700 years BP ous species are at a clear competitive disadvantage therein (De la Cruz (Franco-Múgica and García-Antón, 1994; Franco-Múgica et al., 1998). and Peinado, 1996; Rivas-Martínez et al., 2001, 2011). Taken into account the previous data, we can state that pine forests, The data obtained in the present research therefore clearly support the probably related with P. sylvestris, were significant in the vegetal land- natural and ancient origin of the P. sylvestris forests in the eastern sector of scapes during the first half of the Holocene along the western and central the Central System Range, as they unequivocally demonstrate the presence sectors of the Central System Range. Now, the new data from the Valdojos of this taxon in the first half of the Holocene (approx. between 10,000 and site let us to complete this general overview and we can conclude that 5000 years BP). Furthermore, on considering the ecological characteristics of also in the eastern sector of the Central System Range the scots pine the territory, the hypothesis is supported that the pollen sequences of Pinus was widespread over the firsthalfoftheHolocene. obtained from the nearby cores at Pelagallinas and Somolinos (Franco et al., 2001b; Currás et al., 2012), of an approximate age of 4000 and 2400 years BP respectively, also could correspond to P. sylvestris.Moreover,thishy- 5.2. Palaeodendrochronological inferences at Valdojos site pothesis is coherent with the abundant historical information dating from the XIV century (Montoya Ramírez, 1992), which indicates an Within the European framework, sites with dendrochronologically abundance of pine forests in vast areas of the eastern edge of the Central dated fossils are relatively common (e.g. Leuschner et al., 2002; System Range previous to the initial afforestations half-way through the Nicolussi et al., 2009; Kolář and Rybníček, 2011; Kaiser et al., XX century (Fidalgo Hijano, 1987; Martínez-García, 1999, 2002; 2012); in some of them being Pinus sylvestris themorecommon Martínez-García and Costa Tenorio, 2001). taxon (Miramont et al., 2000; Eronen et al., 2002; Gunnarson, 2008). In southern Europe, however, the much less favourable conditions for the conservation of fossil macroremains have hindered the establish- 5.1.3. The Scots pine in the Central System Range from the early to the mid- ment of palaeodendrochronologies. Holocene To date, the southernmost European sites with dendrochronologically In the Iberian Central System Range few palaeobotanical studies studied woods of Pinus sylvestris group are found in the west of Spain's provide data on the vegetation during the first half of the Holocene, central system -the Gredos mountains- (Rubiales et al., 2007; Génova

Table 3 Macroremains found in the Central System Range in the Lateglacial and Early to mid-Holocene related to P. cf. sylvestris or P. gr. sylvestris. N: number of macroremains found. Age: absolute ages expressed in cal yr BP.

Macroremain N Diagnosis Age Site References

Needle 1 P. cf. sylvestris Lateglacial Charco de Candieira Van der Knaap and Van Leeuwen (1997); ca 14,060–12,850 (Serra da Estrela) Connor et al. (2012) Stomata – P. cf. sylvestris Lateglacial Charco de Candieira Van der Knaap and Van Leeuwen (1997); ca 14,060–12,850 (Serra da Estrela) Connor et al. (2012) Charcoal – P. gr. sylvestris Early Holocene El Maíllo Morales-Molino et al. (2013) ca 10,550–9800 (Francia) Wood 1 P. gr. sylvestris Mid-Holocene Hoyocasero Rubiales and Génova (2015) ca 6415 (Gredos) Wood 46 P. cf. sylvestris and P. gr. sylvestris Early and Mid-Holocene Valdojos The present paper ca 10,025–5371 (Ayllón) 820 M. Génova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822 et al., 2009; Rubiales and Génova, 2015). In different sites a total of 23 an unfavourable period for burial and conservation. Nicolussi et al. floating chronologies was obtained between 6415 and 762 cal yr BP (2009) provide further detail and point out that for the period between (Rubiales and Génova, 2015), whereas at the Valdojos site, the 16 chro- 5650 and 5250 cal yr BP, fewer remains have been found, as this time nologies obtained range from 8156 to 5371 cal yr BP. Thus, the informa- coincided with known periods of glacier advances. tion from this new site provides a 75% increase in the number of floating fossil chronologies from the centre of the Iberian Peninsula with 5.3. Bos primigenius Boj. in the Iberian Peninsula in the first half of the an increase of 1750 years in the time record. Furthermore, at Valdojos Holocene the average of the maximum number of rings per sample is around 261 years, over twice as much as the remains located in the Gredos mountains The aurochs (Bos primigenius) was a species with a broad range of (Rubiales and Génova, 2015). The exceptional features of the Valdojos distribution during the Quaternary, from India to north Africa and macroremains enabled us, for the first time in Spain and in southern Europe; its disappearance has been linked to habitat destruction and Europe, to successfully crossdate some of the floating chronologies and overhunting, and it was last detected in Poland in the year 1627 AD to create 4 composite chronologies. Thus, we can highlight the fact that, (Van Vuure, 2005). It reached the European continent from Africa at in the context of southern Europe, the Valdojos site is highly exceptional the start of the Pleistocene (Martínez-Navarro et al., 2007), although from the dendrochronological point of view, and is extraordinarily rich the Iberian Peninsula, as well as the Italic Peninsula, might have consti- due both to the high number of large remains found and studied and to tuted refuge areas for this taxon during this period (Mona et al., 2010). the number, quality and scope of the floating chronologies obtained. The oldest citation for the Peninsula refers to approximately 700 ka Consequently, following our research at Valdojos, there is now a total of (Estévez and Saña, 1999), and its presence was established during the 39 floating chronologies for the whole Central System Range, covering mid-Pleistocene and particularly the upper Pleistocene at different approximately the period 8156–762 cal yr BP (almost 7500 years). sites distributed throughout the Peninsula (e.g. Arsuaga et al., 2012; Although the data from Valdojos are not sufficiently abundant to Burjachs et al., 2012; Évora, 2013). Most occurrences of the taxon on establish palaeoecological conclusions, it should be pointed out that for the Peninsula are from the Holocene, as occurs for Europe, a fact the whole site the most recent fossil macroremains (6700–5300 cal yr that is associated with the improved postglacial climatic conditions, BP) present the narrowest rings (Table 2 and Fig. 5). This means that, which favoured its expansion (Wright, 2013). For the first half of the approximately 6700 years ago in this sector of the Iberian Central System, Holocene, this taxon has been cited in a small number of Iberian sites environmental changes might have occurred affecting the growth of the (Fig. 8). These are: Cueva de Mazaculos II (10013–7503 cal BC; trees. Among others we can highlight a temperature decrease, coinciding Arroyo and Morales, 2009), Mendandia (6550–4490 BC; Castaños, with a phase of glacier advances (Nicolussi et al., 2009). Another possible 2005), La Sierra de Gibijo (6505–5927 cal BC; Altuna, 1974), Cueva interpretation of the poor growth rates at Valdojos in this final stage de Chaves (4820–4170 cal BC; Castaños, 2004) and Arenaza (Meso- might involve a hypothetical increase in detritic accumulation, because lithic; Arias Cabal and Altuna, 1999). Also cited and assigned to the at other sites containing fossil macroremains from alluvial and colluvial Neolithic–Calcolithic are the Fuente Flores (Cabanilles and Valle, deposits, this factor has been associated with periods of abrupt growth 1988)andLaRenke(Altuna, 2001) sites. Therefore, in sites on the decrease (Miramont et al., 2000). Peninsula from between the Mesolithic and the Neolithic, Valdojos The end of the functioning of the sedimentation system at Valdojos is to date the only one presenting aurochs macroremains in the Cen- might coincide with the age of the most recent fossil macroremains tral System Range and the surrounding area of the centre of the (approximately 5300 cal yr BP). This also occurs in the Middle Durance Peninsula. (Southern Alps, France; Miramont et al., 2000), or in the extensive With regard to its habitat, according to Van Vuure (2005) the abun- dendrochronological study conducted in the Eastern Alps (Nicolussi dant findings of aurochs in the Netherlands in river valleys suggest that et al., 2009). In both cases, very few trunks of fossilised trees were it inhabited flatlands near rivers. Hall (2008) has corroborated this found as from 6000 cal yr BP, due to the fact that this was the start of thesis by means of analysis of numerous sites dated as Late Pleistocene

1 3 2 4 5

Fig. 8. Mesolithic and Neolithic sites cited in the text and presenting aurochs. 1. Cueva de Mazaculos II. 2. La Sierra de Gibijo 3. Arenaza. 4. La Renke y Mendandia. 5 Cueva de Chaves. 6. Fuente Flores. 7. Valdojos (the present paper). M. Génova et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 441 (2016) 811–822 821 and Postglacial in Britain, thus confirming the Van Vuure's thesis that the Carrión, J.S., Munuera, M., Navarro, C., Sáez, F., 2000b. Paleoclimas e historia de la vegetación fl cuaternaria en España a través del análisis polínico. Viejas falacias y nuevos paradigmas. aurochs inhabited riverine atlands. Because of its location in a valley Complutum 11, 115–142. bottom, close to a riverbed, the Valdojos site could have presented similar Carrión, J.S., Fernández, S., González-Samperiz, P., Gil-Romera, G., Badal, E., Carrión-Marco, environmental conditions to those described as a preferential habitat for Y., López-Merino, L., López-Sáez, J.A., Fierro, E., Burjachs, F., 2010. Expected trends and surprises in the Lateglacial and Holocene vegetation history of the Iberian Peninsula this species. and Balearic Islands. Rev. Palaeobot. Palynol. 162, 458–475. Castaños, P., 2004. Estudio arqueozoológico de los macromamíferos del Neolítico de 6. Conclusions lacueva de Chaves (Huesca). Salduie 4, 125–171. Castaños, P., 2005. Estudio arqueozoológico de la fauna de Mendandia (Sáseta, Trevino). In: Alday, A. (Ed.), El campamento prehistórico de Mendandia: Ocupaciones mesolíticas y The results obtained from the analysis of the macroremains found at neolíticas entre el 8500 y el 6400 B.P. B9. Fundación Barandiarán, Vitoria-Gasteiz, Valdojos provide highly relevant data for palaeobotanical interpretation pp. 427–448 Castro, I., Moreno, J.C., Humphries, C., Williams, P., 1996. Strengthening the Natural of the eastern sector of the Central System Range throughout the Holo- and National Park system of Iberia to conserve vascular plants. Bot. J. Linn. Soc. 21, cene. They provide accurate information that enables us to reconstruct 189–206. the vegetal landscape of the territory. The taxon P. cf.sylvestrishas been Connor, S.E., Araújo, J., Van der Knaap, W.O., Van Leeuwen, J.F.N., 2012. Along-termperspec- – identified as one of the principal species in these forests during the first tive on biomass burning in the Serra da Estrela, Portugal. Quat. Sci. Rev. 55, 114 124. Costa, M., Morla, C., Sáinz, H. (Eds.), 1997. Los bosques ibéricos. Una aproximación half of the Holocene, which appears to have formed stable and continuous geobotánica. Planeta, Barcelona. forests. This hypothesis seems to be supported by the ecological charac- Currás, A., Zamora, L., Reed, J.M., García-Soto, E., Ferrero, S., Armengol, X., Mezquita-Joanes, teristics of the area and by the abundant historical information. F., Marqués, M.A., Riera, R., Julià, R., 2012. Climate change and human impact in central Spain during Roman times: High-resolution multi-proxy analysis of a tufa lake record From the dendrochronological perspective the Valdojos site is totally (Somolinos, 1280 m asl). Catena 89, 31–53. exceptional within the southern European and Iberian context, provid- De la Cruz, M., Peinado Lorca, M., 1996. El paisaje vegetal de la Cuenca del río Henares. I. fl Comunidades arbóreas y arbustivas. Wad-al-Hayara 23, 335–396. ing a set of extensive oating chronologies of great palaeoecological fi fi De la Fuente, V., 1985. Vegetación oró la del Occidente de la provincia de Guadalajara interest for the rst half of the Holocene. (España). Lazaroa 8, 123–219. Finally, the remains found of Bos primigenius enlarge the scant Dorado-Valiño, M., 1993. Evolución de la vegetación durante el Holoceno en el catalogue of localities presenting this species between the Mesolithic Valle de Amblés (Ávila) (PhD dissertation) Universidad de Alcalá de Henares, Madrid. and the Neolithic on the Peninsula, Valdojos being to date the only site Eronen, M., Zetterberg, P., Briffa, K., Lindholm, M., Meriläinen, J., Timonen, M., 2002. Part 1: of aurochs in the centre of the Peninsula in this age. The supra-long Scots pine tree-ring record for northern Finnish Lapland; chronology construction and initial inferences. The Holocene 12 (6), 673–680. Estévez, J., Saña, M., 1999. Auerochsenfunde auf der Iberischen Halbinsel. In: Acknowledgments Weniger, G.-C. (Ed.), Archäologie und Biologie des AuerochsenWissenschaftliche Schriften des Neanderthal Museum 1. Köln, pp. 119–131. Évora, M., 2013. Osseous industry and exploitation of animal resources in Southern Iberia The present paper is dedicated to the memory of our friend and – fi during the Upper Palaeolithic. Quat. Int. 318, 33 45. colleague Ru no García, who passed away in 2011 and who initiated Fernández-López, S., 1990. El significado de la autoctonía/aloctonía tafonómica. In: this research with us. We are grateful to Carlos Morla for his continuous Fernández-López, S. (Ed.), Comunicaciones de la Reunión de Tafonomía y Fosilización. assistance. We also wish to thank Arturo Morales Muñiz for his help in Departamento Paleontología, Univ. Complutense, Madrid, pp. 115–124. fi Fernández-Muñoz, S., 2002. Consecuencias socioeconómicas y territoriales de the identi cationofthebones.Weverymuchappreciatetheinformation las repoblaciones forestales en el Alto Sorbe (Guadalajara). Ería 58, referring to the site and the support provided by Joaquín Castelo, Gregorio 183–203. Cerezo and Juan Pablo Santo Domingo de Marcos. 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