Glacial Chronology, Environmental Changes and Implications for Human Occupation During the Upper Pleistocene in the Eastern Cantabrian Mountains

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Quaternary International 364 (2015) 22e34

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Glacial chronology, environmental changes and implications for human occupation during the upper Pleistocene in the eastern Cantabrian Mountains

* Enrique Serrano a, , Manuel Gomez-Lende a, María Jose Gonzalez-Amuchastegui b, María Gonzalez-García a, Juan Jose Gonzalez-Trueba c, Ramon Pellitero d, Ibai Rico b a Universidad de Valladolid, GIR PANGEA, Dpt. Geografía, Paseo Prado de la Magdalena s/n, 47011, Valladolid, Spain b Universidad del País Vasco, Dpt. Geografía, Prehistoria y Arqueología, Francisco Tomas y Valiente s/n, 01006, Vitoria-Gasteiz, Spain c CIESE-C, Universidad de Cantabria, 39520, Comillas, Spain d University of Aberdeen, Dept.of Geography & Environment, School of Geosciences, Aberdeen, Scotland, AB24 3UF, UK article info abstract

Article history: Interest in the evolution of Pleistocene glaciers, environmental change and human occupation in southern Available online 14 October 2014 Europe is now growing thanks to the availability of fresh dating in different mountains. The demise of Ne- anderthals in the north of the Iberian Peninsula and the expansion of anatomically modern humans took place Keywords: between 40 and 30 ka (MIS3) under cold and wet climatic conditions. Hypotheses differ regarding the Glaciations migration and demise of Neanderthals, relating to environmental, human or cultural factors. The aim of this Late Quaternary study is to establish glacial evolution in the eastern Cantabrian Mountains related to environmental changes MIS3 and climatic evolution during the MIS3 and MIS2 and human occupation including the Neanderthals' demise. Human occupation Cantabrian Mountains The study is based on glaciomorphology by glacial reconstructions, morphostratrigraphic sequences, equilibrium line altitude estimation, numerical dating (14C, AMS, and OSL), palaeoclimate sequences and human occupation. Three glacial stages during MIS3 and MIS2 have been established. S-I took place during MIS3 in a cold and wet environment associated with snowfall coming from cold, humid masses from the N, NW and SW and available dates place S-I at 42e31 ka. S-II was characterised by shorter and thicker glaciers, which point to a cold, dry and more stable climate, and this is coetaneous with the European Last Glacial Maximum. S-III was a brief period, still poorly dated but attributed to the Late Glacial, characterised by small glaciers housed at altitude and absent from lower mountains. A lack of harmony has been detected between environmental cooling and population and this may have been a response to environmental changes in N and NW Europe. Neanderthals disappeared from the Cantabrian region at the time of the S-I glacial advance during a cold and wet period at the end of the MIS3. If they had adapted to environmental changes, including abrupt cold events (H6, H5) with important environmental consequences, factors other than environmental ones may have been the cause. The weight of the cultural hypothesis seems more likely in view of the environmental changes, glacial evolution and the continuity of oceanic climates. © 2014 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction thanks to the availability of new dating in different mountains (Perez-Alberti et al., 2004; Hughes and Woodward, 2008; García- Interest in Pleistocene glaciers in the southern European context Ruíz et al., 2010; Palacios et al., 2011; Jimenez et al., 2013). In the and the chronology of the different glacial stages is now increasing Iberian Peninsula considerable evidence has been collected of glacial stages prior to the Last Glacial Maximum (LGM) in the north * Corresponding author. of Europe in both Atlantic and Mediterranean environments E-mail addresses: [email protected], [email protected] (E. Serrano), (Jimenez et al., 2002, 2013; García-Ruíz et al., 2003, 2010; Perez- [email protected] (M. Gomez-Lende), [email protected] Alberti et al., 2004, 2011; Gonzalez-Samp eriz et al., 2006; Jalut (M.J. Gonzalez-Amuchastegui), [email protected] (M. Gonzalez- et al., 2010; Moreno et al., 2010; Serrano et al., 2012, 2013). The García), [email protected] (J.J. Gonzalez-Trueba), r.pellitero- [email protected] (R. Pellitero), [email protected] (I. Rico). present day controversy is on the possible synchrony between the http://dx.doi.org/10.1016/j.quaint.2014.09.039 1040-6182/© 2014 Elsevier Ltd and INQUA. All rights reserved. E. Serrano et al. / Quaternary International 364 (2015) 22e34 23 peninsular and European LGM in the Pyrenees or the Central Sys- direction (Fig. 1). The western area is formed by the Asturian base- tem (Pallas et al., 2006; Calvet et al., 2011; Palacios et al., 2011). ment of slates, quartzites, limestones, sandstones and conglomerate There is a long history of studies on Pleistocene glacial stages in the of Devonian and Carboniferous age. The complex structure and dif- Cantabrian Mountains, which were first attributed to two Quaternary ferential erosion built up isolated massifs on limestones, quartzites glaciations, Riss and Würm (Penck, 1897; Obermaier, 1914; Saenz, and sandstones and eroded depressions on slates and turbidites. 1935; Hernandez-Pacheco, 1944, 1961; Nussbaum and Gigax, 1953; Rivers deeply eroded and cut structures on the north side, while Lotze, 1962) and later to a single recent Quaternary glacial period adapting to structures in the south side. The eastern area was covered (Martínez de Pison and Arenillas,1979;Alonso et al.,1982; Smart,1986, by Mesozoic outcrops: Triassic sandstones and conglomerates, 1998; Frochoso, 1990; Castan~on and Frochoso, 1992; Gale and Hoare, Jurassic and Cretaceous marls, sandstones, limestones, and turbi- 1997; Serrano and Gonzalez-Trueba, 2002; Serrano and Gutierrez, dites. The climate is governed by the exposure of mountain ranges to 2002; Gonzalez-Trueba, 2007a, 2007b). New dates and palaeoglacial the Polar Front, their relative height and proximity to the sea. The reconstructions (Moreno et al., 2010; Jalut et al., 2010; Serrano et al., north side has a hyperhumid oceanic mountain environment with 2012, 2013) show that the glacial maximum of the Cantabrian over 2500 mm/year of rainfall, whereas to the south the climate is Mountains may have been prior to the European LGM. All dates point transitional Atlantic-Mediterranean, with around 600 mm/year of to a distinct behaviour of Cantabrian glaciers compared with those of rainfall. The studied area is an important fluvial and environmental Europe during the end of MIS3 and MIS2. threshold between the Atlantic and Mediterranean basins. The demise of the Neanderthals in the north of the Iberian Peninsula took place between 40 and 30 ka (MIS3) during a 3. Material and methods climate characterised by abrupt changes with periods alternating between cold and warm together with the expansion of the We selected the glaciomorphological and palaeoclimate se- anatomically modern human (AMH) and changes in industrial and quences, and human occupation information from previously pub- cultural typologies. The Neanderthal was specialized in exploiting lished studies. The glacial sequence analyzed comprises: (i) glacial ecotonal ecosystems with a wide variety of resources in small reconstructions based on geomorphological maps (1/10.000 to 1/ areas; moreover, the AMH was able to exploit open landscapes in 25.000 scales) and morphostratrigraphic sequences (Gonzalez- cold and dry environments. The AMH survived and expanded in Trueba, 2007a, b; Pellitero, 2009, 2013, 2014; Gonzalez-Trueba and the Cantabrian Cordillera from 42 to 41 ka cal BP (Sepulchre et al., Serrano, 2010; Gonzalez Trueba et al., 2011; Serrano et al., 2012, 2007; Hoffecker, 2009). According to Finlayson the temperature 2013); (ii) Equilibrium line altitude estimation by the AAR method in patterns and climate variability of the last 85 ka determined eight massifs for each stage (Serrano et al., 2013a, b); (iii) Chronologies changes in habitat and distribution of hominids, cold stable pe- based on numerical dating either by 14C, AMS, U/Th and OSL (Table 1). riods were alternating with extinctions and warm periods with The different dating techniques imply chronological uncertainty about persistence (Finlayson, 2004, p.58). The main discussion on the the maximum ice extent and the LGM in the Iberian mountains. hypothesis regarding the migration and demise of the Neanderthal Although the dating procedures, AMS, 14C, and OSL, pose different focuses on whether it was triggered by the expansion and impo- limitations and problems, such as possible contamination of samples sition of AMH and its direct competition by 'competitive exclusion' of aquatic organic matter affected by old carbon, evolution of com- (Banks et al., 2008); by the assimilation by AMH, which shared plexes or fine deposits undetected during field work, and the results some of its genes, which remains a very premature hypothesis can be questionable, the coherence of the dates obtained by AMS and (Havarti, 2012); or by migration and extinction related to envi- OSL techniques increases the consistency of glacial chronologies; (iv) ronmental changes that occurred during MIS3 and the subsequent Sedimentological archives and a sequence of Loss of Ignition (LOI) in population dynamics of the species (Finlayson, 2004; Finlayson the Picos de Europa (Serrano et al., 2012) used as evidence of envi- and Carrion, 2007). ronmental changes directly related to glacial evolution; and (v) In- formation on environmental changes and human population in the 2. Regional setting eastern Cantabrian Mountains has been obtained from natural deposits and archaeological research (Gutierrez and Serrano,1996, 2007; The Cantabrian Mountains are located in the north of the Iberian Baena et al., 2004; Cabrera et al., 2004; Rasilla de la and Straus, 2004; Peninsula, (43N; 500Wto200W) stretching 700 km in an east-west Garralda, 2005; Straus, 2005; Iriarte et al., 2005; Sanchez Goni~ and

Fig. 1. Location of study area and distribution of studied glaciers by massifs. 1, the Picos de Europa. 2, Cebolleda. 3, Fuentes Carrionas. 4, Pena~ Sagra. 5, Alto Campoo and Valde- cebollas. 6, Pas. 7, Gorbeia. 8, Aralar. 24 E. Serrano et al. / Quaternary International 364 (2015) 22e34 d'Errico, 2005; Uzquiano, 2005; Sepulchre et al., 2007; Uzquiano et al., in precipitation from west to east in the Pleistocene lowered the 2008, 2012; Hoffecker, 2009; Alvarez-Lao and García, 2011; Maroto ELAs to less than 1300 m a.s.l. in the east. The western area is et al., 2012; Garcia Garriga et al., 2012). Several parameters such as characterised by massifs glaciated above 2000 m a.s.l. with ELAs open-air or cave sites, mountain or coastal location, and altitude have of between 1600 and 1800 m a.s.l. been considered in order to differentiate environmental conditions.

Table 1 Dating for the glacial related deposits in the Cantabrian Mountains.

Place Site Tecnique Sediment Depth (m) Age a BP Ages cal a BP Reference

Picos de Europa Aliva U/Th Debris slope e 396,470 ± 24,083 e Villa et al. (2013) Duje U7Th Debris slope e 192,700 þ 3190/e2330 e Castan~on and Frochoso (1996) Aliva AMS Lacustrine 1 6660 ± 40 6720e6640 Serrano et al. (2012) Aliva AMS Lacustrine 1.33 8310 ± 50 9400e9300 Aliva AMS Lacustrine 3 17,300 ± 100 17,400e17,300 Aliva AMS Lacustrine 5.50 21,390 ± 100 21,500e21,390 Aliva AMS Lacustrine 7.80 27,570 ± 320 32,580e31,900 Aliva AMS Lacustrine 10.55 26,090 ± 240 30,660e31,400 Aliva AMS Lacustrine 14.10 27,460 ± 300 31,850e32,426 Aliva AMS Lacustrine 15.50 31,200 ± 440 35,700e34,850 Comeya AMS lacustrine 35.5 40,480 ± 820 44,118 ± 885 Jimenez and Farias (2002) Enol AMS Lacustrine 6 m e 9109 ± 74 Jimenez et al. (2013) AMS Lacustrine 10 e 14,734 ± 326 OSL Cemented sand and silt 42.7 44,966 ± 3337 e Pas Mountain Trueba valley AMS Peat 1 26,082 ± 118 29,149e28,572 Serrano et al. (2013a, b) AMS Peat 0.50 8768 ± 42 7968e7633 AMS Peat 1.40 8186 ± 35 7310e7077 AMS Peat 1.60 10,467 ± 42 10,623e10,420 Soba valley OSL Till e 44,98 ± 2.3 e Frochoso et al. (2012). OSL Till e 41,56 ± 2.4 e OSL Till e 44,53 ± 2.45 e Altos Ason OSL Till e 40,43 ± 5.14 e Zucia OSL Till e 13,42 ± 1.25 e Fuentes Carrionas Vega de Naranco OSL Proglacial/Supraglacial 0.50 36,028 ± 2350 e Pellitero (2011) AMS Lacustrine 5.50 15,614 ± 21 16,950e16,695 AMS Lacustrine (sands) 3 25,591 ± 23 28673e28300 AMS Lacustrine 0.33 6831 ± 50 5750e5660 AMS Lacustrine 19 14,275 ± 49 15,570e15,240 Sanabria Pías OSL Lacustrine 27,000 ± 200 e Perez-Alberti et al. (2011) OSL Lacustrine 3100 ± 300 e OSL Lacustrine 33,000 ± 300 e Sanabria lake AMS Basal clastic unit 25,584 ± 374 Rodríguez et al. (2011) Top of clastic unit 14,494 ± 347 Clastic unit 13,100 Coastic unit 12,300 San Martin AMS Lacustrine 21,833 ± 358 Redes Natural Park Monasterio C14 Lacustrine clastic 35.6e35.5 28,990 ± 230 33,485 ± 362 Jimenez et al. (2013) Tarna Valley OSL Till 23,967 ± 1841 e OSL Till 23,967 ± 1841 e OSL Landslide 22,983 ± 2321 e OSL Lacustrine 42 44,966 ± 3337 e AMS Peat bog 20.640 ± 300 24,579 ± 421 Ancares Radiocarbon dates Peat bog 17,400 Munoz-Sobrino~ et al. (2001) Sil Valley Villaseca AMS Glaciolacustrine e 41,150 Jalut et al. (2010)

Table 2 4. Pleistocene glaciations and chronology Glaciated massifs in the eastern Cantabrian Mountains. Massif Altitude Glaciers N Ice surface Orientations % m a.s.l. In the Cantabrian Mountains, Pleistocene glaciations were Km2 %N S limited to the highest massifs and all glaciers were of small size Picos de Europa Urrielles 2648 24 68.55 20 55 45 (Table 2) compared with those of the Pyrenees or the Alps. Andara 2444 15 30.5 9 57 43 Nevertheless, the Cantabrian Mountains hosted the lowest gla- Cebolleda 2078 11 16.75 5 77 23 ciarized massif in the Iberian Peninsula. Two groups of glacial Fuentes Carrionas 2536 20 105 30.7 45 55 ~ massifcanbedifferentiatedintheeasternCantabrianMountains Pena Sagra 2046 15 11.4 3 87 13 Campoo 2125 10 20 6 15 85 in terms of altitude, longitude and distance from the sea (Fig. 2). Valdecebollas 2143 4 5.8 2 25 75 In the easternmost area, glaciers developed in very low moun- Pas 1718 8 76.5 22.5 67 33 tains, on the Pas Mountain, Gorbeia and Aralar, where cirque, Gorbeia 1481 3 1.15 0.3 100 e alpine and ice domes were emplaced below summits of 1700 Aralar 1413 2 5.35 1.5 100 e e and 1300 m a.s.l. A strong longitudinal gradient and an increase Total 112 341 100 67 33 E. Serrano et al. / Quaternary International 364 (2015) 22e34 25

Fig. 2. Distribution of main glacial parameters of glaciated massifs.

Recent studies with a regional focus centring on Cantabrian a first approximation to the Late Pleistocene glacial chronology. Mountain massifs have provided optically stimulated luminescence During this period, three main Pleistocene phases have been (OSL) and radiocarbon dates for the glaciolacustrine, proglacial, detected (Jimenez et al., 2013; Serrano et al., 2013a,b; Rodríguez- peat bog deposits and moraines. Dates obtained by radiocarbon Rodríguez et al., 2014). (14C, AMS) of discrete organic matter fragments or concentrated pollen to minimize a possible hard-water effect (Jalut et al., 2010; Table 3 Moreno et al., 2010; Serrano et al., 2011, 2012) have been ob- Glacial stages and correlation between massifs in the eastern Cantabrian Mountains. tained in Sil Valley, Picos de Europa and Pas Mountains, and are consistent with U/Th and OSL dates (Pellitero, 2011; Serrano et al., Massifs Pleistocene glacial stages 2012; Frochoso et al., 2013) from the Palentina and Pas Mountains. S-I S-II S-III In the Picos de Europa, a slope deposit on an erosional feature on Picos de Europa, Urrielles I II III limestone has been dated as prior to MIS7 (Frochoso and Castan~on, Picos de Europa, Andara I II IIIa 1996) and 396.47 ± 24.8 ka (MIS11), inferring an age prior to 478 ka IIIb Cebolleda I II IIIa (MIS12 glacial period) for the first glacial features (Villa et al., 2013). IIIb This is an indirect date, and an unclear glacial feature below the Fuentes Carrionas I II III slope deposit requires more data to confirm an ancient glaciation Ib prior to the slope deposits. Ancient glacial deposits have been Campoo I II III described (Serrano, 1991) and dated in the Pyrenees (Pena~ et al., Valdecebollas I II IIIa IIIb 2004; Lewis et al., 2009; Sancho et al., 2008, 2011; Delmas et al., Pas Mountains Ia IIa-IIb III 2011) and have provided the most extensive glaciations with an IV age during MIS6 (190e130 ka). Dates have also been obtained in the Gorbeia I II e NW Iberian Peninsula of around MIS8 and MIS6 (Fernandez- Aralar I II e Mosquera et al., 2000). Pre-Würm glaciations in the Cantabrian Age MIS3 MIS2 MIS2 and Pyrenean Mountains, located between MIS12 and MIS6, have highly uncertain chronologies depending on the different dating 4.1. Stage of maximum expansion (S-I) techniques used, and morphostratigraphic sequences and more deposits, landforms or dates are required. Glaciers occupied the maximum extent during S-I, when the The morphostratigraphic correlation of the glacial landforms ELAs were located at an average altitude between 1580 and 1700 in (Table 3) and dating of different moraine complexes have facilitated the western massif, and 1190 in the Castro Valnera icefield (Fig. 2). 26 E. Serrano et al. / Quaternary International 364 (2015) 22e34

During S-I in Campoo, Valdecebollas, Fuentes Carrionas and deposits (lacustrine, peat bog and till) and using different tech- Cebolleda Cebolleda massifs, alpine and cirque glaciers were niques (AMS and OSL) confirm the maximum extent of glaciers developed, in which the mean altitudes of the ELAs increased with during MIS3, before the LGM of N and NW Europe. distance from the sea. Cebolleda and Fuentes Carrionas massif have If all massifs have homogeneous thermal behaviour, the differ- a sharp NeS dissymmetry and the ELAs are very low to the north. ences between massifs are determined by changes in snowfall Alpine glaciers have large cirques and tongues (between 3.7 and conditions (Serrano et al., 2013a,b). Therefore, snowfall intensity 14.9 km length) reaching 1200 m a.s.l., where the moraine com- and origin (NW, N and SW) would be the determining factor in the plexes are well preserved. On the southern side, only cirque glaciers types of glaciations during Pleistocene glacial stages. Limitations without tongues were generated. Alpine glaciers developed in Alto deriving from distance from the sea, barrier effects, and altitude are Campoo and Valdecebollas, whose fronts were situated at around compensated by SW flows feeding snowfall towards the southern 1300 m with the ELA at 1700 m a.s.l. In Pena~ Sagra, a massif located massifs. This different behaviour could only be explained by a more only 26 km from the sea, alpine glaciers oriented to the north (87%) southerly circulation of the atmospheric currents, with SW fronts were dominant. Similar processes took place in Gorbeia, where reaching the southern massifs (Florineth and Schlüchter, 2000). only three small cirques oriented to the north and generated by leeward spindrift accumulation were shaped by ice (Fig. 3B). 4.2. Stage of alpine glaciers (S.II) In the northernmost massifs, with greater Atlantic influence (Pas, the Picos de Europa and Aralar), lesser distance from the sea This stage was characterised by a small reduction in the extent and no physiographic barriers, ice domes developed. Glaciers of the of glaciers and a moderate ascent of the ELA, between 1200 and Picos de Europa were short, but their fronts reached very low al- 1775 m in the western mountains and 1200 m in the Castro Valnera titudes, 450 and 600 m a.s.l. in the north side and 800 m in the icefield (Fig. 2). The glaciers were of similar proportions in extent to south (Figs. 2 and 3). The maximum expansion of glaciers was the previous phase but with an increase in volume in the frontal defined by summit icefields with outlets reaching 7 km in the Duje, portions. 4 km in the Deva and 4.2 km in Valdiezmo. The Pas Mountain The glacial retreat and re-advance to positions very close to the developed two glaciated sectors, the Castro Valnera massif (1718), previous phase have been revealed by changing directions of glacial where a icefield developed with an outlet dropping to the north flows in the Picos de Europa, two different infill deposits in the (Miera tongue of 5 km length) and the south (Trueba tongue, 15 km Trueba Valley, and the different lithology of the moraines at Vega length) and the Veinte-Pizarras massif (1472 m), at the north of the del Naranco in Fuentes Carrionas (Turú et al., 2007a; Pellitero, 2011; watershed, where cirques and troughs of moderate size developed. Serrano et al., 2012, 2013). A period of glacial equilibrium has been The icefield shows net dissymmetry with the main ice accumula- dated by minimum LOI in the lacustrine sediments of Aliva at be- tions to the east of the dividing line, low altitudes of glacial fronts tween 21,500e21,390 cal a BP, a second stage of deglaciation in Enol (400e600 m a.s.l. to the north and 760 m a.s.l. in the south side) and between 20 and 18 ka (Moreno et al., 2010) and 20,640 ± 300 a BP the ELA at 1190 m a.s.l. (Fig. 2). It was a large glacier as low as in the Alto Nalon (Jimenez et al., 2002). A glacial advance has been 1700 m and reaching front altitudes analogous to the western established between 32 and 18 ka (Jalut et al., 2010) in the Pyrenees massifs or Pyrenean glaciers. The Aralar Range (Ugarte, 1992; Rico, and the Cantabrian Mountains. It could belong to MIS2 cold stage, 2012) was occupied by a small ice dome around 3.9 km2 with or to the transition from MIS3 to MIS2 (H2), possibly prior to the several tongues reaching the 850 m a.s.l. The low altitude and LGM of N and NW Europe or very close to it. The explanations of eastern longitude of glacial processes does not provide an expla- this cooling and glacial stage may be the altitudinal decrease of the nation of the existence of colder conditions than in the western- frozen sea that led to the prevalence of cold but rather dry air most Pyrenean massifs. Distance from the sea, physiography and masses, generating intense cold conditions, dry winters, absence of the longitudinal gradient of snowfall from west to east explain the air masses from the SW and a relative decline in the snowfall from development of glaciers at unusually low altitudes in the context of the N and NW (Serrano et al., 2013a,b). a hyperhumid climate characterised by strong winter snowfall and a cold summer. 4.3. High mountain advance, stage of cirque glaciers (S-III) The threshold of the lowest glaciated peak was at 1800 m a.s.l. in the Picos de Europa and Campoo, and around 1400 m a.s.l. in the Northern oriented glacial cirques over 1800e2000 m, especially Pas Mountains. Campoo, Cebolleda, Pena~ Sagra, Valdecebollas, those protected by walls 200e250 m high in shaded and high sites, Gorbeia and Aralar denoted a topoclimate influence in the location present well conserved moraine complexes that reveal a cirque of cirque and alpine glaciers, dominating in the northern ones. In glacier stage. These landforms belong to a period of equilibrium and the Valdecebollas and Fuentes Carrionas massifs, located to the minor advance with ELAs at 2050e2130 m a.s.l. Ice fields and ice south of the watershed, due to their windward position and mod- domes disappeared during this moderate cooling period, no gla- erate altitude only a SW flowing atmospheric circulation can ciers remained in Gorbeia and Aralar, and only small alpine and explain the oceanic character of the glaciated massif, confirmed by cirque glaciers were left on the northern slope of the Pas Moun- a pattern of lower ELAs in the north and northeast glaciers. tains. Topoclimatic factors determined the development of glaciers. Results of dating (Table 1) in the Picos de Europa place the The cold permitted the genesis of small rock glaciers but the glacial maximum prior to 40 ka in Enol (Moreno et al., 2010); snowfall was not enough for the development of large glaciers. 40,480 ± 820 BP in Comeya (Jimenez and Farias, 2002); and A high degree of uncertainty remains regarding the chronology 35,700e34,850 cal BP in el Duje (Serrano et al., 2012). In the of this period because it is based on indirect data. An equilibrium in Fuentes Carrionas the minimum age is of 36,028 ± 2350 a BP for the the Ca content between 14.5 and 13.5 ka in Enol (Moreno et al., glacial maximum (Pellitero, 2011; Serrano et al., 2012). The Trueba 2010) and a moderate minimum LOI in the lacustrine sediments Valley (Pas Mountains) yields an age prior to 29,149e28,572 cal a of Aliva circa 13.9 ka (Serrano et al., 2012) are interpreted as cooling. BP (Serrano et al., 2012, 2013) and on the north side an age of 44,980 ± 2.3 ka (Frochoso et al., 2012). Further west, dates point to 5. Human occupation ages of 28,990 ± 230 a BP in Redes (Jimenez et al., 2002, 2013), around 44 ka in the Sil Valley (Jalut et al., 2010) and prior to 25.6 ka There is discussion between authors who think of a local tech- in Sanabria (Rodríguez et al., 2011). All dates, made on different nological and cultural transition between Middle and Upper E. Serrano et al. / Quaternary International 364 (2015) 22e34 27

Fig. 3. Glacial extent and evolution in the larger (A, The Picos de Europa) and smaller (B, Gorbeia) glaciated massifs.

Palaeolithic techno-complexes, with Aurignacian emerging as a due to problems related to dating, cultural attribution, or a lack of local evolution from the Mousterian (Cabrera and Bernaldo de reliable radiocarbon dates, and the supposed survival of Neander- Quiros, 1990;Saenz de Buruaga, 1991, 2004; Cabrera et al., 2001) thals in other European regions is not supported by recent radio- and authors who think that there is no reliable evidence suggesting carbon dating (Maroto et al., 2012). this continuity (Maroto et al., 2012). The coexistence of Neander- Neanderthal groups are documented in the Iberian Peninsula thals and AMH in Iberian Peninsula sites can currently be dismissed from ~170 ka to ~30 ka, which reﬂects inter- and intra-population 28 E. Serrano et al. / Quaternary International 364 (2015) 22e34 variability, and although most Neanderthal remains were found Aurignacian was in early development (Garcia Garriga et al., 2012). with Mousterian industries, a few appeared with the so-called But doubts remain about the exact chronology of fossils, dated 'transitional cultures' between Middle and Upper Palaeolithic between 45.7 ka and 38.5 ka, and Mousterian cultures. The archaic (Garralda, 2005). The main Neanderthal ﬁndings of the Iberian Aurignacian of El Castillo Cave would be, at ~40 ka, among the most Peninsula very often have chronostratigraphical problems, not only recent in Europe, but the context of Level 18 (see Table 4) is ques- those found a long time ago but also remains recently discovered. tioned by some authors. They consider a mixture of two techno- Only in six sites of the Cantabrian Mountains and the coast have logically differentiated assemblages, with Aurignacian occupation remains been found of Neanderthals (Table 4), all dated between in the upper part and a Mousterian presence in the lower part of the 45.7 and 38.5 ka (Garralda, 2005). sequence (Garcia Garriga et al., 2012). The most accurate dating of a

Table 4 Neanderthal remains located in the North of Spain (Garralda, 2005).

Site Province Dating Material Culture MIS Reference

Lezetxiki Gipuzcoa e Bones Pre Mousterian MIS4 Baldeon (1993) Teeth MIS3 Arrillor Cave Alava 45,700 þ 1200 a BP Teeth Mousterian MIS3 Bermúdez de Castro and 45,400 þ 1800 a BP Teeth Mousterian MIS3 Saenz de Buruaga (1999) Axlor Shelter Biscaye 42,010 þ 1280 a BP Bones Mousterian MIS3 Basabe (1973) Teeth Gonzalez-Urquijo et al. (2006) El Castillo Cave Cantabria e Bones Mousterian MIS3 Cabrera et al. (2004) Teeth Maíllo et al. (2004) 40 ka Bones Transitional MIS3 38.5 ka Teeth Aurignaciena Sidron Cave Asturias 43,129 þ 129 cal a BP Bones Mousterian MIS3 Lalueza et al. (2005) Teeth

a Level 18 El Castillo cave, in Cabrera et al. (2004).

Human occupation of the Cantabrian region in the study period Neanderthal fossil by AMS in the Sidron Cave, 43,129 ± 129 cal a BP was characterised by important changes in population density (Garralda, 2005; Lalueza et al., 2005) is included in the glacial (Finlayson and Giles, 2000; Finlayson, 2004; Straus, 2005). The period S-I. Finally, the geographical distribution of sites shows the maps of sites (Baena et al., 2004; Cabrera et al., 2004; Rasilla de la latest Neanderthals inhabiting the harshest, but also the most and Straus, 2004; Straus, 2005; Gutierrez and Serrano, 2007) reveal diverse environments, and not the temperate conditions of the high Mousterian density and a greater proliferation of sites in southern areas (Maroto et al., 2012). mountain regions (Garcia Garriga et al., 2012; Maroto et al., 2012). d'Errico and Sanchez Goni~ (2003) and Finlayson et al. (2008) In this sense, the Mousterian occupation reached the high valleys, consider climatic deterioration during H4 to be the main cause of with sites in Nestares or Mave in Campoo (Gutierrez and Serrano, the contraction and disappearance of the Neanderthals, together 2007, Díez-Martín et al., 2011; Yravedra et al., 2013), a natural with MIS2 climatic deterioration and the development of open connection between the Cantabrian coast and the Duero, and the landscapes, perhaps more suitable for AMH, and competing with inner mountain at low altitudes in Picos de Europa or Pas Valley. each other (d'Errico and Sanchez Goni,~ 2003; Banks et al., 2008). The moderate density of Neanderthals in the Cantabrian region However, conditions were probably not extreme and other authors 39 ka ago (Finlayson, 2004) coincides with the combination of sustain that with great diversity in geography and ecosystems be- Mousterian and Gravettian cultures and the change in conditions tween 40 and 30 ka in MIS3, the key historical event was the from forest landscapes with closed woods of dominant Mousterian expansion of AMH and Upper Palaeolithic technology (Garcia culture to open Gravettian-dominant ones (Finlayson, 2004) and Garriga et al., 2012), not climatic variability. Therefore, some ar- also a moderate presence of AMH. chaeologists are in agreement on one explanation of the extinction In the northern Iberian Peninsula, Maroto et al. (2012) described or assimilation of the Neanderthals in the northern Iberian Penin- the first Upper Palaeolithic assemblages appearing mainly in low- sula, based on the relations between neighbouring communities land areas linked to natural corridors and thick Middle Palaeolithic and the alteration of the equilibrium (Garcia Garriga et al., 2012). sequences where Middle Palaeolithic populations were replaced by They argue that if Neanderthal withdrawal was the response to Upper Palaeolithic ones. This is probably the case of the El Castillo climate change, flows of cultural groups over time should be and Lezetxiki fossils, all of them very near the passages to or from detected. Archaeologists have observed that when Neanderthals or southwestern European regions (Garralda, 2005). After 40 ka, these Middle Palaeolithic levels disappeared from a particular region, it assemblages tend to appear at higher altitudes, in mountain land- was forever. Garcia Garriga et al. (2012) point to an agreement with scapes with clearly diverse ecological conditions characterised by a model of competition among groups rather than climate environments with contrasting features in terms of their biological adaptation. potential to support humans, with long occupational hiatus, where In the Aurignacian, a fall in population density took place in there was no cultural replacement, and the extinction of the Ne- contrast with the stability of the Acheulean and Mousterian, as well anderthals cannot be explained by the pressure of Upper Palae- as a fall in the number of sites and a distancing from the mountain, olithic populations (Garcia Garriga et al., 2012; Maroto et al., 2012). though this also occurred on the coast (Rasilla and Straus, 2004). It Although most Neanderthal finds are related to Mousterian cul- was not only a fall in resources and migration towards lower areas, tures, in El Castillo and Lezetxiki there are documented remains of but a generalized depopulation in which the possible emigration to 'Transitional Aurignacian' or 'Transitional Mousterian' techno- the north of the AMH would not be substituted by the Neanderthal. complexes (Garralda, 2005; Garcia Garriga et al., 2012). Two key Around 33 ka, the Neanderthals disappeared from the Cantabrian sequences, El Castillo and El Esquilleu, show the survival of Middle region, coinciding with a high density of AMH in the north of the Palaeolithic communities in inland areas, while in coastal areas the Iberian Peninsula. E. Serrano et al. / Quaternary International 364 (2015) 22e34 29

The mapping of the Gravettian shows a moderate increase in dependant on changes between precipitations and wetness. sites, mainly at low altitudes on the Cantabrian slope near the high Changes between glacial stages SI and SII also derive from changes mountain, with the re-occupation of inland valleys. In the Solutrean in moisture linked to variations in the atmospheric dynamic, there was a sharp population increase attributable to the ‘refuge mainly of the latitude of NW flows. Between 65 ka and the LGM, effect’ (Straus, 2000) deriving from southward migrations resulting short cold water stages (Heinrich events) have been reported, one from cooling and glacial expansion in the north. This phase co- of which between 39 and 40 ka was of similar intensity to that of incides with the complete extinction of the Neanderthals to the the LGM. south and a sharp fall in population in north and northwest Europe. The regional responses to climate variability have been pointed In the Magdalenian, density increased in the north of Europe and to as causes of different glacial periods in southern Europe (Hughes north of Iberia, mountain areas becoming occupied at the end of the et al., 2006; Delmas et al., 2008; Serrano et al., 2012; García-Ruíz LGM. et al., 2013). Small glaciers characterising the Cantabrian Moun- The premises adopted by Maroto et al. (2012), first the consid- tains react more rapidly than large glaciers, with almost immediate eration that the Middle Palaeolithic techno-complexes were pro- expansion or contraction of the ice mass as a consequence of duced by Neanderthals and second, that the Upper Palaeolithic was changes in precipitation or temperature. In the Pyrenees (García- the product of AMH, have been adopted by us to consider the Ruíz et al., 2013) it has been pointed out that if, during some of disappearance of the Neanderthals. Both premises consider that the abrupt changes (Dansgaard et al., 1993) the Scandinavian ice MIS3 is characterised by wide climatic variability with a highly cap and valley glaciers had undergone sustained growth during changing environment in which there were glaciers at the summits, MIS3 due to the greater inertia, with stabilization or slower growth forcing humans to readjust their behaviour and develop new of the ice mass, other small Mediterranean glaciers would have adaptive strategies to occupy and exploit the territory. retreated during the warm periods because of their greater sensi- tivity to climate fluctuations. In the Cantabrian Mountains, the 6. Discussion hyperhumid climate and the reception of SW cold and wet air masses contribute to the regional responses of glaciers depending Climatic and glacial evolution, geographical location and phys- on cooling or moisture increase (Serrano et al., 2012, 2013). The iography in the eastern Cantabrian Mountains have given rise to a location of the Polar Front during the MIS4 and 3, at approximately wide variety of ecosystems and landscapes over the last 70 ka. The 46N, favoured meridian circulation (Florineth and Schlüchter, differing environments arose because of the oceanic climate and 2000), which explains the different response to the Polar Front. existence of Pleistocene glaciers during S-I, S-II and S-III. Glaciers Variations in the growth of glaciers have been recognized not only were relegated to the interior of the mountains, at low altitudes in in the Cantabrian Mountains, but also in the Pyrenees, the Vosges, the north (400e600 m) where they shared territories with humans, and the northern and western Alps (Seret et al., 1990; Preusser and higher ones in the south (700e1000 m). et al., 2007; Ivy-Ochs et al., 2008). Changes in the Polar Front and The S-I glaciation is dated at 42e31 ka ago, a period coinciding the dry conditions after 30 ka were a limiting factor to glacial with MIS3 with the presence of dated Neanderthal fossils expansion in humid or Mediterranean environments of the (40e38 ka) and Mousterian cultures in the north of the Iberian southernmost latitudes of Europe and glaciers did not reach the Peninsula. Glacial chronological results are similar to the Pyrenees, positions they did during MIS3. where dates indicate that the glacial maximum occurred during In the eastern Cantabrian Mountains, the more intense snowfall MIS4 and MIS3, earlier than in central and northern Europe (MIS2) supplied by cold and partially humid air masses from the N and (Pena~ et al., 2004; Sancho et al., 2008, 2011; Lewis et al., 2009; NW, as well as less cold but more humid ones from the SW, may García-Ruíz et al., 2010, 2013). The LGM was restricted to the have been the cause of a more extensive S-I glacial advance, be- headwaters (García-Ruíz et al., 2003) and although evidence of the tween 42 and 30 ka ago, than the LGM. The MIS3 period has been LGM is scarce (García-Ruíz et al., 2003, 2010; Lewis et al., 2009), the characterised in the Esquilleu Cave by open landscapes with landforms related to LGM valleys occur several kilometres up- arboreal taxa (Pinus, Betula, Juniperus) discontinuously distributed. stream of the maximum ice extent features (Garcia Garriga et al., The morphological features and dominant humid conditions fav- 2012). Studies in the Mediterranean mountains based on 14C and oured the survival of mesophilous and hygrophilous taxa OSL dating techniques led to similar conclusions (Mardones and (Uzquiano et al., 2008) and provided enough vegetable biomass Jalut, 1983; Andrieu et al., 1988; Jalut et al., 1992, 2010; Hughes and housed related fauna to support periodic seasonal human oc- et al., 2006; Hughes and Woodward, 2008) and cosmogenic sur- cupations (Uzquiano et al., 2012). A mild and moist period has been face exposure ages have provided ages coherent with an early detected in the Atlantic during 38.6e36.5 cal ka BP (GI8) with the maximum ice extent in the Pyrenees (Pallas et al., 2010; Delmas development of a weak deciduous forest (Banks et al., 2008), and a et al., 2011, 2012). All of this leads to the conclusion that a favourable moisture period has been detected in the Esquilleu Cave maximum ice extent occurred in the Pyrenees much earlier than (Phase III, layers XIII-V) by a decrease in Pinus and an increase in the global LGM. Nevertheless, not all dates show a maximum mesophilous taxa between 39 and 34.3 ka BP (Uzquiano et al., advance in MIS3. Studies based on 10Be exposure in the Pyrenees 2012), a fact which may be more pronounced due to the regional indicate an ice advance maximum in the LGM (MIS 2). All cases are climate between the sea and the mountains in the easternmost obtained in the eastern Pyrenees, but the debate regarding the Cantabrian Mountains. The vegetation in the Cantabrian area dur- chronology of the maximum ice extent in the Pyrenees remains ing MIS3 is an open forest on poorly developed soils with Pinus on open (Pallas et al., 2006, 2010; Delmas et al., 2008; García-Ruíz dry subtracts, Betula with Sorbus on humid ones, and occasionally et al., 2010; Calvet et al., 2011). hygrophilous taxa such as Salix and Alnus, with Corylus, Quercus, The presence of glaciers during MIS3 implied geographical and Castanea (Iriarte et al., 2005; Sanchez Goni~ and d'Errico, 2005; variations (latitudinal and altitudinal) on the distribution of gla- Uzquiano, 2005; Uzquiano et al., 2008, 2012). Between 39 and 36 ka ciers and environments. The important latitudinal changes and there was a pre-forest vegetation cover characterised by colonizing glaciers at lower altitudes in the easternmost area would be re- species such as Pinus, Sorbus and Betula (Uzquiano, 2005). flected in geographical variations in vegetation landscapes. There- During the S-I, prior to 38.5 ka, Neanderthals were living in the fore, the altitudinal differences between the glaciation threshold Cantabrian Mountains following an isolation period or migration to altitude, ELAs and front altitudes can be considered as being the south, but they disappeared from the Cantabrian area 30 E. Serrano et al. / Quaternary International 364 (2015) 22e34 coinciding with environmental changes (Fig. 4) from closed to open species that remained in the north of the Iberian Peninsula until landscapes (Uzquiano, 2005; Sanchez Goni~ and d'Errico, 2005; 10 ka (Alvarez-Lao and García, 2011). However, to the north the Iriarte et al., 2005). Corridors passing toward the Duero basin had wide variety of ecosystems and the location of the glaciers, at the been used during previous periods (Gutierrez and Serrano, 2007; top of mountains with a natural passage free of ice, would not Díez-Martín et al., 2008). AMH reached the area during a wet and necessarily appear to be a limiting factor. cold period using the Cantabrian coast and mountains as a refuge The only new and very different event during the MIS3 was prior to 37 ka BP. The early arrival of the Aurignacians has been AMH taking refuge in the northern Iberian Peninsula (Fig. 4). The interpreted as a relatively rapid substitution of the Neanderthals by hypothesis of ‘competitive exclusion’ (Banks et al., 2008) seems AMH in the north of the Iberian Peninsula (Garcia Garriga et al., possible, though other authors do not consider the evidence to be 2012; Maroto et al., 2012), but the co-existence of AMH and Ne- compelling (Stewart, 2005). Relationships between often obscure anderthals in the same region may span from the initial S-I until the environmental and cultural changes (Straus, 2005) are, therefore, warming period at the end of the S-I. During the S-I, climatic one possible explanation for the demise of the Neanderthal popu- instability, fragmentation of the remnant Neanderthal population lation. On a regional scale we can consider the inability to adapt and the existence of environment non-analogues with modern culturally to the new environment and abrupt conditions (climate ones (Finlayson, 2004; Stewart, 2005) may have generated intense variability, AMH population, faunal migration and extinctions), environmental stress on Neanderthal populations, leading to overcome in the recent past but now collapsed. southward migrations. If the Neanderthals were able to remain in The glaciers retreated and partially abandoned the mountain the Iberian Peninsula by adapting to environmental changes such valleys until the new glacial stage S-II, a glacial advance at the as the steppe environment between 200 and 100 ka as described by transition to MIS2. The cooling of the LGM coincided with a period some authors, the cold events in MIS4 earlier than 70 ka, H6 and of intense cold and glacial advance in all massifs studied. The MIS3 (H5), some of which were abrupt changes between warm intense cold was accompanied by a decrease in altitude of the phases with swift environmental changes, it is difficult to reach the frozen sea that led to the prevalence of cold but very dry air masses conclusion that they could not adapt to cold and wet events (H4, from N and NW during the winter and a relative decline in snowfall H3). It has been pointed out that changes in the ecosystems the and air masses from the SW (Ruddimann and McYntire, 1981; Neanderthals belonged to would generate an absence of resources Florineth and Schlüchter, 2000). During that period, the glaciers culminating in their demise, partially coinciding with the extinc- were less extensive but thicker, reflecting a more stable cold period tion of associated fauna (Finlayson, 2004; Stewart, 2005; Finlayson characterised by cold and dry conditions, with stepped landscape et al., 2008). Nevertheless, we have seen how orographic charac- reaching the lower part of mountain valleys. The lowest and most teristics and the longitudinal gradient cause a wide variety of continental massifs evolved towards marginal glaciation immedi- ecotones between the steppe and forest environments in a cold and ately after the glacial maximum. Glacial advance is coetaneous to wet Atlantic climate. Neanderthals failed to adapt to the new an open landscape occupied by AMH and cold-adapted mammal environmental conditions, although they had, no doubt, been species, which sheltered in the Cantabrian area until 17.5 cal ka BP considerably more flexible in the past (Straus, 2005). Conditions to (Alvarez-Lao and García, 2011). the south of the Cantabrian Mountains were possibly less rich, The last Pleistocene glacial stage (S-III) extended only over the made up of extensive steppes occupied by cold-adapted mammal highest ranges, whereas the easternmost massifs and low altitude

Fig. 4. Cultural and climate events in the Cantabrian region (Synthesized from Finlayson and Giles (2000); Finlayson (2004); Straus (2005); Sepulchre et al. (2007); Serrano et al. (2012, 2013)). E. Serrano et al. / Quaternary International 364 (2015) 22e34 31 ranges were free of ice during this phase. The presence of cirque reconstruction of glacial advances and retreats and climatic varia- glaciers and rock glaciers appears to be a general feature in the tions are not in accordance with responses in Palaeolithic popula- Cantabrian Mountains. Glaciers developed in Campoo, Fuentes tion density. Glacial stages coincide with population increases at Carrionas, Cebolleda and the Picos de Europa. The cooling meant the end of the Mousterian culture, in the Gravettian and Solutrean, the landscape was dominated by herbaceous vegetation but with and in the Magdalenian. little representation of cryoxeric taxons. The oceanic environment Glacial advances and equilibriums behave differently with re- remained and open landscapes with little forestation were domi- gard to northern Europe: S-I took place during MIS3 in a cold and nant (Iriarte et al., 2005). During the Late Glacial in the north of the wet environment associated with snowfall coming from cold, hu- Iberian Peninsula the ‘Magdalenian Boom’ (Straus et al., 2002) took mid masses from the N, NW and SW, and open landscapes in inner place, with an increase of human occupation favoured by the and coastal sites. The accentuated gradient of longitudinal pre- moderately wet and cold climatic conditions. Glaciers, located high cipitations meant that in the easternmost mountains glaciers were in the mountains, would no longer constrain human activities. generated at very low altitudes in favour of snowfall. Available The environmental changes in N and NW Europe explain the dates situate S-I at 42e31 ka, coinciding with the permanence and increase in population during cold phases in the Cantabrian expansion of Mousterian and Aurignacian cultures. Adaptation to Mountains, as AMH flowed south as a relatively cold and dry new environmental conditions could not be very different to what optimal refuge. This did not happen to the Neanderthals, who occurred during the previous phases of MIS3 and MIS4 (H5 and H6 disappeared from the Cantabrian region at the time of the S-I events), when Neanderthals still occupied the territories, mainly on glacial advance (Fig. 5), a cold, wet period at the end of the MIS3. the coast. If, during previous periods (MIS4, MIS5) they had adapted to Later, during S-II, characterised by shorter, thicker glaciers environmental changes, including abrupt cold events (H6, H5) denoting a cold, dry and more stable climate with glaciers fed by with important environmental consequences, the inability to snowfall from cold, dry air masses from the N and NW, the Nean- adapt to the changes that took place in H4 in the Cantabrian derthal and Middle Palaeolithic cultures withdrew. The climate was Mountain, with the intensification of cold but, above all, the colder and drier than S-I, but glaciers occupied only the high increase in humidity, it may be due to two factors, the compet- mountains. There were no environmental conditions different to S- itive exclusion imposed by AMH, who migrated to the south in I that would cause further difficulty to the Neanderthal occupation. search of better conditions, or to the cultural limitations of a S-II is coetaneous with the European LGM and AMH travelled from human group incapable of adapting with innovative responses to north and northwest Europe to the south, increasing the population the new environmental and cultural conditions, and to the on the Cantabrian coast. S-III was a brief period attributed to the presence of AMH. The weight of one of these hypotheses, pri- LGM, which is still poorly dated but related to the Younger Dryas marily cultural, seems more likely in view of the environmental when AMH population once more expanded throughout the north changes, glacial evolution and the continuity of oceanic climates. of the Iberian Peninsula. The cold period in northern Europe may More environmental and cultural records are needed to be more possibly have led to human flow to the south in search of favourable accurate in the chronologies and possible human responses to ecosystems in the wetter and warmer coasts and mountains cultural or environmental changes in northern Iberian Peninsula. compared with central and northern Europe. In this period the Following the territorial occupation by AMH during the cold and glaciers, small in size, were housed at altitude and disappeared dry S-II, the AMH population increased in response to the from the lower mountains. advance of the glaciers in northern Europe and migrations The reasons behind this lack of harmony between environ- thereafter. Finally, the last glacial advance, S-III, had no regional mental cooling and population may have come in response to influence and coincided with the population and cultural environmental changes in N and NW Europe, since on the Canta- expansion of the Magdalenian. brian coast the climate would have been relatively harsh with respect to that of the north. It would, thus, constitute an optimal 7. Conclusion refuge for AMH in spite of its relative cold or dryness. This did not happen to the Neanderthals, who disappeared from the Cantabrian In the Late Pleistocene in the Cantabrian Mountains during MIS3 region at the time of the S-I glacial advance (Fig. 5), a cold, wet and MIS2, three glacial stages have been established. The period at the end of the MIS3.

Fig. 5. Glacial, landscape and human evolution in the study area. (1), Moreno et al. (2010). (2), Serrano et al. (2012, 2013). (3), Uzquiano (2005), Uzquiano et al. (2008, 2012). (4), Finlayson (2004); Garralda (2005). (5) Hoffecker (2009). (6) Chronology of dated Neanderthal remains in Cantabrian Region. 1 and 2, El Castillo Cave. 3, Axlor Shelter. 4, El Sidron Cave. 5, Arrillor Cave. 32 E. Serrano et al. / Quaternary International 364 (2015) 22e34

If, during previous periods (MIS4, MIS5) they could adapt to cold Delmas, M., Gunnell, Y., Braucher, R., Calvet, M., Bourles, D., 2008. Exposure age and abrupt environmental changes, why did they fail to adapt to chronology of the last glaciation in the eastern Pyrenees. Quaternary Research 69, 231e241. changes of H4, characterised by cold and increased humidity? This Delmas, M., Calvet, M., Gunnell, Y., Braucher, R., Bourles, D., 2011. Palaeogeography may be explained either by competition with AMH, migrating and 10Be exposure age chronology of Middle and Late Pleistocene glacier southwards from Europe, or by the cultural limitations of a human systems in the northern Pyrenees: implications for reconstructing regional palaeoclimates. Palaeogeography, Palaeoclimatology, Palaeoecology 305, group incapable of adapting with innovative responses to the new 109e122. environmental and cultural conditions. The expansion of AMH Delmas, M., Calvet, M., Gunnell, Y., Braucher, R., Bourles, D., 2012. Les glaciations quaternaires dans les Pyren ees Ariegeoises: approche historiographique, during the cold and dry S-II and S-III shows the relationships be- fl donnes paleogeographiques et chronologiques nouvelles. Quaternaire 23, tween glacier advance and human ows from northern Europe 61e81. during the Late Pleistocene. Díez-Martín, F., Sanchez Yustos, P., Gomez Gonzalez, J.A., Gomez de la Rua, D., 2008. We must get a better chronological and environmental knowl- Earlier palaeolithic settlement patterns: landscape archaeology on the river Duero Basin plateaus (Castilla y Leon, Spain). Journal of World Prehistory 21, edge on glacial evolution, because at present our only reliable 103e137. knowledge relates to the extent of glaciers. It is necessary to Díez-Martín, F., Sanchez, P., Gomez, J.A., Gomez De La Rúa, D., Yravedra, J., Díaz, I., establish links between glaciers and cultural records as well as 2011. La ocupacion Neandertal en El Can~on de la Horadada (Mave, Palencia). e achieving a more accurate chronology between glacial and cultural Nuevas perspectivas arqueologicas en Cueva Corazon. Munibe 62, 65 85. d'Errico, F., Sanchez Goni,~ M.F., 2003. Neandertal extinction and the millennial scale changes. Effort must be made to develop an accurate glacial climatic variability of OIS 3. Quaternary Science Reviews 22, 769e788. chronostratigraphy. Fernandez-Mosquera, D., Martí, K., Vidal-Romaní, J.R., Weigel, A., 2000. Late Pleis- tocene deglaciation chronology in the NW of the Iberian Peninsula using cosmic-ray produced 21Ne in quartz. Nuclear Instruments and Methods in Physics Research 172, 832e837. Acknowledgements Finlayson, C., 2004. Neanderthals and Modern Humans. An Ecological and Evolu- tionary Perspective. Cambridge University Press, Cambridge. This study was supported by research funds from R þ D Pro- Finlayson, C., Carrion, J.S., 2007. Rapid ecological turnover and its impact on gramme of the Spanish Ministry of Science and Innovation (project Neanderthal and other human populations. Trends in Ecology and Evolution 22 (4), 213e222. CGL-2010-19729), and the Spanish Ministry of the Environment Finlayson, C., Giles, F., 2000. The Southern iberian Peninsulain the late Pleistocene: (project OAPN-053/2010). Authors express their gratitude to two geography, ecology and human occupation. In: Stringer, C.B., Barton, R.N., anonymous referees who helped us to improvement the Finlayson, C. (Eds.), Nenaderthals on the Edge. Oxbow books, Oxford, pp. 139e154. manuscript. Finlayson, C., Fa, D., Jimenez Espejo, F., Carrion, J.S., Finlayson, G., Giles Pacheco, F., Rodríguez Vidal, J., Stringer, C., Martínez Ruíz, F., 2008. Gorhams cave, Gibraltar. The persistence of a Neanderthal population. Quaternary International 181, References 64e71. Florineth, D., Schlüchter, C., 2000. Alpine evidence for atmospheric circulation patterns in Europe during the Last Glacial Maximum. Quaternary Research 54, Alonso, F., Arenillas, M., Saenz, C., 1982. La morfología glaciar en las Montanas~ de 295e308. Castilla La Vieja y Leon. In: I Congreso de Geografía de Castilla-Leon. Consejo Frochoso, M., 1990. Geomorfología del Valle del Nansa. Universidad de Cantabria, General de Castilla y Leon, Burgos, pp. 23e41. Santander. Alvarez-Lao, D.J., García, N., 2011. Geographical distribution of Pleistocene cold- Frochoso, M., Castan~on, J.C., 1996. El relieve heredado de la glaciacion cuaternaria en adapted large mammal faunas in the Iberian Peninsula. Quaternary Interna- el macizo de Pena~ Prieta (Cordillera Cantabrica). Polígonos 6, 25e43. tional 233, 159e170. Frochoso, M., Castan~on, J.C., 1998. El relieve glaciar de la Cordillera Cantabrica. In: Andrieu, V., Hubschman, J., Jalut, G., Herail, G., 1988. Chronologie de la deglaciation Gomez-Ortiz, A., Perez-Alberti, A. (Eds.), Las huellas glaciares de las montanas~ des Pyren es françaises. Dynamique de sedimentation et contenu pollinique des espanolas.~ Universidad de Santiago de Compostela, Santiago, pp. 65e137. paleolacs: application al ’interpretation du retrait glaciaire. Bulletin de l'Asso- Frochoso, M., Gonzalez-Pellejero, R., Allende, F., 2012. Pleistocene glacial ciation Française pour l’Etude du Quaternaire 2/3, 55e67. morphology and timing of Last Glacial Cycle in Cantabrian Mountains (Northern Baena, J., Cabrera, V., Carrion, E., 2004. Las sociedades neandertales durante el final Spain): new chronological data from the Ason area. Central European Journal of del Paleolítico Medio. Kobie 8, 91e140. Geosciences 5 (1), 12e27. Baldeon, A., 1993. El yacimiento de Lezetxiki (Gipuzkoa, País Vasco). Los niveles Frochoso, M., Gonzalez-Pellejero, R., Allende, F., 2013. Pleistocene glacial musterienses. Munibe 45, 3e97. morphology and timing of Last Glacial Cycle in Cantabrian Mountains (Northern Basabe, J.M., 1973. Dientes humanos del Musteriense de Axlor (Dima, Vizcaya). Spain): new chronological data from the Ason Area. Central European Journal of Trabajos de Antropología 16 (4), 187e207. Geosciences 5 (1), 12e27. Banks, W.E., d’Errico, F., Peterson, A.T., Kageyama, M., Sima, A., Sanchez-Go ni,~ M.F., Gale, S.J., Hoare, P.G., 1997. The glacial history of the Northwest Picos de Europa on 2008. Neanderthal extinction by competitive exclusion. PLoS One 3 (12), e3972. Northern Spain. Zeitschrift fur Geomorphologie 41, 81e96. Bermúdez de Castro, J.M., Saenz de Buruaga, A., 1999. Etude preliminaire du site Garcia Garriga, J., Martínez Molina, K., Baena, J., 2012. Neanderthal survival in the pleistocene superieur a hominide d'Arrillor (Pays Basque, Espagne). L'Antro- North of the Iberian Peninsula? reflections from a Catalan and Cantabrian pologie 103 (4), 633e639. perspective. Journal of World Prehistory 25 (2), 81e121. Cabrera, V., Arrizabalaga, A., Bernaldo de Quiros, F., Maíllo, J.M., 2004. La transicion García-Ruíz, J.M., Valero, B.L., Martí, C., Gonzalez, P., 2003. Asynchroneity of al paleolítico superior y la evolucion de los contextos aurinacienses~ (50.000- maximum glacier advances in the central Spanish Pyrenees. Journal of Qua- 27.000 BP). Kobie 8, 141e208. ternary Science 18, 61e72. Cabrera, V., Bernaldo de Quiros, F., 1990. Donnees sur la transition entre le García-Ruíz, J.M., Moreno, A., Gonzalez, P., Valero, B., Martí, C., 2010. La cronología Paleolithique moyen et le Paleolithique superieur de la region cantabrique: del último ciclo glaciar en las montanas~ del sur de Europa. Una revision. Cua- revision critique. In: Farizy, C. (Ed.), Paleolithique moyen recent et Paleolithique ternario y Geomorfología 24 (1e2), 35e46. superieur ancien en Europe. Ruptures et transitions: examen critique des García-Ruíz, J.M., Martí-Bono, C., Pena,~ J.L., Sancho, C., Rhodes, E.J., Valero- documents archeologiques, pp. 185e188. Musee de Prehistoire d’Ile de France, Garces, B.L., Gonzalez-Samp eriz, P., Moreno, A., 2013. Glacial and fluvial de- Nemours. posits in the Aragon valley, central-western Pyrenees: chronology of the Pyr- Calvet, M., Delmas, M., Gunnell, Y., Braucher, R., Bourles, D., 2011. Recent advances in enean late Pleistocene glaciers. Geografiska Annaler: series A. Physical research on Quaternary glaciations in the Pyrenees. In: Ehlers, J., Gibbard, P.L., Geography 95 (1), 15e32. Hughes, P.D. (Eds.), Quaternary Glaciations-extent and Chronology, De- Garralda, M.D., 2005. Los Neandertales en la Península Iberica. Munibe 57, 289e314. velopments in Quaternary Sciences, vol. 15. Elsevier, Amsterdam, pp. 127e139. Gonzalez Samperiz, P., Valero, B.L., Moreno, A., Jalut, G., García-Ruíz, J.M., Martí, C., Castan~on, J.C., Frochoso, M., 1992. La glaciacion Würm en las montanas~ cantabricas. Delgado, A., Navas, A., Otto, T., Dedoubat, J., 2006. Climate variability in the In: Cearreta, C., Ugarte, F. (Eds.), The Late Quaternary in the Western Pyrenean Spanish Pyrenees during the last 30,000 yr revealed by the El Portalet sequence. Region. Universidad del País Vasco, Bilbao, pp. 319e332. Quaternary Research 66, 38e52. Castan~on, J.C., Frochoso, M., 1996. Hugo Obermaier y el glaciarismo pleistoceno. In: Gonzalez-Trueba, J.J., 2007a. El pasiaje natural del macizo central de los Picos de Moure, A. (Ed.), El hombre fosil 80 anos~ despues. Universidad de Cantabria, Europa. Gobierno de Cantabria, Santander. Santander, pp. 153e176. Gonzalez-Trueba, J.J., 2007b. Geomorfología del Macizo Central del Parque Nacional Dansgaard, W., Johnsen, S.J., Clausen, H.B., Dahl-Jensen, D., Gundestrup, N.S., de Picos de Europa. OAPN-Ministerio de Medio Ambiente, Madrid. Hammer, C.U., Hvidberg, C.S., Steffensen, J.P., Sveinbiornnsd€ ottir, A.E., Jouzel, J., Gonzalez-Trueba, J.J., Serrano, E., 2010. Geomorfología del Macizo Oriental del Bond, G., 1993. Evidence for general instability of past climate from a 250-Kyr Parque Nacional Picos de Europa. OAPN-Ministerio de Medio Ambiente, Madrid. ice-core record. Nature 364, 218e220. E. Serrano et al. / Quaternary International 364 (2015) 22e34 33

Gutierrez, A., Serrano, E., 1996. El yacimiento del Paleolítico Medio de la “Ermita del 2010. The last glaciation in the Picos de Europa Nacional Park (Cantabrian Abra” (Campoo De Suso, Cantabria). Aproximacion cultural, cronologica y Mountains, nortern Spain). Journal of Quaternary Science 25, 1076e1091. geomorfologica. Cuaternario y Geomorfología 12 (3e4), 27e39. Munoz~ Sobrino, C., Ramil-Rego, P., Rodriguez Guitian, M.A., 2001. Vegetation in the Gutierrez, A., Serrano, E., 2007. Nuevas aportaciones al estudio del paleolítico medio mountains of northwest Iberia during the last glacial-interglacial transition. en el valle de Campoo (Cantabria): aproximacion cultural, cronologica y geo- Vegetation History and Archaeobotany 10, 7e21. morfologica. Zona Arqueologica 7 (2), 234e251. Nussbaum, F., Gigax, F., 1953. La glaciation quaternaire dans la Cordillera Cantab- Gonzalez Trueba, J.J., Serrano, E., García, M., Gomez, M., 2011. Geomorfología del rique (Espagne du Nord). Estudios Geograficos 14, 261e270. Macizo Occidental del Parque Nacional Picos de Europa. OAPN-Ministerio de Obermaier, H., 1914. Estudio de los glaciares de los Picos de Europa. Museo Nacional Medio Ambiente, Madrid. de Ciencias Naturales, Madrid. Gonzalez-Urquijo, J.E., Ibanez,~ J.J., Ríos, J., Bourguignon, L., 2006. Aportes de las Palacios, D., De Marcos, J., Vazquez, L., 2011. Last glacial maximum and deglaciation nuevas excavaciones en Axlor sobre el final del Paleolítico Medio. In: of Sierra de Gredos, Central Iberian Peninsula. Quaternary International 233, Cabrera, V., Bernardo de Quiros, F., Maillo, J.M. (Eds.), En el centenario de 16e26. la Cueva del. Castillo: el ocaso de los neandertales. UNED, Madrid, Pallas, R., Rodes, A., Braucher, R., Carcailler, J., Ortuno,~ M., Bordonau, J., Bourles, D., pp. 269e290. Vilaplana, J.M., Masana, E., Santanach, P., 2006. Late Pleistocene and Holocene Havarti, K., 2012. What happened to the Neanderthals? Nature Education Knowl- glaciation in the Pyrenees: a critical review and new evidence from 10Be edge 3 (10), 13e14. exposure ages, south-central Pyrenees. Quaternary Science Reviews 25, Hernandez-Pacheco, F., 1944. Fisiografía, geología y glaciarismo cuaternario en las 2937e2963. montanas~ de Reinosa. Memorias de la Real Academia de Ciencias Físicas Exactas Pallas, R., Rodes, A., Braucher, R., Bourles, D., Delmas, M., Calvet, M., Gunnell, Y., y Naturales, Madrid. 2010. Small, isolated glacial catchments as priority targets for cosmogenic Hernandez-Pacheco, F., 1961. La pequena~ cuenca glaciar de Pena~ Lusa en la surface exposure dating of Pleistocene climate fluctuations, southeastern Pyr- Cordillera Cantabrica, Santander. Boletín Real Sociedad Espanola~ de Historia enees. Geology 38, 891e894. Natural secc. Geología 59 (2), 191e205. Pellitero, R., 2009. Application of an alpine geomorphological mapping system to an Hoffecker, J.F., 2009. The spread of modern humans in Europe. Proceedings, Na- Atlantic mountain environment: the Curavacas Massif (Cantabrian Range, tional Academy of Sciences 106e38, 16040e16045. Northwest Spain). Journal of Maps 5, 194e205. Hughes, P.D., Woodward, J.C., 2008. Timing of glaciation in the Mediterranean Pellitero, R., 2011. El complejo morrenico de Vega Naranco (Leon, Cordillera Can- mountains during the last cold stage. Journal of Quaternary Science 23, tabrica). Evolucion y correlacion de fases glaciares durante el Último Maximo 575e588. Glaciar. In: Turu, V., Constante, A. (Eds.), El Cuaternario en Espana~ y areas afines, Hughes, P.D., Woodward, J.C., Gibbard, P.L., 2006. Glacial history of the Mediterra- Avances en 2011. AEQUA-Fundacion M. Chevalier, Andorra, pp. 7e9. nean mountains. Progress in Physical Geography 30, 334e364. Pellitero, R., 2013. Evolucion finicuaternaria del glaciarismo en el macizo de Fuentes Iriarte, M.J., Gomez Orellana, L., Munoz~ Sobrino, C., Ramil, P., Arrizabalaga, A., 2005. Carrionas (Cordillera Cantabrica), propuesta cronologica y paleoambiental. La dinamica de la vegetacion durante la transicion cultural del Paleolítico medio Cuaternario y Geomorfología 27 (1e2), 71e90. al Paleolítico superior. In: Montes, R., Laceras, J.A. (Eds.), Neandertales can- Pellitero, R., 2014. Geomorphology and geomorphological landscapes of Fuentes tabricos, estado de la cuestion. Museo Nacional y Centro de Investigacion de Carrionas. Journal of Maps 10, 313e323. Altamira, Santander, pp. 231e253. Pena,~ J.L., Sancho, C., Lewis, C., McDonald, E., Rhodes, E., 2004. Datos cronologicos de Ivy-Ochs, S., Kerschner, H., Reuther, A., Preusser, F., Heine, K., Maisch, M., las morrenas terminales del glaciar del Gallego y su relacion con las terrazas Kubik, P.W., Schlüchter, C., 2008. Chronology of the last glacial cycle in the fluvioglaciares (Pirineo de Huesca). In: Pena,~ J.L., Longares, L.A., Sanchez, M. European Alps. Journal of Quaternary Science 23, 559e573. (Eds.), Geografía Física de Aragon. Universidad de Zaragoza e Institucion Fer- Jalut, G., Montserrat, J., Fontugne, M., Delibrias, G., Vilaplana, J., Julia, R., 1992. Glacial nando El Catolico, Zaragoza, pp. 71e84. to interglacial vegetation changes in the northern and southern Pyrenees: Penck, A., 1897. Die Picos de Europa und das kantabrische Gebirge. Geographische deglaciation, vegetation cover and chronology. Quaternary Science Reviews 11, Zeitschrift 278e281. 449e480. Perez-Alberti, A., Valcarcel, M., Blanco, R., 2004. Pleistocene glaciation in Spain. In: Jalut, G., Turu, I., Michels, V., Dedoubat, J.J., Otto, T., Ezquerra, J., Fontugne, M., Ehlers, J., Gibbard, P.L. (Eds.), Quaternary Glaciations- Extent and Chronology. Belet, J.M., Bonnet, L., García de Celis, A., Redondo-Vega, J.M., Vidal-Romaní, J.R., Elsevier, London, pp. 389e394. Santos, L., 2010. Palaeoenvironmental studies in NW Iberia (Cantabrian Range): Perez-Alberti, A., Valcarcel, M., Martini, P., Pascucci, V., Andreucci, S., 2011. Upper vegetation history and synthetic approach of the last deglaciation phases in the Pleistocene glacial valley-junction sediments at Pias, Trevinca Mountains, NW Western Mediterranean. Palaeogeography, Palaeoclimatology, Palaeoecology Spain. In: Martini, I.P., French, H.M., Perez Alberti, A. (Eds.), Ice-marginal and 297, 330e350. Periglacial Processes and Sediments, Geological Society, Special Publications, Jimenez, M., Farias, P., 2002. New radiometric and geomorphologic evidences of a 354, London, pp. 93e110. last glacial maximum older than 18 ka in SW European mountains: the example Preusser, F., Blei, A., Graf, H., Schlüchter, C., 2007. Luminiscence dating of Würmian of Redes Natural Park (Cantabrian Mountains, NW Spain). Geodinamica Acta 15, (Weichselian) proglacial sediments from Switzerland: methodological aspects 93e101. and stratigraphical conclusions. Boreas 36, 130e142. Jimenez, M., Ruíz, M.B., Farias, P., Dorado, M., Gil, M.J., Valdeomillos, A., 2002. Rasilla de la, M., Straus, L.G., 2004. El poblamiento en la region cantabrica en torno Palaeoenvironmental research in Cantabrian Mountains: Redes Natural Park al Último Maximo Glaciar: gravetiense y solutrense. Kobie 8, 209e242. and Comella basin. In: Ruiz, B., Dorado, M., Valdeolmillos, A., Gil, M.J., Bardají, T., Rico, I., 2012. Glacial morphology and evolution in the Arritzaga valley (Aralar Bustamante, I., Martínez, J. (Eds.), Quaternary Climatic Changes and Environ- range, Gipuzkoa). Cuaternario y Geomorfología 25 (1e2), 83e104. mental Crises in the Mediterranean Region. Universidad de Alcala de Henares, Rodríguez, L., Jimenez, M., Domínguez, M.J., Rico, M.T., Valero, B., 2011. Last degla- Madrid, pp. 229e240. ciation in north-western Spain: new chronological and geomorphologic evi- Jimenez, M., Rodríguez, L., García-Ruiz, J.M., Domínguez, M.J., Farias, P., Valero, B., dence from the Sanabria region. Geomorphology 135, 48e65. Moreno, A., Rico, M., Valcarcel, M., 2013. A review of glacial geomorphology and Rodríguez-Rodríguez, L., Jimenez, M., Dominguez, M.J., Aranburu, A., 2014. Research chronology in northern Spain: timing and regional variability during the last history on glacial geomorphology and geochronology of the Cantabrian glacial cycle. Geomorphology 196, 50e64. Mountains, north Iberia (43-42N/7-2W). Quaternary International. http:// Lalueza, C., Sampietro, M.L., Caramelli, D., Puder, Y., Lari, M., Calafell, F., Martinez- dx.doi.org/10.1016/j.quaint.2014.06.007. Maza, C., Bastir, M., Fortea, J., Rasilla, M., Bertranpetit, J., Rosas, A., 2005. Ruddimann, W.F., McYntire, A., 1981. The North Atlantic Ocean during the last Neandertal Evolutionary Genetics; Mitochondrial DNA data from the Iberian deglaciation. Palaeogeogrphy, Palaeoclimatolgy, Palaeoecology 35, 145e214. Peninsula. Molecular Biology and Evolution 22 (4), 1e5. Saenz, C., 1935. Restos glaciares de Castro Valnera. Boletín Real Sociedad Espanola~ Lewis, C.J., McDonald, E.V., Sancho, C., Pena,~ J.L., Rhodes, E.J., 2009. Climatic impli- de Historia Natural 35, 236e237. cations of correlated upper Pleistocene glacial and fluvial deposits on the Cinca Sanchez Goni,~ M.F., d’Errico, F., 2005. La historia de la vegetacion y del clima del and Gallego Rivers (NE Spain) based on OSL dating and soil stratigraphy. Global último ciclo climatico (OIS 5-OIS 1/140-10 Ka. BP) en la Península Iberica y su and Planetary Change 67, 141e152. posible impacto sobre los grupos paleolíticos. In: Montes, R., Laceras, J.A. (Eds.), Lotze, F., 1962. Pleistocene Vergletscherungen im ostteil des Kantabrischen gebirges Neandertales cantabricos, estado de la cuestion. Museo Nacional y Centro de (Spanien). Akademie der Wissencshaften und der Literatur, Mainz, p. 22. Investigacion de Altamira, Santander, pp. 115e129. Maíllo, J.M., Cabrera, V., Bernaldo de Quiros, F., 2004. Le debitage lamellaire dans le Sancho, C., Pena,~ J.L., Lewis, C., McDonald, E., Rhodes, E., 2008. Actividad glaciar y Mousterien final de Cantabrie (Espagne): le cas de El Castillo et Cueva Morin. desarrollo de terrazas durante el Pleistoceo Superior en Pirineos-Cuenca del L'Anthropologie 108, 367e393. Ebro. Geo-Temas 10, 763e766. Mardones, M., Jalut, G., 1983. La tourbiere de Biscaye (alt. 409 m, Hautes Pyren ees): Sancho, C., Pena-Monn~ e, J.L., Rhodes, E., Arenas, C., Pardo, G., García-Ruiz, J.M., approche paleoecologique des 45,000 dernieres annees. Pollen et Spores 25, Martí-Bono, C.E., 2011. El registro glaciolacustre de Linas de Broto (Valle del Ara, 163e212. Pirineo Central, Huesca): Nuevas aportaciones. In: El Cuaternario en Espana~ y Maroto, J., Vaquero, M., Arrizabalaga, A., Baena, J., Baquedano, E., Jorda, J., Julia, R., areas afines. avances en 2011. Fundacion Henri Chevalier, Andorra, pp. 11e14. Montes, R., Van Der Plicht, J., Rasines, P., Wood, R., 2012. Current Issues. In: Late Sepulchre, P., Ramstein, G., Kageyama, M., Vanhaeren, M., Krinner, G., Sanchez- Middle Palaeolithic Chronology: New Assessments from Northern Iberia. Qua- Goni,~ M.F., d'Errico, F., 2007. H4 abrupt event and late Neanderthal presence in ternary International, vol. 247, pp. 15e25. Iberia. Earth and Planetary Science Letters 258, 283e292. Martínez de Pison, E., Arenillas, M., 1979. Algunos problemas de morfología glaciar Seret, G., Dricot, J., Wansard, G., 1990. Evidence for an early glacial maximum in the en la Espana~ Atlantica. Acta Geologica Hispanica 14, 445e450. French Vosgues during the last glacial cycle. Nature 346, 453e456. Moreno, A., Valero, B.L., Jimenez, M., Domínguez, M.J., Mata, M.P., Navas, A., Serrano, E., 1991. Glacial evolution of the Upper Gallego valley (Panticosa mountains Gonzalez-Samp eriz, P., Stoll, H., Farias, P., Morellon, M., Corella, J.P., Rico, M.T., and Ribera de Biescas, Aragonese Pyrenees, Spain). Pirineos 138, 83e104. 34 E. Serrano et al. / Quaternary International 364 (2015) 22e34

Serrano, E., Gonzalez-Trueba, J.J., 2002. Morfología y evolucion glaciar en los Picos Straus, L.G., Gonzalez-Morales, M.R., Fano, M.A., García-Gelabert, M.P., 2002. Last de Europa. In: Redondo, J.M., Gomez, A., Gonzalez, R.S., Carrera, P. (Eds.), El glacial human settlement in eastern cantabria (Northern Spain). Journal of modelado de origen glaciar en las montanas~ leonesas. Universidad de Leon, Archaeological Science 29, 1403e1414. Leon, pp. 249e268. Turú, V., Boulton, G.S., Ros, X., Pena,~ J.L., Martí, C.E., Bordonau, J., Serrano, E., Serrano, E., Gutierrez, A., 2002. El glaciarismo pleistoceno en la vertiente meridional Sancho, C., Constante, A., Pous, J., Fabregas, J., Gonzalez, J.J., Palomar, J., de la Cordillera Cantabrica (montanas~ de Palencia, Cantabria y Burgos). In: Herrero, R., García-Ruiz, J.M., 2007. Structure des grands bassins glaciaires dans Geomorfología y Paisaje. VII Reunion Nacional de Geomorfología. SEG-Uni- le Nord de la Peninsule Iberique: comparaison entre les vallees d’Andorre versidad de Valladolid, Valladolid, pp. 91e161. (Pyren ees Orientales), du Gallego (Pyren ees Centrales) et du Trueba (Chaîne Serrano, E., Gonzalez-Trueba, J.J., Turu, V., Ros, X., 2011. Cronología glaciar pleisto- Cantabrique). Quaternaire 18e4, 309e325. cena en el valle de Trueba (Cordillera Cantabrica): primeras dataciones. In: Ugarte, F.M., 1992. Glacial and periglacial morphogenesis in the Basque Mountains. Turu, V., Constante, A. (Eds.), El Cuaternario en Espana~ y Areas Aﬁnes, pp. 3e6. In: Cearreta, A., Ugarte, F.M. (Eds.), The Late Quaternary in the Western Pyr- Avances en 2011. AEQUA-Fundacion M. Chevalier, Andorra. enean Region. Universidad del País Vasco, Bilbao, pp. 235e255. Serrano, E., Gonzalez-Trueba, J.J., Gonzalez-Gonz alez, M., 2012. Mountain glaciation Uzquiano, P., 2005. El registro antracologico durante la transicion Musteriense- and paleoclimate reconstruction in the Picos de Europa (Iberian Peninsula, SW Paleolítico Superior Inicial en la Region Cantabrica: vegetacion, paleoambiente Europe). Quaternary Research 78 (2), 303e314. y modos de vida alrededor del fuego. In: Montes, R., Laceras, J.A. (Eds.), Nean- Serrano, E., Gonzalez-Trueba, J.J., Pellitero, R., Gonzalez-García, M., y Gomez- derthales cantabricos, estado de la cuestion. Museo Nacional y Centro de Lende, M., 2013a. Quaternary glacial evolution in the Central Cantabrian Investigacion de Altamira, Santander, pp. 255e274. Mountains (Northern Spain). Geomorphology 196, 65e82. Uzquiano, P., Arbizu, M., Arsuaga, J.L., Adan, G., Aranburu, A., Iriarte, E., 2008. Datos Serrano, E., Gomez-Lende, M., Gonzalez-Trueba, J.J., Turu, V., Ros, X., 2013b. Fluc- paleoﬂorísticos en la Cuenca media del Nalon entre 40-32 Ka BP: Antra- tuaciones glaciares pleistocenas y cronología en las Montanas~ Pasiegas coanalisis de la Cueva del Conde (Santo Adriano, Asturias). Cuaternario y (Cordillera Cantabrica). Cuaternario y Geomorfología 27, 91e110. Geomorfología 22 (3e4), 121e133. Smart, P.L., 1986. Origin and development of glacio-karst closed depressions in the Uzquiano, P., Yravedra, J., Ruiz, B., Gil, M.J., Sese, C., 2012. Human behaviour and Picos de Europa, Spain. Zeitschrift für Geomorphologie 30, 423e443. adaptations to MIS3 environments trends (>53-30 ka BP) at Esquilleu cave Stewart, J.R., 2005. The ecology and adaptation of Neanderthals during the non- (Cantabria, Northern Spain). Quaternary International 252, 82e89. analogue environment of Oxygen Isotope Stage 3. Quaternary International Villa, E., Stoll, H., Farias, P., Adrados, L., Edwards, R.L., Cheng, H., 2013. Age and 137, 35e46. significance of the Quaternary cemented deposits of the Duje Valley (Picos de Straus, L.G., 2000. A quarter ecentury of research on the Solutrean of vasco- Europa, Northern Spain). Quaternary Research 79, 1e5. cantabria, Iberia and beyond. Journal of Anthropology Research 56, 39e58. Yravedra, J., Díez-Martín, F., Sanchez, P., Gomez De La Rúa, D., Díaz, I., Gomez, J.A., Straus, L.G., 2005. The Upper Paleolithic of Cantabrian, Spain. Evolutionary An- 2013. Estrategias de subsistencia durante el Paleolítico Medio en la submeseta thropology 14, 145e158. norte. La Cueva Corazon (Mave, Palencia). Zephyrus 61, 45e60.