Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada (Valencia, Spain): The evidence from charred plant and micromammal remains

Quaternary Science Reviews xxx (xxxx) xxx

Contents lists available at ScienceDirect

Quaternary Science Reviews journal homepage: www.elsevier.com/locate/quascirev

Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada (Valencia, Spain): The evidence from charred plant and micromammal remains
n Marco a, *, Pere Guillem Calatayud b, Aleix Eixea a, Yolanda Carrio ~o Zilha ~o d, e, f, Carmen M. Martínez-Varea a, Carmen Tormo c, Ernestina Badal a, Joa a Valentín Villaverde a ria, Arqueologia i Histo ria Antiga, Av. Blasco Iban ~ ez 28, 46010, Valencia, PREMEDOC - GIUV2015-213, Universitat de Val encia, Departament de Prehisto Spain b
Area de Arqueología y Paleontología, IVCR, CulurArts Generalitat, Genaro Lahuerta 25, 46010, Valencia, Spain c
n Prehisto
rica, C/Corona, 36, Valencia, Spain Museo de Prehistoria, Servicio de Investigacio d
Catalana de Recerca i Estudis Avançats (ICREA), Passeig Lluís Companys 23, 08010, Barcelona, Spain Institucio e ria i Arqueologia, Facultat de Geografia i Histo ria, c/Montalegre 6, 08001, Barcelona, Spain Universitat de Barcelona, Departament d’Histo f UNIARQ - Centro de Arqueologia da Universidade de Lisboa, Faculdade de Letras de Lisboa, Universidade de Lisboa, Alameda da Universidade, 1600-214, Lisboa, Portugal

a r t i c l e i n f o

a b s t r a c t

Article history: Received 24 July 2018 Received in revised form 29 November 2018 Accepted 29 November 2018 Available online xxx

The Abrigo de la Quebrada rock shelter was occupied by Neanderthal groups during the early Upper Pleistocene, yielding evidence for their subsistence practices and local resource exploitation. This paper focuses on the plant macroremains and the micromammals, which provide information about occupation patterns, the surrounding landscape, the use of resources, and the environment. Mountain pine forests and permanent grass formations containing humid zones and open spaces that would have harboured an eurythermal microfauna were the dominant landscape type. Cold-climate pines provided most of the firewood. The data are consistent with a recurrent, seasonal occupation pattern, in which the rock shelter was used for short periods in the context of an annual round characterized by a high degree of mobility. © 2018 Elsevier Ltd. All rights reserved.

Keywords: Abrigo de la Quebrada Neanderthal Woodland exploitation Charcoal Seeds Micromammals Taphonomy

1. Introduction There is no shortage of literature on the persistence of Nean
n derthals in the Iberian Peninsula (Finlayson et al., 2008; Galva et al., 2014; Higham et al., 2014; Hublin, 2015; Jennings et al., €ris et al., 2011; Maroto et al., 2012; Zilh~ 2011; Jo ao et al., 2017), their cognitive and organisational abilities (Conard et al., 2009; d’Errico et al., 2003; Hoffmann et al., 2018; Jaubert et al., 2016; ~o, 2007; McBrearty and Brooks, 2000; Peresani et al., 2011; Zilha Zilh~ ao et al., 2010), including how they exploited their environment

* Corresponding author.
n E-mail addresses: [email protected], [email protected] (Y. Carrio Marco).

and made use of regional resources (Delagnes and Rendu, 2011;
blot-Augustins, 1993, 1997; Gaudzinski, 2006; Geneste, 1985, Fe 1988; Grayson and Delpech, 2003; Kuhn, 1995; Rendu et al., 2011; Roebroeks, 1988; Stiner et al., 1999; Turq, 1992), or their ability to make fire (Dibble et al., 2017, 2018; Sorensen, 2017; Sorensen et al., 2018). Knowledge of the environmental conditions is essential when addressing such matters. Thanks to regional pollen sequences and other proxies, Mediterranean Iberia's Middle Palaeolithic environments are well known (Cacho et al., 2006; Desprat ~ i, 2008; Moreno et al., et al., 2013; Fletcher and S
anchez Gon ~ i et al., 2008, 2013), and the correlation of the 2005; S
anchez Gon regional sequence with the Greeland ice record (Andersen et al., 2004; Rasmussen et al., 2006, 2014) is well established. However, there are still large gaps in the information about the immediate

https://doi.org/10.1016/j.quascirev.2018.11.032 0277-3791/© 2018 Elsevier Ltd. All rights reserved.


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

2

environment of settlement localities and, despite the basic role it
, 2002; Badal et al., 2011; must have played in subsistence (Allue Gonzalez-Samperiz et al., 2010; Uzquiano et al., 2008, 2012, VidalMatutano et al., 2015, 2017), the use of plants. This is largely due to preservation problems and, often, the unsuitability of the sampling protocols used. The Abrigo de la Quebrada (Chelva, Valencia) was discovered in 2004. Its archaeological excavation, completed in 2015, revealed an early Upper Pleistocene sequence of human occupations, with some hiatuses. In the region, paleoenvironmental data for this time interval remain scarce (Badal et al., 2012). Here, we combine the examination of anthropogenic (charcoal and seeds from fire features) and non-anthropogenic (naturally accumulated micromammal remains, which, due to their sensivity to climate change, are excellent sources of environmental information) data to reconstruct the site's landscape context, the use of the plant resources it offered, and how both changed through time. Prior anthracological work (Badal et al., 2012; Vidal-Matutano, ~o et al., 2016, 2017) 2017; Vidal-Matutano et al., 2015, 2017; Zilha has suggested that, in Mediterranean Iberia, very open coniferous formations, mainly pines and junipers, prevailed through the Mousterian, so it is of particular interest to ascertain how the resources of the forests were managed and how wood gathering adapted to their physiognomy and composition. For our landscape reconstruction, we redress the cultural bias inherent to the wood charcoal data through consideration of the composition of the microfaunal assemblage. Together with the archaeological data, ́ ́ these two records make for one of the most informative stratigraphic sequences currently known in the region. 2. The site of Abrigo de La Quebrada 2.1. Biogeographical context The Abrigo de la Quebrada is located near Chelva, 65 km northwest of the city of Valencia (Fig. 1). The site lies on the left-hand side of the Rambla de Ahillas ravine, 728 m asl (above present sea level). It is a sheltered, fairly horizontal, south-dipping, 38 m-long and 29 m-wide platform. Given its position (facing NW and at the bottom of a narrow, steep gorge), direct exposure to sunlight is limited. The Rambla de Ahillas ravine, which leads to a dead end and forms a sort of natural trap, opens onto a wide plain known as Plano de Arquela (Villaverde et al., 2017). Currently, bioclimatic conditions at the locality are mesoMediterranean (average annual temperature of 14.8
C) and dry (average annual precipitation of 400 mm). The vegetation cover features a strong duality between the ravine's bottom and the surrounding mountains and plains. Under the microclimate of the ravine, the vegetation is leafy and riverside plants, such as oleander (Nerium oleander), willow (Salix sp.), reeds (Juncus sp.), wild roses (Rosa sp.) and butcher's broom (Ruscus aculeatus) proliferate, in association with climbing plants such as honeysuckle (Lonicera implexa), wild madder (Rubia peregrina), brambles (Rubus sp.) and ivy (Hedera helix). Poplars (Populus sp.) and tamarisks (Tamarix sp.) thrive in areas where the ravine is wider. Along the edges of the ravine and in its sunnier parts, taller shrubs and trees form small groves of typically Mediterranean species d pines (Pinus halepensis and P. pinaster) and holm oak (Quercus rotundifolia), accompanied by e.g. juniper (Juniperus oxycedrus, J. phoenicea), rosemary (Rosmarinus officinalis), kermes oak (Quercus coccifera), Mediterranean buckthorn (Rhamnus alaternus), grey-leaved cistus (Cistus albidus), and gorse (Ulex parviflorus) d among which such warm-loving taxa as the lentisk (Pistacia lentiscus) can also be found. The landscape around the shelter is varied, comprising both

very steep terrain and wide plains, and, accordingly, plant formations are also diverse. Combined with the configuration of the ravine, favourable to trap-hunting, the variety of animal and plant resources available in the different biotopes within reach underpins the attractiveness of the site for human occupation (Eixea et al., 2011e2012; Villaverde et al., 2017). 2.2. Stratigraphic framework The stratigraphy of the site is composed of a total of nine units distributed over a thickness of 3 m (Fig. 2, Table 1). Level I, at the top, is disturbed by penning. Below, levels II to V, VII, VIII and IX contain Middle Palaeolithic stone tool assemblages and faunal remains, levels III-V being the richest. Level VI, which is almost 1 m thick, is sterile. The preliminary results from sedimentological studies, in progress, suggest the succession accumulated through the operation of low intensity flooding episodes alternating with phases of mild cryoclastic wall degradation. In some areas, the percolation of water led to the formation of carbonated crusts, which also often coat the surfaces of archaeological remains. The bioturbation features and the non-anthropogenically accumulated microfaunal remains found alonsgide the archeological material denote phases of stasis and a generally slow sedimentation rate, especially in the upper levels. Level VI, however, is distinct (it formed rapidly in the context of higher-energy fluvial deposition), while the presence of large blocks in levels VII-VIII reflects recession of the overhang; currently, the drip line is in row 5 of the excavation grid (Eixea et al., 2011e2012). Radiocarbon accelerator mass spectrometry (AMS) dating of charcoal fragments yielded ages of 40,500 ± 530 BP (Beta-244003; ABA-treated) and >50,800 (OxA-24854; ABOx-processed) for level III, of 43,930 ± 750 BP (Beta- 244002; ABA-treated) and >51,600 (OxA-24855; ABOx-processed) for level IV, and of >47,100 (OxA25583; ABOx-processed) for level V (Villaverde et al., 2008; Eixea et al., 2011e2012; Real et al., 2018, Table 2). For archaeologically sterile level VI, we have two OSL results on the feldspar fraction of 80.0 ± 4.7 ka (C-L3900) and 83.2 ± 5.4 ka (C-L3898), placing its deposition in MIS-5a or MIS-5b (Real et al., 2018). These results constrain the deposition of levels IIeV to the interval between MIS5b and early MIS-3, and of levels VIIeVIII to MIS-5a, MIS-5b or earlier but within the Last Interglacial (as the small mammal assemblage therein is of Upper, not Middle Pleistocene composition; Tormo and Guillem, 2015). Thus, where the length of time represented by the deposit is concerned, the chronometric data are consistent with it representing (a) the ca.75,000 years comprised between the minimum radiocarbon ages for levels III-V and the onset of MIS-5e, in one extreme, (b) it all belonging in the 10,000 years of MIS-5a, in the other extreme, and (c) a number of intermediate scenarios. From an archeological standpoint, levels III-V are similar and contrast strongly with levels VII-IX in terms of lithic technology, raw material economy, occupation patterns and, albeit to a lesser extent, faunal composition (Villaverde et al., 2017). In levels III-V, stone tools were produced using discoid and recurrent centripetal Levallois methods; lateral sidescrapers, Levallois points, and Mousterian points are the dominant tooltypes. The abundance of cortical fragments, cores, flakes of different sizes and refitted elements suggest that knapping was carried out in situ. Raw-material was procured mostly within a radius of 5 km. Some flint coming from sources situated more than 100 km away is also found (Eixea et al., 2011, 2014, 2016; Villaverde et al., 2012). The faunal study of level IV identified remains from eight different families: Bovideae, Equideae, Cervidae, Leporidae, Testudinidae, Suidae, Rhinocerotidae, and Canidae, with ibex


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

3

Fig. 1. Location of Abrigo de la Quebrada. A. Geographical location. B. Overview of the ravine showing the position of the site. C. Overview of the shelter during excavation. D. Site plan and excavation grid (the Roman numerals indicate the level reached in each colour-coded area). (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.)

(Capra sp.), horse (Equus ferus), and deer (Cervus elaphus) being the most abundant taxa (Sanchis et al., 2013; Real et al., 2018). The fragmentation of lithic reduction sequences, the reuse of previously discarded pieces and the recycling of blanks and tools into cores suggest that the upper levels reflect frequent, repeated occupations with intensive on-site processing. A similar pattern is observed in other Middle Palaeolithic contexts of Mediterranean Spain, such as Cova Beneito (Martínez Valle, 1996), level IV of Bolomor (Blasco, 2008), and level O of Abric Romani (Gabucio et al., 2014). At Quebrada, the ungulate seasonality data suggest that the site was visited in spring and autumn. In level VIII, the lower density of lithics and fauna reflects more

sporadic and shorter occupations. Nevertheless, a faster sedimentation rate allowed a much better preservation of the record's spatial structure (Eixea et al., 2011e2012, Villaverde et al., 2017). 3. Material and methods 3.1. Charcoal analysis The anthracological analysis presented here concerns the complete sequence, except levels I-II (because of their observed, or suspected, lack of stratigraphic integrity) and IX (in which plant remains were too scant for quantititave assessment). The data for


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

4

Fig. 2. Abrigo de la Quebrada: plan of the surface of level IV (left) and stratigraphic section in square G3 (right).

Table 1 Abrigo de la Quebrada. Excavation and sampling information for the different stratigraphic levels. “Liters floted” refers to the volume of sediment collected for flotation, whence come the present study's plant charcoal and micromammal remains. Level

Thickness (in cm)

N. of spits

N. of combustion structures

Liters of sediment floated

II III IV V VI VII VIII

10e15 10e15 20e25 35e40 60e80 40e50 60e70

2 2 4 7 29 3 10

24 20 29 17 e 5 18

67 94 68 539 244 525 1047

Table 2 Abrigo de la Quebrada. Dating results for levels III to VI. Taxon

Level

Method

d13C

Age

Laboratory reference

Reference

Pinus Pinus Pinus Pinus Pinus e

III III IV IV V VI

C14 C14 C14 C14 C14 OSL

22,9 21,03 22,9 22,62 22,82 e

40500 ± 530 >50800 43930 ± 750 >51600 >47100 79000 ± 500

Beta-244003 OxA-24854 Beta-244002 OxA-24855 OxA-25583 C-L3898

Villaverde et al. (2008) Villaverde et al. (2008) Villaverde et al. (2008) Eixea et al., 2011e2012 Eixea et al., 2011e2012 Real et al. (2018)

e

VI

OSL

e

82000 ± 500

C-L3900

Real et al. (2018)

nigra-sylvestris nigra-sylvestris nigra-sylvestris nigra-sylvestris nigra-sylvestris

ABA ABOx ABA ABOx ABOx

levels III-IV expand on those preliminarily published by Badal et al. (2012). The material corresponds to dispersed fragments derived from combustion features denoted by ash-, black- or red-stained areas within which, however, no charcoal was preserved d either because of post-depositional scatter or because the wood was entirely reduced to ash. The charcoal for radiocarbon dating was manually recovered during the excavation or dry sieving of the sediments, for presubmission botanical identification of the samples. A 1 m2-column, >2500 L in total, was processed in a flotation machine with meshes of 1 mm and 0.25 mm to pick up the heavy residue and the light fraction, respectively. The number of charcoal fragments per

spit varies (generally, >100), but is in all cases sufficient to characterise the flora of the surroundings. The carpological remains are scarce: despite the sampling effort, only a few were retrieved, all in the lower levels. For standard analysis (Vernet et al., 1979), the charcoal was broken manually to produce fresh, diagnostic sections that were analysed under a Leica DM6000 M optical microscope with Brightfield, Darkfield and Polarisation contrast modes; the Leica Application Suite (LAS X) software platform was used for measurement. No chemical treatment was applied. A Hitachi S-4100 scanning electron microscope (SEM) with spotlight of field emission and QUANTAX 200 digital image acquisition system, held at


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

the University of Valencia's Central Service for the Support of Experimental Research (SCSIE) was used for the observation of specific features and to take pictures. Taxonomic identification was assisted by specialised plant anatomy bibliography (Crivellano and Schweingruber, 2013; Gale and Cutler, 2000; Schweingruber, 1990) and reference to the collection of modern charred woods and seeds held at the University of Valencia's Laboratory of Archaeology. The frequency of identified wood taxa was assessed as percentages of fragment counts (Chabal, 1988; Chabal et al., 1999). The diagram was drawn using software developed by Mimaen at the Polytechnic University of Valencia. 3.2. Micromammals The micromammal remains analysed here were recovered in levels I-VIII. The results for the material retrieved during the 2004, 2007, 2009 and 2014 field seasons are laid out in a previous publication (Tormo and Guillem, 2015). This study focuses on levels VII and VIII, whence most (39% and 51.4%, respectively) come (Fig. 3). Levels I-V contain few remains, mostly postcranial fragments of difficult taxonomic assignment. Two molars belonging to Microtus cabrerae have been determined in levels II and V. In Level VI, only two unidentified micromammal remains were found d a diaphyseal fragment and a metapodial fragment. The remains were recovered by water sieving in a three-sieve stack with 1 cm, 0.5 cm and 0.05 cm meshes. After drying, the sediment was sorted with the aid of a Luxo magnifying glass. The drawing and the metric study of the teeth was performed using Adobe Photoshop CS6 image processing software and a Nikon SMZ-U stereoscopic microscope with a built-in Nikon Digital Sight camera. The measurements were taken on anatomically oriented, 25  photos of the occlusal surface of the teeth. They are expressed in millimetres and correspond to the maximum length (L) and

5

width (W), with the variability of the dimensions for each molar being also specified.
pez Martínez (1980) was The nomenclature proposed by Lo followed. In descriptions, the abbreviations used for the teeth are I, M and P, which denote incisor, molar and premolar, respectively, with uppercase and lowercase letters denoting upper teeth and lower teeth. The alterations caused by predation (following Andrews, 1990; representation of the remains, type of fractures, degree of digestion), by post-depositional processes, and by exposure to fire were assessed and noted. The following indexes were calculated:  Index of Relative Abundance (IAR, proposed by Dodson and Wexlar, 1979): Ri ¼ Ni/(MNI x Ei), where Ni is the number of each skeletal element found in the sample, MNI is the minimum number of individuals and Ei the number of each skeletal element estimated for an individual.  Ratio of postcranial (Pc) to cranial (C) elements: Pc/C index, where the postcranial elements are humeri, radii, ulnae, femora and tibiae, and the cranial elements are maxillae, mandibles and isolated molars; to correct for their relative abundance in the skeleton, the sum of the postcranial elements is multiplied by eight and that of the cranial elements by five (Andrews, 1990).  Ratio of proximal limb elements (H: humerus, F: femur) to number of cranio-maxillary (CMx) and mandibular (Md) remains: HþF/CMxþMd index.  Preferential loss of distal elements: RþT/HþF index (R: radius, T: tibia, H: humerus, F: femur).  Ratio of isolated molars (IM) to empty alveolar spaces in the
ndez-Jalvo and mandibles and maxillae (EA): IM/EA index (Ferna Andrews, 1992). The fractures found on long bones, maxillae and mandibles were sar (2010). categorised according to Benna

Fig. 3. Abrigo de la Quebrada. The micromammal assemblage: spatial provenience, Number of Remains (NR), and taxa.


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

6

4. Results 4.1. Identified flora A total of 3765 charcoal fragments from 25 spits have been analysed, leading to the identification of 12 woody taxa: Ephedra sp. (joint pine), Ericaceae (heather), Juniperus sp. (juniper), monocotyledons, Pinus nigra and/or P. sylvestris (black and/or Scots pines), Pinus tp. pinaster (maritime pine), Pinus sp., cf. Pistacia sp., evergreen Quercus sp. (holm oak, kermes oak), Quercus sp., cf. Rosmarinus officinalis (rosemary) and Viburnum sp. (Fig. 4). Both trees and shrubs are found in this list, but the latter are fewer.

Carpological remains of Celtis, of possible Fabaceae, and of other angiosperms, were also identified. The highest taxonomic diversity is found in level VIII, which yielded 11 anthacological and five carpological taxa, and the lowest in level III (which only yielded Pinus nigra/P. sylvestris and conifer remains). In general, there seems to be a positive correlation between taxonomic richness and number of fragments analysed (Fig. 5). All levels are of similar composition. Pines account for between 80 and 90% of the assemblages, sometimes more. Conversely, the presence of angiosperms is always very low (5e10%).

Fig. 4. Abrigo de la Quebrada. SEM photographs of some woody taxa. Conifers: 1. Juniperus sp., cross section 90; 2. Juniperus sp., tangential section 300; 3. Juniperus sp., radial section 900; 4. Pinus nigra and/or P. sylvestris, cross section 150; 5. Pinus nigra and/or P. sylvestris, radial section 400; 6. Pinus pinaster, radial section 900; Angiosperms: 7. Ericacea, cross section 200; 8. Ericacea, tangential section 800; 9. Pistacia sp., cross section 130; 10. Quercus sp. evergreen, cross section 130; 11. Viburnum sp., cross section 150; 12. Viburnum sp., radial section 350.


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

7

Fig. 5. Abrigo de la Quebrada. Taxonomic diversity as a function of the number of charcoal fragments analysed.

In some cases, pines could be identified to species or group of species (Pinus nigra and/or P. sylvestris, Pinus tp. pinaster), which makes it possible to reconstruct environmental conditions with more detail. Cold-climate pines abound throughout. Given the accompanying taxa and the geographical situation of the rock shelter, we assign them to Pinus nigra and/or Pinus sylvestris. Anatomically, these species are very much like other cryophilous pines (Pinus uncinata, Pinus mugo, etc.), from which they can nonetheless be distinguished based on transverse plane (secretory ducts distributed in the latewood) and radial plane (presence of windowlike cross-field pits and denticulate marginal ray tracheids) views (Schweingruber, 1990). Most pine charcoals, however, could only be identified at the genus level (Pinus sp.), due to their small size, lack of consistency of the woody tissue, or other kinds of alterations concealing the anatomical structure of the cross-fields. Some fragments of Pinus tp. pinaster have been identified, based on the presence of 2e4 pinoid pits per cross-field (Schweingruber, 1990). All come from spit 5 of level VIII. In modern times, P. nigra and P. pinaster co-occur on sandy patches of the Iberian System (Costa et al., 1997). Viburnum tinus and V. lantana, which could well be the species of the genus identified at Quebrada, are characteristic of the understorey of such forests. Juniperus is discontinuously present. In the regional Upper Palaeolithic, the several species of juniper known in Iberia, nowadays typical of distinct types of environments, have been shown to coexist within the sites' wood gathering catchments (Badal García and Martínez Varea, 2018; Martínez Varea and Badal García, 2018). However, it is only based on their reproductive organs that the genus's species can be identified in a charred plant assemblage, and no such remains were retrieved in Quebrada. Two stems with overlapping scale-like leaves could correspond to Juniperus sabina or J. phoenicea (Fig. 6-A). Most of the identified angiosperm remains come from levels VIIVIII. These include Ericaceae, a mostly heliophilous family that comprises a wide range of genera and species typical of diverse environments. Other shrubs, such as rosemary and ephedra, have also been identified. The few fragments of evergreen Quercus indicate the presence of holm oaks and kermes oaks, which can also be found in pine forests. The carpological remains show the presence of Celtis sp. (Fig. 6-B), cf. Fabaceae, and three other indeterminate angiosperms. The frequency diagram in Fig. 7 summarizes the evidence. The similarity of the assemblages is well apparent, with only the

Fig. 6. Abrigo de la Quebrada. Scale-like juniper leaves (A) and Celtis endocarp (B) recovered in.

almost complete lack of angiosperms in both the charcoal and seed remains found above Level VI being worthy of special notice. 4.2. Micromammals A total of 382 micromammal remains were found, almost all of them in levels VII (39%) and VIII (51.4%). The minimum number of individuals for these two levels is 12 and 15, respectively. Based on the study of molars, five species of rodents were identified: Arvicola sapidus, Microtus cabrerae, Terricola duodecimcostatus, Eliomys quercinus and Apodemus sylvaticus. The Arvicola M2 from Quebrada has the typical features of A. sapidus, as the enamel is thicker in the distal than the mesial part
pez García, 2008; Cuenca-Besco
s et al., 2008) of the triangles (Lo (Fig. 8, nos. 1e3). We identify M. cabrerae based on the length of the only m1 (3.5 mm) (Table 3), which falls within the values that define M. cabrerae. The m1 of the Middle Pleistocene form M. brecciensis does not exceed 3 mm in length (Laplana and Sevilla, 2013), while, in the final Middle Pleistocene site of Preresa (Moreno et al., 2017), where M. cabrerae is already present, the average length of 50 m1 is
et al., 2011). 3.10 mm (minimum, 2.81, maximum 3.51 mm; Sese Following Laplana and Sevilla (2013), this Quebrada measurements


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032

8


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

Fig. 7. Abrigo de la Quebrada. Frequency diagram of anthracological and carpological remains.

further suggest assignment to the Upper Pleistocene. These inferences are corroborated by the fact that the site's only Microtus m3 has closed triangles, while the Middle Pleistocene form M. brecciensis features confluent triangles (Fig. 8, nos. 4e11). The Terricola duodecimcostatus m1 from Quebrada has three closed triangles in front of the posterior loop, while the next two, T4 and T5, are confluent, forming the so-called pitymyan rhombus. The neck of the anterior loop is wide, due to the reduction of reentrant angles A8 and A9. T6 is open and minimally inclined and triangles T6 and T7 display a symmetrical and parallel disposition, forming a second pitymyan rhombus that is broadly confluent with the anterior loop. In the one measurable instance, the labial width is 0.42 mm and the lingual width is 0.55 mm, while the width of the neck connecting T6-T7 with the AC, taken on 2 M, is 0.42 and 0.44 mm. The L/W ratio is 3.91. All these values are within the
pez-García, 2008) (Fig. 8 nos. variability of T. duodecimcostatus (Lo 12e14). Eliomys quercinus molars have a concave occlusal surface with transverse ridges along the tooth from the labial to lingual surface and the main cusps at each end are well developed (Fig. 8 nos. 15e18). The measurements are within the range of variability of the present-day species (Table 3). In the Apodemus m1 from Quebrada, T4 and T7 are joined and T1 is hardly displaced. In the M2, T9 is well developed in the only molar preserved. Both morphological features are characteristic of A. sylvaticus (Pasquier, 1974) (Fig. 8, nos. 19e26). Only the assemblage from levels VII-VIII is amenable to paleoenvironmental interpretation. Among the taxa identified, two species (M. cabrerae and T. duodecimcostatus) have clearly Mediterranean ecological requirements. Another, A. sapidus, implies the presence of a permanent water course with a riverbank where it could dig its tunnels and featuring an herbaceous vegetation or the plants with non-woody stems upon which it feeds. In the Mediterranean region, A. sapidus is currently found in the Pyrenees and lbez, 1987). in coastal streams (Gosa

M. cabrerae requires areas with permanently damp soil allowing for seasonal pools with green vegetation to form year-round. This vole would have lived in meso- and supra-Mediterranean bioclimatic zones, at an altitude of approximately 250e1500 m, depending on latitude and climatic conditions. Today, in the Valencia Region, it is found in river sources, springs, the base of ravines, disused fields and roadside ditches or forest tracks (Blanco,
ndez-Salvador, 2007; Garrido-García and Soriguer, 1998: Ferna 2015) with a plant cover predominantly made up of grasses (Brachypodium phoenicoides) with scattered bulrushes (Scirpoides holoschoenus) and elmleaf blackberry (Rubus ulmifolius), at an altitude of 514e1082 masl d in meso-Mediterranean and supraMediterranean bioclimatic zones with a dry-subhumid climate (Belenguer et al., 2016). T. duodecimcostatus is a burrowing taxon linked to diggable soils that are neither very sandy nor very stony. It digs its tunnels in open spaces. This species is endemic to the Iberian Peninsula; elsewhere, only in south-east France has its presence been recorded. It feeds on roots, tubers, bulbs and any juicy plant elements. It is a species typical of Mediterranean environments that is not usually found above an altitude of 900e1000 m in Catalonia (Gos albez, 1987), although in Sierra Nevada it can be found as high as 3000 m asl (Cotilla and Palomo, 2007). E. quercinus is a highly generalist species that occupies a wide spectrum of terrestrial and arboreal niches. It lives close to houses, in stony areas, shrubland and forests of holm oaks, cork oaks, pines and deciduous trees. It is located from sea level up to an altitude of 1500 m. It has an omnivorous diet and is found across Iberia (Moreno, 2007). A. sylvaticus is also a highly generalist species, although it mainly occupies areas with good shrub or tree cover at the edge of forests lbez, of deciduous trees, pines and holm oaks or kermes oaks (Gosa 1987; Jubete, 2007).


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

9

Fig. 8. Abrigo de la Quebrada micromammal remains. A. sapidus: 1. Right M1, 2. Left m2, 3. Right m2. M. cabrerae: 4. Left M3, 5. Right m1, 6. Right m1, 7. Right m2, 8. Left m2, 9. Right m3. T. duodecimcostatus: 10. Left M1, 11. Right M2, 12. Left M3, 13. Right m1, 14. Left m1. E. quercinus: 15. Left P4, 16e17. Left M2, 18. Right m2. A. sylvaticus: 19. Right M1, 20. Right m1, 21. Right M1, M2 and M3, 22. Right m1 and m2, 23. Left m1, 24. Right m2, 25. Right m1, 26. Right m2.


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

10

Table 3 Abrigo de la Quebrada. Molar measurements of M. cabrerae, T. duodecimcostatus, E. quercinus and A. sylvaticus. N: Number of molars; L: Length; B: Breadth. M. cabrerae

N.

M2

1 1 2 2 1 1 3 3 N 1

M3 m1 m2 T. duodecimcostatus M1 M2 M3 m1 m2 E. quercinus P4 M2 m2 A. sylvaticus M1 M2 M3 m1 m2

1 1 1 1 1 1 1 1 N 1 1 2 2 1 1 N 2 2 1 1 1 1 5 5 3 3

Max. L B L B L B L B

2073 1123

1202 Max.

L B L B L B L B L B Max. L B L B L B L B L B L B L B L B

1562 2003

Max. 2251 1476

1958 1309 1366 1215

Med. 1,64 0,931 2004 1079 3,5 1448 1791 1129 Med. 2,18 1845 1098 1,47 0,814 3035 1009 1,64 0,82 Med. 1098 1631 1521 1942 1476 1893 Med. 2144 1385 1,23 1,23 1094 0,932 1,02 1208 1333 1188

Min.

1934

1,64 1013 Min.

Min.

1479 1,88

Min. 2036 1293

1874 1162 1298 1146

Wexlar, 1979; Guillem, 1996; Kowalski, 1990). At Quebrada, based on current ethological data, such predators most likely were small carnivores and birds of prey d both diurnal (Falconiformes) and nocturnal (Strigiformes) d that use caves and rock shelters as places of refuge and, in the case of birds, also as nesting places. Table 4 shows a summary of the results for three variables (representation of skeletal elements, fracture type, and degree of digestion) used to determine the origin of the accumulation. The low values of the representation indexes, the high level of fragmentation, and the strong degree of digestion of the remains are well apparent. According to the method proposed by Andrews (1990) d and bearing in mind that we are extrapolating from modern predator samples, that postdepositional processes can distort the record, and that the formation of micromammal aggregates is not always related to predation d these values are like those generated by small carnivorous mammals. The absence of insectivores in the sample is consistent with this inference, as they do not tend to form part of the diet of such mammals. The number of micromammal remains found is inversely correlated with the intensity of human occupation. Indeed, the basal levels (VII and VIII) yielded few lithic materials and, while the remains of micrommamals and of Leporidae accumulated by birds of prey (Villaverde et al., 2017) are numerous, its macrofaunal remains feature carnivore marks, suggesting human occupation was sporadic. The opposite is the case in Levels III-V. In the Valencia Region, this pattern is widespread. A case in point is the Neolithic site of Cova de l'Or, where the abundant presence of unaltered Chiroptera remains reflects natural death and
rez a very sporadic presence of humans and their livestock (Pe Ripoll, 2016). A Middle Paleolithic example is Cova Negra, where the presence of bat neonates implies the presence of breeding colonies, which is inconsistent with prolonged human presence (Villaverde et al., 1996). At other sites, such as Abric de la Falguera (Guillem, 2006) or Cingle del Mas Cremat (Guillem, 2010), the most numerous micromammal accumulations are found at times when human presence was low key, and the sites were in use by such carnivores as the fox (Vulpes vulpes).

5. Discussion 5.1. Taphonomic processes and intensity of human occupation 5.1.1. Origin of the micromammal accumulations Micromammal remains derive from the pellets and excrements left by the predators that consume microvertebrates (Andrews, 1983, 1990; Brodkorb, 1959; Chaline and Mein, 1979; Dodson and

5.1.2. Preservation and alteration of plant remains Most plant biomass disappears with use, e.g. as food or fuel. Even though much of the biomass disappears during combustion (preservation is only of the wood that was going through the carbonisation phase when the fire went out), the fuel's woody remains are more frequently found in archaeological deposits than the nonwoody organs (seeds, leaves, etc.) that are used or consumed raw.

Table 4 Abrigo de la Quebrada (levels VII and VIII). Taphonomic indicators of the origin of the micromammal accumulations. Pc: Postcrania; C: Crania; H: Humerus; F: Femur; Mx: Maxilla; Md: Mandible; R: Radius; T: Tibia; IM: Isolated molars (molars no inserted in the mandibles and maxillae); EA: Empty alveolar spaces in the mandibles and maxillae.

Representation

Fractures

Digestion

Variables analysed

LEVEL VII

LEVEL VIII

Relative Abundance Index Pc/C H þ F/Mx þ Md R þ T/H þ F IM/EA Complete crania-maxilla Maxilla with zygomatic missing Complete mandible Mandibles with ascending ramus missing Complete postcranial elements Complete molars Isolated molars Teeth broken (incisors and molars) Postcrania digested

14,67 0,92 3 0,08 4,4 0% 100% 0% 100% 6,30% 50% 92,90% 81,70% 75,70% ¼ Strong-Extreme 71,42% ¼ Strong-Extreme

14,25 0,56 3,3 0,3 4,7 0% 100% 0% 100% 12% 47,10% 97% 84,20% 66,30% ¼ Strong-Extreme 83,60% ¼ Strong-Extreme

Incisors digested


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

Plant matter is also particularly sensitive to biodegradation by microbial and fungal activity (Aranguren et al., 2018; Lev et al., 2005; Melamed et al., 2016; Sievers, 2006). Other postdepositional factors, e.g. trampling or exposure to the elements, also influence the size and preservation of plant remains (Chrzazvez et al., 2014). At Quebrada, differential preservation of plant remains is observed, especially when the lower (VII to IX) and the upper (III to V) levels are compared. This is due to variation in sedimentary dynamics and occupation patterns. In the lower levels, especially in level VIII, a fast sedimentation rate explains the large number of well-preserved hearths and the significant presence of charred and uncharred seeds, especially in spits 56 to 52 of level VIII and spit 46 of level VII, in which the superimposition of combustion episodes is denser. In the upper levels (especially in level IV), the archaeosedimentary evidence suggests repeated, seasonal occupations and a low sedimentation rate (Villaverde et al., 2017), leaving the archaeological remains exposed to weather, trampling, and palimpsest formation for longer strecthes of time. Seeds are most affected by these taphonomic and conservation biases. At Quebrada, microbial and fungal activity is detected throughout, but it is more visible in the upper than in the lower levels' charcoal. Judging from the humus particles present through the sequence (Fig. 9, nos. 1e2), the loss of organic matter is principally due to such activity. These humus particles are fragments with a carbonaceous appearance containing the remains of anatomically unconnected plant cells that are highly altered and

11

compacted into a vitrified, porous, shapeless mass. The meso- and microfauna of the soil (insects, bacteria and fungi) seen in Quebrada's plant macroremains (Fig. 9, nos. 3e6) is the agent that broke down the organic matter. From their presence in wood charcoal samples that provided dates consistent with the associated stone tools, we infer that these microorganisms are coeval, i.e., of Middle Palaeolithic age, not recent intrusions. One of them (Fig. 9, no. 3) is probably a ubiquist of the soil flora that has been identified at sites from different time periods and regions, across Europe (from Iberia to Poland) and in North America (Badal and Atienza, 2005; Moskal, 2010; Vidal-Matutano et al., 2015). Humus particles are most abundant in levels V and IV, whose sedimentation rate was low, allowing for longer periods of microbial activity, while the lower levels's faster accumulation would have provided plant macroremains with a better defense from microbial activity in contact with the air. This contrast could well explain the upper levels' lack of carpological remains. The ratio between the number of identified coniferous and angiosperm taxa may also relate to differential preservation. Conifers absolutely predominate over angiosperms (which appear very sporadically) in all the levels, as noted at other Mousterian
n (Zilha ~o et al., 2016) and Abric del Pastor sites, such as Cueva Anto (Vidal-Matutano et al., 2015). This predominance may reflect the composition of the biomass available in the wood-gathering territory of the sites' inhabitants, as cold conditions prevailed through most of those sequences, implying decreased environmental biodiversity. Cryophilous pines are also predominant in

Fig. 9. Abrigo de la Quebrada. Charcoal alterations. 1. Humus x60; 2. Humus x350; 3. Fungal chain x25000; 4. Bacteria x4500; 5. Microorganism x6000; 6. Microorganism x4000; 7. Rosmarinus officinalis, radial cracks on cross section x150; 8. Pinus nigra and/or P. sylvestris, vitrified cross section x400; 9. Juniperus sp., collapsed cells on cross section x1500.


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032

12


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

Mediterranean Iberia's Upper Palaeolithic sites, in which, however, the proportion of angiosperms is much higher (Badal García and Martínez Varea, 2018; Badal et al., 2012; Martínez Varea and ~o et al., 2017). A possible explanation for Badal García, 2018; Zilha this pattern is that the two types of wood deteriorate differentially, with angiosperms' being more prone to fragmentation and postdepositional alteration, as has been demonstrated in both experimental and archeological contexts (Chrzazvez et al., 2014; VidalMatutano et al., 2015). The higher frequency of angiosperm charcoal and seeds observed in the basal levels of Quebrada, in which the dynamics of the accumulation was more favorable to the preservation of plant macroremains, is consistent with this hypothesis. In general, the charcoal fragments recovered in all levels of Abrigo de la Quebrada are small, on average ca. 2 mm, which hinders identification and explains why many fragments could not be classified beyond the conifer, angiosperm, or family level. Other contributing factors are charcoal anatomy alterations that, probably, result from processes that take place during carbonization d for instance, the variably visible but frequent radial contraction cracks illustrated in Fig. 9 (no. 7). Age is another factor, one that seems to underpin the “fusion-compaction” (or vitrification) of cellular tissues (Fig. 9, no. 8) caused by the intense combustion
ry-Parisot, 2001), probably exacerbated by the process itself (The levels’ subsequent sealing and by palimpsest formation, resulting in the formation of highly thermo-altered accumulations (Eixea et al., 2011e2012). Cellular structures sealed by mineral and organic inclusions are also frequently observed (Fig. 9, no. 9). Uncharred hackberry (Celtis sp.) seeds were also found in the Quebrada deposit. The presence of such uncharred endocarps has been noted at several Lower and Middle Palaeolithic sites: among
et al., 2015), Cova del Bolomor (Ferna
ndez others, Atapuerca (Allue Peris et al., 2000), El Salt (Vidal-Matutano et al., 2018) and Cueva ~ o (García Moreno et al., 2014), in Spain, or Douara Cave del Nin (Akazawa, 1987; Matsutani, 1987), Dimanisi (Messager et al., 2008) and Theopetra (Kotzamani, 2009; Tsartsidou et al., 2015). These seeds’ high mineral content d they are basically composed of aragonite (Jahren et al., 1998; Messager et al., 2010; Shillito et al., 2009; Wang et al., 1997), one of the most frequent biogenic minerals (Weiner and Dove, 2003) d means that the uncharred Celtis endocarps are amenable to preservation in the fossil record. However, such remains have never been documented anthracologically, and none was ever directly dated. Whether they are coeval with the Paleolithic deposits they were found in must therefore remain an open issue, at Quebrada as much as elsewhere. 5.2. Plant resource management Most Quebrada's combustion structures appear as superimposed carbonaceous extensions resulting from the scattering and alteration of hearths successively lit in broadly the same spot, one that appears to have been the most appropriate to function as the hub of the occupation, around which activities were carried out and objects were discarded. In some cases, this is demonstrated by the reddening of the underlying sediment, with their shallowness and lack of a prepared perimeter explaining the easy scatter of the charcoal, caused by repeated use and attendant trampling (Eixea et al., 2011e2012). This redundancy d otherwise apparent in the abundance of fire-altered bones and flints d shows that the site's occupants were efficient fire managers, fully capable of keeping it alight, of controlling it, and of using it for a diverse range of purposes. This conclusion is consistent with the evidence that Middle and early Upper Pleistocene Europeans mastered pyrotechnology to the extent necessary to make fire (Sorensen et al., 2018), to use it for the modification of the physical properties of the wood

(Aranguren et al., 2018), or to manufacture such artificial rawmaterials as birch bark pitch (Schenck and Groom, 2018). The composition of the charcoal assemblages produced by the above processes allows several, non-exclusive hypotheses to be put forth: (a) the local vegetation underwent little, if any structural change throughout; (b) coniferous trees (primarily pines) were the preferred fuel; and (c) the preservation of angiosperm wood varied significantly. Assessing these hypotheses requires detailed knowledge of the kinds of plant formations extant in the region during the Mousterian. At present, however, such detailed knowledge is unavailable; the information at hand is insufficient to draw detailed maps of the period's vegetation, even though our record is clearly partial and incomplete. Indeed, the plant resources exploited by Middle Paleolithic people must have been rather more diverse than what we can tell from what is preserved in archaeological deposits (Vidal-Matutano et al., 2015). With regards to Quebrada, our hypothesis is that the identified taxa do reflect their abundance in the shelter's immediate setting, which would have been a pine forest. However, the range of species used was probably wider, as must also have been both the range of plant organs and the purposes to which plant resources were procured. It is well known, for instance, that wooden tools existed in the Lower and Middle Paleolithic, as recently highlighted by the Italian material reported in Aranguren et al. (2018); thus, it cannot be excluded that some of the charcoal comes from the burning of wooden tools. It is also unlikely that plants were not eaten. In this regard, it should be noted that Celtis sp. drupes are edible. If indeed coeval, those found at Quebrada must have been harvested as food, at the end of summer/beginning of autumn, which is when the fruit of the Mediterranean hackberry tree ripens d thereby also providing valuable information on the seasonality of the site's human occupation. This information is consistent with other indicators (faunal dentitions, presence of fruit phytoliths) suggesting occupation outside the winter period (Villaverde et al., 2017; Esteban et al., 2017). The zooarcheological and raw-material procurement data suggest that, in a context of significant territorial mobility, Quebrada was used to exploit the animal species available in the immediate surroundings d ibex, horse and deer being the principal targets (Eixea et al., 2011e2012, Sanchis et al., 2013; Villaverde et al., 2017; Real et al., 2018). Most flint is also local (Eixea et al., 2011), coming from within an area of 5e8 km, although much of it would have been accessible within a smaller radius of approximately 2 km. The documented woody taxa would also have been found within a similar radius, mostly in the surrounding plateaus rather down in the adjacent walls of the Barranco de Ahillas gorge. The composition of Quebrada's paleobotanical remains is consistent with these inferences. The plant resources documented were mostly local, as is the rule in prehistoric societies, especially when collected as fuel (Chabal, 1997). Given the economic and social importance of fire d not only for cooking, the processing of adhesives or the fending off predators, but also to provide heat, light and a focus for sharing and other forms of personal interaction d wood gathering must have been a recurrent, planned, daily activity in which almost all members of the group participated. Dead wood, which does not require drying and, in natural forests, is plentiful, must have made-up most of the collected fuel. Even though likely, this inference, however, cannot be verified with the Quebrada charcoals; xylophage galleries have been identified, but they are not numerous, which means that whether they were excavated in the living tree or after collection cannot be established
ry-Parisot, 2001). (The The pine forests around Quebrada would have harbored a range of angiosperm plants more diverse than reflected in the site's


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

anthracological record. This under-representation may be due to preservation issues, but it is equally plausible that angiosperms would have been only occasionaly collected, or seldom brought home, as implied by the fact that they are also scarcely represented among phytoliths (Esteban et al., 2017) d almost 80% of which are of herbaceous plants. The introduction of the latter probably responded to different needs, namely bedding, food, or the lighting of fires. These wood-gathering patterns seem to be constant through the Quebrada sequence. Despite the contrasts in type of occupation seen when the upper levels are compared with the basal ones, no significant difference between them exists with regards to plant exploitation patterns. Since cryophilous pines provided most of the wood in all levels and none contains a clearly warm-climate flora, we can thus infer that (a) the site was occupied during stadial periods, and (b) during the particular stadials at stake, local plant communities remained stable and/or were exploited by humans in a manner that remained stable d just as is the case with the local fauna. 5.3. Paleoenvironmental reconstruction Marine cores in the Alboran Sea and the Atlantic Ocean document various abrupt climatic fluctuations during the Last Glacial period, with alternating cold (Heinrich Events) and warm events (Dansgaard/Oeschger) (Cacho et al., 1999; Lebreiro et al., 1996). The continental pollen data corroborate that these climatic changes affected the plant cover of Iberia (Fletcher et al., 2010; GonzalezSamperiz et al., 2010; Moreno et al., 2012), and so do the charcoal records available for the Middle Palaeolithic of the peninsula. Most such records coincide in revealing a stadial, cold-climate vegetation; only a few archaeobotanical assemblages record a warmclimate flora. Causes for this scarcity may well be reside in that, during warm periods, either humans did not use the areas containing the caves and rock shelters that yielded the available evidence, or hiatuses in sedimentation, and attendant palimpsest formation, have hindered the formation therein of adequate anthracological archives. At Quebrada, the high frequency of cryophilous pines suggests that they were readily available in the surrounding area. The available pollen sequences are consistent with a general predominance of conifer formations in the region (although, due to the nature of the material, a wider range of species is represented therein). At Salines, Villena and Cova Beneito, for instance, we see very high percentages of pine and juniper among the woody taxa, although the vegetation is dominated by herbaceous plants such as
n et al., 2012). In the present day, cryophilous pines Poaceae (Carrio in the Iberian Range grow above altitudes of 1500 m and extend to the upper tree line (Costa et al., 1997). Together with these pines, Juniperus phoenicea and J. sabina can give rise to open forests above 1400 m, under markedly continental climatic conditions. Based on this evidence, the area around Quebrada would have ́ been under dry or sub-humid supra-Mediterranean environmental ́ conditions through the time of human occupation. From the scarcity of shrubs and small trees an open pine parkland growing on poor soils or with little edaphic development, typical of harsh climatic periods, interspersed with patches of steppe, may be inferred. That almost 80% of phytoliths are from herbaceous plants (Esteban et al., 2017) supports the existence of open forests, consistent with the presence of such micromammals as M. cabrerae and T. duodecimcostatus which would have required biomes with permanent herbaceous plants and open spaces (these taxa are eurythermal and can also live under cold climatic conditions). Apodemus sp. and Eliomys quercinus would have lived in the more densely forested parts of the landscape, where junipers, holm oaks,

13

kermes oaks and other woody taxa also thrived. The lower strata of the vegetation would have consisted of different species of heather, probably in the form of bushes. Indeed, a notable presence of some Ericaceae, which are present in the Quebrada charcoal assemblages, is seen in regional pollen sequences. The presence of A. sapidus points to a wet environment featuring permanent water courses, the banks of which would have formed the species’ ideal habitat. Based on the cryophilous taxa and the other bioindicators found in the Mousterian sites of Mediterranean Iberia, a climatic boundary running at approximately latitude 40ºN is readily apparent. To the south, the wood charcoal from sites located between sea level and 700e800 m asl is dominated, during MIS-4 and MIS-3, by remains of conifers, mostly cryophilous pines (Pinus sylvestris-nigra) and juniper, with a few angiosperms (Badal et al., 2012; VidalMatutano, 2017; Vidal Matutano et al., 2015, 2017). The low diversity of shrubs and small plants may indicate the presence of open herbaceous steppe formations, in line with the pollen record (Gonzalez-Samperiz et al., 2010). These conditions are well apparent in Mediterranean Iberia's eastern and southern sites (Aparicio, 1981; Badal, 1984, Carrion et al., 2008). Pines are excellent climatic bioindicators. At present, Pinus sylvestris and P. nigra grow above 1000e1200 m asl on the mountains of Iberia under supra-Mediterranean (mean annual temperature 8e13
C) or oro-Mediterranean (mean annual temperature 4e8
C) conditions (Rivas-Martínez, 1987). P. sylvestris is more cryophilous than P. nigra, and requires more humid conditions; for these reasons, it can be found on the highest and most humid mountains of the Peninsula. These taxa would have been well-adapted to the dry, cold climate that prevailed in Iberia during the Last Glacial period because pines are capable of storing considerable amounts of water in their trunks, which helps them survive through long periods of drought; their presence pines in archaeological sites from MIS-5a, MIS-4 and MIS-3 indicates a Mediterranean-type precipitation regime defined by a summer water deficit. When we compare the present-day distribution of Pinus nigra and P. sylvestris in the Iberian Peninsula with the location of Mousterian sites, it becomes apparent that, during the colder oscillations of the Upper Pleistocene, the sequence of bioclimatic belts would have been displaced by up to 1000 m in altitude, implying up to 10
C decrease in temperature. At such times, most of southern Iberia would have been characterised by oro-supra-Mediterranean bioclimatic conditions. Warmer conditions would have persisted, ́ ́ however, in a narrow band extending along the coastal areas of Andalusia and Portugal, where thermophilous species thrived ́ throughout d cf. the wood charcoal and cones of P. pinea identified across the Gorham's Cave sequence (Metcalfe, 1958; Carrion et al., 2008), or the Olea europaea seed identified and radiocarbon dated to 37,220 ± 290 BP (OxA-12272) at Higueral de la Valleja (Carrion et al., 2008; Jennings et al., 2009). 5.4. Chronostratigraphic implications Dry, hot conditions during the first half of MIS-5a, coinciding with the long Greenland Interstadial (GI) 21, between 78 and 85 ka,
n by the predomiare denoted in sub-complex AS5 of Cueva Anto nance of Pinus halepensis and Juniperus sp. in association with Ephedra sp. (Zilh~ ao et al., 2016). This combination suggests an open steppe landscape with scattered pine stands. Due to its sedimentary characteristics, sterile level VI of Quebrada may well have been deposited during a warm episode, conceivably the same repre
n; the available OSL dates sented by sub-complex AS5 of Anto (80.0 ± 4.7 and 83.2 ± 5.4 ka; pooled mean 81.4 ± 2.5 ka; Table 2) are consistent with this hypothesis. The radiocarbon dates for levels V-III of Quebrada are consistent with deposition during an interval straddling the end of MIS-5a,


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032

14


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio

MIS-4 and MIS-3. However, in these levels, Testudo hermanni is a significant presence, one that would seem inconsistent with a postMIS-5a age because, in Mediterranean Iberia, this taxon is no longer found in faunal lists from sites that post-date the Last Interglacial. This evidence pleads for levels V-III of Quebrada to be correlated
n, which accumulated with sub-complexes AS3-AS2 of Cueva Anto during the progressively cooler oscillations marking the end of MIS5a, between 78 and 71 ka. Indeed, in the Greenland ice record, stadial conditions prevail during the intervals 77.8-76.4 ka (Greenland Stadial (GS) 21.1) and 74.1-72.3 ka (GS-20) (Rasmussen et al., 2014), while replacement of Pinus halepensis with P. nigra correspondingly characterises the vegetation cover reflected in the
n. A charcoal and pollen from sub-complexes AS3 and AS2 of Anto similar pattern is seen at Abric del Pastor, which dates beyond 75 ka and where the identified flora is dominated by steppic plants (Juniperus) and features 10e20% of Pinus nigra. In central Iberia, cryophilous pines (Pinus sylvestris), in association with Betula, have also been identified in last interglacial deposits, albeit situated at a higher altitude, e.g. at the site of Cueva del Camino (1114 masl; Arsuaga et al., 2010; Gonzalez-Samperiz et al., 2010; Badal et al., 2011). Thus, coupled with the tortoise evidence and the lack of significant discontinuity in sedimentary accumulation, the compari
n and Pastor records supports placing Quebrada son with the Anto levels V-III at the end of MIS-5a. This suggestion needs to be additionally supported by chronometric dating but, if confirmed, implies that the site's current radiocarbon dating record is entirely made-up of vastly underestimated minimum ages. 6. Conclusions The evidence from the anthropogenic (plant macroremains) and natural (microfauna) inputs provides enough data for a reconstruction of the environmental context of the short, repeated human occupations recorded in the Abrigo de la Quebrada sequence. The anthracological results show not only that pine wood was the preferred fuel during the accumulation of levels VII-VIII and V-III but also that angiosperms were present in the pine forest developed under supra-Mediterranean conditions extant in the site's immediate environment d which, as indicated by the evidence from both microfauna and phytoliths, included humid areas and open spaces with permanent herbaceous plants. This landscape provided all the resources required by the site's inhabitants, including lithic raw-materials and meat, which were also locally ́ ́ acquired. The flora represented in the wood charcoal assemblage matches that from other Mousterian sites of Mediterranean Iberian that are securely dated to stadial periods. At such times, Pinus sylvestris woodlands would have been the most extensive formation to the north of latitude 40ºN, while to the south P. sylvestris and P. nigra stands would have co-existed. Thermophilous and xeric bioindicators such as Pinus halepensis, Pinus pinea, Ephedra, Juniperus, Fabaceae and Olea europaea would have found refuge in the southernmost parts of the peninsula only. Conversely, during interstadial periods, Pinus halepensis extended northwards and the cryophilous pines retreated to higher elevations. This process is
n (Zilha ~o well illustrated by the MIS-5a sequence of Cueva Anto et al., 2016), where the xeric aspect of the flora recorded in subcomplex AS5, dated to GI-21, suggests that, during the Last Interglacial, south-east Iberia was already one of the driest parts of the Peninsula. The plant remains from Abrigo de la Quebrada have also made it possible to carry out a more in-depth analysis of matters related to preservation issues. The differences found in this regard between the upper (III and IV) and basal (VII to IX) levels are determined by

taphonomic factors d the dynamics of the sedimentary accumulation and of the site's human occupation, i.e., the lower sedimentation rate and the higher intensity of the activities recorded in the upper levels versus the higher sedimentation rate and more sporadic nature of the human presence recorded in the basal levels. In both cases, however, occupations were short and seasonal. Regardless of whether they date to glacial, interglacial, stadial or interstadial periods, all the Mousterian sites of Mediterranean Iberia have yielded plentiful evidence of a controlled use of fire. Quebrada adds to this evidence, and further contributes to show that, during stadials, the habitat of regional Neanderthal groups extended onto mountainous inland areas, as documented also at El Salt (at 680 masl) (Vidal-Matutano et al., 2018). Here, however, occupation is thought to have taken place in the autumn/winter months (Vidal-Matutano et al., 2017), the reverse of Quebrada. This contrast hints at settlement systems characterized by significant year-round mobility over extensive territories. Acknowledgements Archaeological research at Abrigo de la Quebrada was funded by
n (HAR2011-24878; Spanish Ministerio de Ciencia e Innovacio CGL2012-34717; HAR2014-5267º1-P; HAR2017-85153/P) and by Generalitat Valenciana (Grant PROMETEOII/2013/016; PROMETEO/
n Marco is beneficiary of a Ramo
n y Cajal 2017/060). Y. Carrio research fellowship funded by the Spanish Ministerio de Economía, Industria y Competitividad (Grant RYC-2014-16027). C.M. Martí
de Talent” of VLC-Campus nez-Varea is beneficiary of an “Atraccio ncia (Grant UV-INV-PREpredoctoral grant of Universitat de Vale DOC15-265753). AE is funded by the APOSTD Postdoctoral (Grant APOST2018/121) by Generalitat Valenciana). References Akazawa, T., 1987. The ecology of the middle paleolithic occupation at Douara cave, Syria. In: Bull. of Tokyo University Mus, vol. 29, pp. 155e166.
, E., Ca
ceres, I., Expo
sito, I., Canals, A., Rodríguez, A., Rosell, J., Bermúdez De Allue Castro, J.M., Carbonell, E., 2015. Celtis remains from the lower Pleistocene of gran dolina, Atapuerca (burgos, Spain). J. Archaeol. Sci. 53, 570e577.
, E., 2002. Din

n y explotacio
n del combustible leneoso Allue amica de la vegetacio durante el Pleistoceno Superior y el Holoceno del noreste de la Península
rica a partir del an

gico. Ph.D. Thesis. Universitat Rovira i Ibe alisis antracolo Virgili, Tarragona. Andersen, K.K., Azuma, N., Barnola, J.-M., Bigler, M., Biscaye, P., Caillon, N., Chappellaz, J., Clausen, H.B., Dahl-Jensen, D., Fischer, H., Flückiger, J., Fritzsche, D., Fujii, Y., Goto-Azuma, K., Grønvold, K., Gundestrup, N.S., Hansson, M., Huber, C., Hvidberg, C.S., Johnsen, S.J., Jonsell, U., Jouzel, J., Kipfstuhl, S., Landais, A., Leuenberger, M., Lorrain, R., Masson-Delmotte, V., Miller, H., Motoyama, H., Narita, H., Popp, T., Rasmussen, S.O., Raynaud, D., Rothlisberger, R., Ruth, U., Samyn, D., Schwander, J., Shoji, H., Siggard€ rnsdo
ttir, A.E., Svensson, A., Andersen, M.-L., Steffensen, J.P., Stocker, T., Sveinbjo Takata, M., Tison, J.-L., Thorsteinsson, Th, Watanabe, O., Wilhelms, F., White, J.W.C., 2004. High-resolution record of Northern Hemisphere climate extending into the last interglacial period. Nature 431, 147e151. Andrews, P., 1983. Small mammal faunal diversity at olduvai gorge, Tanzania. In: Animals and archaeology, vol. 1, pp. 77e85. Hunters and their Prey 163. Andrews, P., 1990. Owls, Caves and Fossils. The University of Chicago Press. Aparicio, J., 1981. Primeras dataciones C.14 para el musteriense valenciano. Arch. Prehist. Levantina 16, 2e38. Aranguren, B., Revedin, A., Amico, N., Cavulli, F., Giachi, G., Grimaldi, S., Macchioni, N., Santaniello, F., 2018. Early Neanderthal wooden artifacts from Italy. In: Proc. Natl. Acad. Sci. U. S. A. Feb 2018, p. 201716068. https://doi.org/ 10.1073/pnas.1716068115.
rez-Gonza
lez, A., Sala, M.T., García, N., Alvarez-Lao
, D., Arsuaga, J.L., Baquedano, E., Pe Laplana, C., Huguet, R., Sevilla, P., Maldonado, E., Blain, H.-A., Quam, R., Ruiz Zapata, M.B., Salas, P., Gil, M.J., Uzquiano, P., Pantoja, A., 2010. El yacimiento
gico del Pleistoceno Superior de la Cueva del Camino en el arqueopaleontolo
gica, Nº. 13, Calvero de la Higuera (Pinilla del Valle, Madrid). In: Zona arqueolo
n de científicos sobre cubiles de hiena (y otros grandes Actas de la 1a Reunio
gicos de la Península Ibe
rica, carnívoros) en los yacimientos arqueolo pp. 422e442.
n al estudio de la vegetacio
n prehisto
rica del sur de Badal, E., 1984. Contribucio
s del ana
lisis antracolo
gico. Master Thesis. Valencia y norte de Alicante a trave Universidad de Valencia.


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio
lisis microsco
pico de coprolitos de herbívoros halBadal, E., Atienza, V., 2005. Ana
gicos. In: VI Congreso Iberico de Arqueometria, lados en contextos arqueolo pp. 283e294. ~o, J., 2011. The fire of Iberian Neanderthals. Wood Badal, E., Villaverde, V., Zilha charcoal from three new Mousterian sites in the Iberian Peninsula. In: Badal, E.,
n, Y., Grau, H., Macías, M., Ntinou, M. (Eds.), 5th International Meeting of Carrio Charcoal Analysis. The Charcoal as Cultural and Biological Heritage, Sagvntvm ncia, pp. 77e78. Extra 11. Universitat de Vale Badal, E., Villaverde, V., Zilh~ ao, J., 2012. Middle Palaeolithic wood charcoal from three sites in South and West Iberian: biogeographic implications. In: Badal, E.,
n, Y., Macías, M., Ntinou, M., coords (Eds.), Wood and Charcoal. Evidence Carrio ncia, for Human and Natural History, Sagvntvm Extra-13. Universitat de Vale pp. 13e24. Badal García, E., Martínez Varea, C. Ma, 2018. Different parts of the same plants. Charcoals and seeds from Cova de les Cendres (Alicante, Spain). Quat. Int. 463 (B), 391e400.
n, Y., Figueiral, I., Rodríguez-Ariza, M, O., 2012. Pinares y Badal García, E., Carrio enebrales. El paisaje del Solutrense en Iberia. In: Espacio Tiempo y Forma. Serie
poca 5, pp. 263e276. I, Prehistoria y Arqueología, Nueva e
pez-Alabau, A., Guillem, P.M., Barona, J., Belda, A. y, Belenguer, R., Monsalve, M.A., Lo
pez-Iborra, G., 2016. Nuevas aportaciones al conocimiento de la distribucio
n Lo del topillo de Cabrera Iberomys cabrerae (Thomas, 1906) en el Levante peninsular. GALEMYS 28. https://doi.org/10.7325/Galemys.2016.N1. Benn asar, M., 2010. Tafonomía de micromamíferos del Pleistoceno Inferior de la Sierra de Atapuerca (Burgos): La Sima del Elefante y la Gran Dolina. PhD Thesis. Universitat Rovira i Virgili. ~ a II. Editorial Planeta, Barcelona. Blanco, J.C., 1998. Mamíferos de Espan ́ ́ IV of bolomor cave Blasco, R., 2008. Human consumption of tortoises at level (Valencia, Spain). J. Archaeol. Sci. 35, 2839e2848. Brodkorb, P., 1959. Pleistocene birds from new providence island Bahamas. In: Bulletin of the Florida State Museum. Biological Sci. Ser, 4, pp. 349e371. Cacho, I., Grimalt, J.O., Pelejero, C., Canals, M., Sierro, F.J., Flores, J.A., Shackleton, N.J., 1999. Dansgaard-oeschger and heinrich event imprints in Alboran Sea temperatures. Paleoceanography 14, 698e705. Cacho, I., Shackleton, N., Elderfield, H., Sierro, F.J., Grimalt, J.O., 2006. Glacial rapid variability in deep-water temperature and d18O from the Western Mediterranean Sea. Quat. Sci. Rev. 25, 3294e3311.
n, J.S., (Coord), Fern

lez-Sampe
riz, P., Lo
pez-Merino, L., Carrio andez, S., Gonza ~ a, L., Burjachs, F., Lo
pez-Sa
ez, J.A., García-Anto
n, M., Carrio
n Marco, Y., Pen
n, E., Allue
, E., Badal, E., Dupre
, M., Fierro, E., Uzquiano, P., Postigo, J.M., Barro
nez Moreno, G., Gil Munuera, M., Rubiales, J.M., García Amorena, I., Jime Romera, G., Leroy, S., García-Martínez, M.S., Montoya, E., Fletcher, W., Yll, E., ~ alba, C., Gil García, M.J., Pe
rez Vieira, M., Rodríguez-Ariza, M.O., Anderson, S., Pen
mez Manzaneque, F., Sanz, A., Albert, R.M., Díez, M.J., Morales-Molino, C., Go Parra, I., Ruiz Zapata, B., Riera, S., Zapata, L., Ejarque, A., Vegas, T., Rull, V.,
rez Díaz, S., Scott, L., Abel Schaad, D., Andrade, A., Manzano, S., Navarro, C., Pe
ndez-Mateo, L., Sa
nchez Baena, J.J., Riquelme, J.A., Iglesias, R., Moreno, E., Herna
nez Espejo, F., ValleFranco, F., Chaín, C., Figueiral, I., Grau, E., Matos, M., Jime
ndez, M., Rivas-Carballo, R., Arribas, A., Garrido, G., Mun ~ iz, F., Herna
rez Jorda
, G., Miras, Y., 2012. Paleoflora y Finlayson, G., Finlayson, C., Ruiz, M., Pe
n de la Península Ibe
rica e Islas Baleares: Plioceno-Cuaternario. Paleovegetacio Ministerio de Economía y Competitividad, Madrid. http://paleofloraiberica.net/ INICIO.html.
n, J.S., Finlayson, C., Ferna
ndez, S., Finlayson, G., Allue
, E., Lo
pez-Sa
ez, J.A., Carrio
pez- García, P., Gil-Romera, G., Bailey, G., Gonza
lez-Sampe
riz, P., 2008. Lo A coastal reservoir of biodiversity for Late Pleistocene human populations: palaeoecological investigations in Gorham's Cave (Gibraltar) in the context of Iberian Peninsula. Quat. Sci. Rev. 27, 2118e2135.
lever les charbons de bois pour la pe
riode Chabal, L., 1988. Pourquoi et comment pre
thodes utilise
es sur le site de Lattes (He
rault). In: Lattara, 1, antique: les me pp. 187e222. ^ts et socie
te
s en Languedoc (Ne
olithique final, Antiquite
tarChabal, L., 1997. Fore
thode et pale
oe
cologie. Documents d'Arche
ologie dive), L’anthracologie, me Française, Paris. Chabal, L., Fabre, L., Terral, J.-F., Thery-Parisot, I., 1999. L'Anthracologie. In: re, A. (Ed.), La Botanique. Errance, París, pp. 43e104 (Dir.). Ferdie
volution. Doin, Paris. Chaline, J., Mein, P., 1979. Les rongeurs et l’e
ry-Parisot, I., Fiorucci, G., Terral, J.F., Thibaut, B., 2014. Impact of Chrzazvez, J., The post-depositional processes on charcoal fragmentation and archaeobotanical implications: experimental approach combining charcoal analysis and biomechanics. J. Archaeol. Sci. 44, 30e42. Conard, N.J., Malina, M., Münzel, S.C., 2009. New flutes document the earliest musical tradition in southwestern Germany. Nature 460, 737e740.
ricos. Una Interpretacio
n GeoCosta, M., Morla, C., Sainz, H., 1997. Los Bosques Ibe
nica. Planeta, Barcelona. bota Cotilla, I., Palomo, L.J., 2007. Microtus duodecimcostatus (de Selys longchamps, 1839). In: Palomo, L.J., Gisbert, J., Blanco, J.C. (Eds.), Atlas y libro rojo de los ~ a, Ministerio de Medio Ambiente. Tragsa, Madrid, mamíferos terrestres de Espan pp. 422e425. Crivellano, A., Schweingruber, F.H., 2013. Atlas of Wood, Bark and Pith Anatomy of Eastern Mediterranean Trees and Shrubs with a Special Focus on Cyprus. Springer, Heidelberg.
s, G., Straus, L.G., Gonza
lez Morales, M. y, García Pimienta, J.C., 2008. Cuenca-Besco ~ os mamíferos del final del Cuaternario en Cantabria: La Cueva del Los pequen
n (Ramales de la Victoria). Rev. Esp. Palaontol. 23 (1), 91e126. Miro

15

d'Errico, F., Henshilwood, C., Lawson, G., Vanhaeren, M., Soressi, M., Bresson, F., Tillier, A.-M., Maureille, B., Nowell, A., Backwell, L., Lakarra, J.A., Julien, M., 2003. The search for the origin of symbolism, music and language: a multidisciplinary endeavour. J. World Prehist 17, 1e70. Delagnes, A., Rendu, W., 2011. Shifts in Neandertal mobility, technology and s ubsistence strategies in western France. J. Archaeol. Sci. 38, 1771e1783. Desprat, S., Combourieu-Nebout, N., Essallami, L., Sicre, M.A., Dormoy, I., Peyron, O., Siani, G., Bout Roumazeilles, V., Turon, J.L., 2013. Deglacial and Holocene vegetation and climatic changes in the southern Central Mediterranean from a direct landesea correlation. Clim. Past 9, 767e787. Dibble, H.L., Abodolahzadeh, A., Aldeias, V., Goldberg, P., McPherron, S.P., Sandgathe, D.M., 2017. How did hominins adapt to ice age Europe without fire? Curr. Anthropol. 58, 278e287. Dibble, H.L., Sandgathe, D., Goldberg, P., McPherrron, S., Aldeias, V., 2018. Were western european neandertals able to make fire? J. Paleol. Archaeol. 1, 54e79. Dodson, P., Wexlar, D., 1979. Taphonomic investigation of owls pellets. Paleobiology 5, 275e284. ~o, J., 2011. Aproximacio
n al aprovisionamiento de Eixea, A., Villaverde, V., Zilha materias primas líticas en el yacimiento del Paleolítico medio del Abrigo de la Quebrada (Chelva, Valencia). Trab. Prehist. 68 (1), 65e78. ~o, J., Sanchis Serra, A., Morales Pe
rez, J.V., Real Eixea, A., Villaverde, V., Zilha  Zapata, M.M., 2011-2012. El nivel IV del abrigo de la Margalef, C., Bergada
lisis microespacial y valoracio
n del uso del Quebrada (Chelva, Valencia): ana espacio en los yacimientos del Paleolítico Medio valenciano. Mainake 33, 127e158.
n, C., Zilha ~o, J., 2014. Middle Palaeolithic flint proEixea, A., Villaverde, V., Rolda curement in Central Mediterranean Iberia: implications for human mobility. J. Lithic Stud. 1, 103e115. ~o, J., 2016. Not only flint: Levallois on quartzite and Eixea, A., Villaverde, V., Zilha limestone at abrigo de la Quebrada (Valencia, Spain): implications for neandertal behavior. J. Anthropol. Res. 72 (1), 24e57. ~o, J., Villaverde, V., 2017. Neanderthal use of Esteban, I., Albert, R.M., Eixea, A., Zilha plants and past vegetation reconstruction at the Middle Paleolithic site of Abrigo de la Quebrada (Chelva, Valencia, Spain). Archaeol. Anthropol. Sci. 9, 265e278.
blot-Augustins, J., 1993. Mobility strategies in the late Middle Palaeolithic of Fe central Europe and western Europe: elements of stability and variability. J. Anthropol. Archaeol. 12, 211e265.
blot-Augustins, J., 1997. La circulation des matie res premie res au Pale
olithique Fe ge). (E.R.A.U.L., Lie
ndez Peris, J., Guillem Calatayud, P., Martínez Valle, R., 2000. Cova del Bolomor Ferna (Tavernes de la Valldigna-Valencia). In: Datos cronoestratigr
aficos y culturales
rica, Actas de una secuenciadel Pleistoceno medio, Paleolítico da Península Ibe do 3º Congresso de Arqueologia Peninsular, vol. II. ADECAP, Porto, pp. 81e94.
ndez-Jalvo, Y., Andrews, P., 1992. Small mammal taphonomy of gran dolina, Ferna Atapuerca (burgos), Spain. J. Archaeol. Sci. 19, 407e428.
ndez-Salvador, R., 2007. Microtus cabrerae thomas, 1906. Topillo de Cabrera. Ferna In: Palomo, L.J., Gisbert, J., Blanco, J.C. (Eds.), Atlas y Libro Rojo de los mamíferos ~ a, D.G.C. .N.A.-SECEM-SECEMU, Madrid, pp. 429e433. terrestres de Espan
nez, F., Carrio
n, J., Finlayson, G., Giles, F., Rodríguez, J., Finlayson, C., Fa, D.A., Jime Stringer, C., Martínez, F., 2008. Gorham's Cave, Gibraltar. The persistence of a Neanderthal population. Quat. Int. 181, 64e71.
nchez Gon ~ i, M.F., 2008. Orbital- and sub-orbital-scale climate imFletcher, W.J., Sa pacts on vegetation of the western Mediterranean basin over the last 48,000 yr. Quat. Res. 70 (3), 451e464.
nchez-Gonei, M.F., Allen, J.R.M., Cheddadi, R., Combourieu Fletcher, W.J., Sa Nebout, N., Huntley, B., Lawson, I., Londeix, L., Magri, D., Margari, V., Müller, U.C., Naughton, F., Novenko, E., Roucoux, K., Tzedakis, P.C., 2010. Millennial-scale variability during the last glacial in vegetation records from Europe. Quat. Sci. Rev. 29, 2839e2864.
, P., Vallverdú, J., 2014. From small bone Gabucio, M.J., C
aceres, I., Rosell, J., Saladie fragments to Neanderthal activity areas: the case of Level O of the Abric Romaní (Capellades, Barcelona, Spain). Quat. Int. 330, 36e51. Gale, R., Cutler, D., 2000. Plants in Archaeology. Identification Manual of Vegetative Plants Materials Used in Europe and Southern Mediterranean to C. 1500. Westbury and Royal botanic Gardens, Kew.
n, B., Hern
Galva andez, C.M., Mallol, C., Mercier, N., Sistiaga, A., Soler, V., 2014. New evidence of early neanderthal disappearance in the iberian peninsula. J. Hum. Evol. 75, 16e27. García Moreno, A., Ortíz, J.E., Rios Garaizar, J., De Torres, T., Marín Arroyo, A.B.,
pez-Do
riga, I., 2014. La secuencia musteriense de la Cueva del Nin ~ o (Aýna, Lo
rica. Albacete) y el poblamiento neandertal en el sureste de la Península Ibe Trab. Prehist. 71 (2), 221e241. Garrido-García, J.A., Soriguer, R., 2015. Topillo de Cabrera Iberomys cabrerae ~ a, SECEM (Thomas, 1906). In: Guía de indicios de los mamíferos de Espan (accessed 20 july 2015). http://www.secem.es/wp-content/uploads/2015/07/ 020.-Iberomys-cabrerae.pdf. Gaudzinski, S., 2006. Monospecific or species-dominated faunal assemblages during the Middle paleolithic in Europe. In: Hovers, E., Kuhn, S.L. (Eds.), Transitions before the Transition. Evolution and Stability in the Middle Paleolithic and Middle Stone Age. Springer, Sante Barbara, CA, pp. 137e147.
riennes du Pe
rigord : une Geneste, J.-M., 1985. Analyse lithique d’industries mouste approche technologique du comportement des groupes humains au
olithique moyen. PhD Thesis. Universite
de Bordeaux I. Pale mes d’approvisionnement en matie res premie res au Geneste, J.M., 1988. Syste


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032

16


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio


olithique moyen et au Pale
olithique supe
rieur en Aquitaine. In: Otte, M. Pale
andertal. E.R.A.U.L., Lie ge, pp. 61e70. (Ed.), L'Homme de Ne
lez-Sampe
riz, P., Leroy, S.A.G., Carrio
n, J.S., Ferna
ndez, S., García-Anto
n, M., Gonza
s, B., Figueiral, I., 2010. Steppes, saGil-García, M.J., Uzquiano, P., Valero-Garce vannahs, forests and phytodiversity reservoirs during the Pleistocene in the Iberian Peninsula. Rev. Palaeobot. Palynol. 162, 427e457. lbez, J., 1987. Insectivors i Rosegadors de Catalunya. Metodología d'estudi i Gosa cat aleg faunístic. Ketres, Barcelona. Grayson, D.K., Delpech, F., 2003. Ungulates and the middle-to-upper paleolithic transition at grotte XVI (dordogne, France). J. Archaeol. Sci. 30, 1633e1648. Guillem, P.M., 1996. Micromamíferos cuaternarios del País Valenci a: Tafonomía,
n paleoambiental. PhD Thesis. Universitat de Bioestratigrafía y Reconstruccio ncia. Vale Guillem, P.M., 2006. Los micromamíferos (Insectivora, Rodentia y Chiroptera) del ~ o fragmento de la historia de la vida. In: Abric de la Falguera, un pequen García, O., Aura, J.E., Coords (Eds.), El Abric de la Falguera (Alcoi, Alacant). 8.000 ~ os de ocupacio
n humana en la cabecera del río Alcoi, Diputacio
n Provincial an de Alicante. Ayuntamiento de Alcoy, C.A.M., pp. 118e123 Guillem, P.M., 2010. Los micromamíferos (Rodentia, Insectivora y Chiroptera) del Cingle del Mas Cremat. In: Vizcaíno, D. (Ed.), El Cingle del Mas Cremat (Portell
n). Un asentamiento en altura con ocupaciones del Mesode Morella, Castello lítico reciente al Neolítico final, Renomar S.A. y EIN Mediterr
aneo S.L, pp. 126e145. Higham, T., Douka, K., Wood, R., Ramsey, C.B., Brock, F., Basell, L., Camps, M., Arrizabalaga, A., Baena, J., Barroso-Ruíz, C., Bergman, C., Boitard, C., Boscato, P.,
s, M., Conard, N.J., Draily, C., Froment, A., Galv
Caparro an, B., Gambassini, P., Garcia-Moreno, A., Grimaldi, S., Haesaerts, P., Holt, B., Iriarte-Chiapusso, M.-J.,
Pardo, J.F., Maíllo-Ferna
ndez, J.-M., Marom, A., Maroto, J., Jelinek, A., Jorda
ndez, M., Metz, L., Morin, E., Moroni, A., Negrino, F., Panagopoulou, E., Mene Peresani, M., Pirson, S., de la Rasilla, M., Riel-Salvatore, J., Ronchitelli, A., Santamaria, D., Semal, P., Slimak, L., Soler, J., Soler, N., Villaluenga, A., Pinhasi, R., Jacobi, R., 2014. The timing and spatiotemporal patterning of Neanderthal disappearance. Nature 512, 306e309. ~o, J., Hoffmann, D.L., Standish, C.D., García-Diez, M., Pettitt, P.B., Milton, J.A., Zilha
lez, J.J., Cantalejo-Duarte, P., Collado, H., de Balbín, R., Alcolea-Gonza ~ oz, J., Weniger, G.-Ch, Pike, A.W.G., 2018. U-Th Lorblanchet, M., Ramos-Mun dating of carbonate crusts reveals Neandertal origin of Iberian cave art. Science 359, 912e915. Hublin, J.-J., 2015. The modern human colonization of western Eurasia: when and where? Quat. Sci. Rev. 118, 194e210. Jahren, A.H., Gabel, M.L., Amundson, R., 1998. Biomineralization in seeds: developmental trends in isotopic signatures of hackberry. Paleogeogr. Paleoclimatol. Paleoecol. 138, 259e269. Jaubert, J., Verheyden, S., Genty, D., Soulier, M., Cheng, H., Blamart, D., Burlet, C., Camus, H., Delaby, S., Deldicque, D., Edwards, R.L., 2016. Early neanderthal constructions deep in bruniquel cave in southwestern France. Nature 534, 111e114. Jennings, R.P., Giles, F., Barton, R.N.E., Collcutt, S.N., Gale, R., Gleed-Owen, C.P.,
rrez-Lo
pez, J.M., Higham, T.F.G., Parker, A., Price, C., Rhodes, E., SantiagoGutie
rez, A., Schwenninger, J.L., Turner, E., 2009. New dates and palae- oenvirPe onmental evidence for the Middle to Upper Palaeolithic occupation of Higueral de la Valleja Cave, southern Spain. Quat. Sci. Rev. 28 (9e10), 830e839. Jennings, R., Finlayson, C., Fa, D., Finlayson, G., 2011. Southern Iberia as a refuge for the last Neanderthal populations. J. Biogeogr. 38, 1873e1885. €ris, O., Street, M., Terberger, Th, Weninger, B., 2011. Radiocarbon dating the Middle Jo to Upper Palaeolithic transition: the demise of the last Neanderthals and the first appearance of anatomically modern humans in Europe. In: Condemi, S., Weniger, G.C. (Eds.), Continuity and Discontinuity in the Peopling of Europe. One Hundred Fifty Years of Neanderthal Study, Vertebrate Paleobiology and Paleoanthropology. Springer, Dordrecht, pp. 239e298. Jubete, F., 2007. Apodemus sylvaticus (Linnaeus, 1758). Ficha del Libro Rojo. In: Palomo, L.J., Gisbert, J., Blanco, J.C. (Eds.), Atlas y Libro Rojo de los mamíferos ~ a, Direccio
n General para la Biodiversidad-SECEM-SECEMU, terrestres de Espan pp. 449e451. Madrid. Kotzamani, G., 2009. From Gathering to Cultivation: Archaeobotanical Research on the Early Plant Exploitation and the Beginning of Agri- Culture in Greece (Theopetra Schisto, Sidari, Dervenia). Dissertation. University of Thessaloniki, Greece (in Greek). Kowalski, K., 1990. Some problems of the taphonomy of small mammals. In: Feijfar, O., Heinrich, W.D. (Eds.), International Symposium on the Evolution, Phylogeny and Biostratigraphy of Arvicolids, pp. 285e296. Praha. Kuhn, S.L., 1995. Mousterian Lithic Technonlogy: an Ecological Perspective. Princeton University Press, Princeton, N.J.
n de la historia biogeogra
fica de Laplana, C., Sevilla, P., 2013. La documentacio
s de su registro fo
sil. Mamm Rev. 43, 1e14. Microtus cabrerae a trave Lebreiro, S.M., Moreno, J.C., McCave, I.N., Weaver, P.P.E., 1996. Evidence for heinrich layers off Portugal (tore seamount: 39N, 12W). Mar. Geol. 131, 47e56. Lev, E., Kislev, M.E., Bar-Yosef, O., 2005. Mousterian vegetal food in kebara cave, Mt. Carmel. J. Archaeol. Sci. 32, 475e484.
pez García, J.M., 2008. Evolucio
n de la diversidad taxono
mica de los microLo
rica y cambios paleoambientales durante el mamíferos en la Península Ibe Pleistoceno superior. PhD Thesis. Universitat Rovira i Virgili.
pez Martínez, N., 1980. Los micromamíferos (Rodentia, Insectivora y Chiroptera) Lo
n Achelense de Aridos I (Arganda, Madrid). In: Santonja, M., del sitio de ocupacio
pez Martínez, N., Pe
rez Go
nzalez, A. (Eds.), Ocupaciones Achelenses en el Lo

Valle del Jarama (Arganda, Madrid), Publicaciones de la Excelentísima
n de Madrid 1, pp. 161e202. Diputacio , R., Maroto, J., Vaquero, M., Arrizabalaga, A., Baena, J., Baquedano, E., Jord a, J., Julia Montes, R., Van Der Plicht, J., Rasines, P., Wood, R., 2012. Current issues in late Middle palaeolithic chronology: new assessments from northern Iberia. Quat. Int. 247, 15e25. Martínez Valle, R., 1996. Fauna del Pleistoceno superior en el País Valenciano:
micos, huella de manipulacio
n y valoracio
n paleoambiental. PhD aspectos econo ncia. Thesis. Universitat de Vale Martínez Varea, C.M., Badal García, E., 2018. Plant use at the end of the Upper Palaeolithic: archaeobotanical remains from Cova de les Cendres (TeuladaMoraira, Alicante, Spain). Veg. Hist. Archaeobotany 27 (1), 3e14. Matsutani, A., 1987. Plant remains from the 1984 excavations at Douara cave. In: Akazawa, T., Sakaguchi, Y. (Eds.), Paleolithic Site of Doura Cave, Bull. n. 21, Tokyo, pp. 117e122. McBrearty, S., Brooks, A.S., 2000. The revolution that wasn't: a new interpretation of the origin of modern human behavior. J. Hum. Evol. 39, 453e563. Melamed, Y., Kislev, M.E., Geffen, E., Lev-Yadun, S., Goren-Inbar, N., 2016. The plant component of an Acheulian diet at Gesher Benot Ya'aqov, Israel. Proc. Natl. Acad. Sci. U. S. A 113 (51), 14674e14679. Messager, E., Lordkipanidze, D., Ferring, C.R., Deniaux, B., 2008. Fossil fruit identification by SEM investigations, a tool for palaeoenvironmental reconstruction of Dmanisi site, Georgia. J. Archaeol. Sci. 35, 2715e2725. €hlich, F., Deniaux, B., Lordkipanidze, D., Voinchet, P., Messager, E., Badou, A., Fro 2010. Fruit and seed biomineralization and its effect on preservation. Archaeol. Anthropol. Sci. 2, 25e34. Metcalfe, C.R., 1958. Gorham's Cave: repport on the plant remains. Bull. Inst. Archaeol. 4, 219.
n careto. In: Palomo, L.J., Moreno, S., 2007. Eliomys quercinus linnaeus, 1766, liro Gisbert, J., Blanco, J.C. (Eds.), Atlas y Libro Rojo de los Mamíferos Terrestres de ~ a. Direccio
n General para la Biodiversidad, SECEM. SECEMU, Madrid, Espan pp. 392e394.
nchez Gon ~ i, M.F., Shackleton, N.J., Moreno, A., Cacho, I., Canals, M., Grimalt, J.O., Sa Sierro, F.J., 2005. Links between marine and atmospheric pocesses oscillating on a millenial time-scale. A multi-poxy study of the last 50,000 yr from the Alboran Sea (Western Mediterranean Sea). Quat. Sci. Rev. 24, 1623e1636.
lez-Sampe
riz, P., Morello
n, M., Valero-Garce
s, B.L., Fletcher, W.J., Moreno, A., Gonza 2012. Northern Iberian abrupt climate change dynamics during the last glacial cycle: a view from lacustrine sediments. Quat. Sci. Rev. 36, 139e153.
rez Moreno, D., Duval, M., Rubio-Jara, S., Panera, J., Bahain, J.J., Shao, Q., Pe res, C., 2017. ESR dating of Middle Pleistocene archaeoGonz
alez, A., Falgue paleontological sites from the Manzanares and Jarama river valleys (Madrid basin, Spain). Quat. Int. https://doi.org/10.1016/j.quaint.2017.09.003.
gicos Moskal, M., 2010. Los bosques holocenos en Europa Central: estudios antracolo
gicos de Polonia, Eslovaquia y Hungria. PhD Thesis. de yacimientos arqueolo ncia. Universitat de Vale
volutive d'un sousgenere de Muridae, Apodemus Pasquier, L., 1974. Dynamique e res dentaires de populations fos(Sylvaemus). Etude biometrique des caracte
de Montpellier. siles et actuelles d'Europe occidentale. PhD Thesis. Universite Peresani, M., Fiore, I., Gala, M., Romandini, M., Tagliacozzo, A., 2011. Late Neandertals and the intentional removal of feathers as evidenced from bird bone taphonomy at Fumane cave 44ky BP, Italy. Proc. Natl. Acad. Sci. 108, 3888e3893.
rez Ripoll, M., 2016. Los niveles de corral en el yacimiento neolítico de la Cova de Pe
n aportada por el estudio de la fauna. l'Or sectores k34, k35 y k36. Informacio Del neolític a l’edat del bronze en el Mediterrani occidental. Estudis en homenatge a Bernat Martí Oliver. TV SIP, 119: 119.124. Rasmussen, S.O., Andersen, K.K., Svensson, A.M., Steffensen, J.P., Vinther, B.M., Clausen, H.B., Siggaard-Andersen, M.-L., Johnsen, S.J., Larsen, L.B., Dahl€thlisberger, R., Fischer, H., Goto-Azuma, K., Jensen, D., Bigler, M., Ro Hansson, M.E., Ruth, U., 2006. A new Greenland ice core chronology for the last glacial termination. J. Geophys. Res. 111, D06102. Rasmussen, S.O., Bigler, M., Simon, P., Blockley, S.P., Blunier, T., Buchardt, S.L., Clausen, H.B., Cvijanovic, I., Dahl-Jensen, D., Johnsen, S.J., Fischer, H., Gkinis, V., Guillevic, M., Hoek, W.Z., Lowe, J.J., Pedro, J.B., Popp, T., Seierstad, I.K., Steffensen, J.P., Svensson, A.M., Vallelonga, P., Vinther, B.M., Walker, M.J.C., Wheatley, J.J., Winstrup, M., 2014. A stratigraphic framework for abrupt climatic changes during the Last Glacial period based on three synchronized Greenland ice-core records: refining and extending the INTIMATE event stratigraphy. Quat. Sci. Rev. 106, 14e28. ~o, J., 2018. Abrigo de la Real, C., Eixea, A., Sanchis, A., Morales, J.V., Villaverde, V., Zilha Quebrada level IV (Valencia, Spain): interpreting a Middle palaeolithic palimpsest from a zooarchaeological and lithic perspective. J. Palaeol. Archaeol. https://doi.org/10.1007/s41982-018-0012-z. Rendu, W., Costamagno, S., Meignen, L., Soulier, M.C., 2011. Monospecific faunal spectra in Mousterian contexts: implications for social behavior. Quat. Int. 247, 50e58.
n de Espan ~ a 1: Rivas-Martínez, S., 1987. Memoria del mapa de series de vegetacio 400.000. I.C.O.N.A., Madrid. Roebroeks, W., 1988. From Find Scatters to Early Hominid Behaviour. A Study of
de re (The Middle Palaeolithic Riverside Settlements at Maastricht-belve Netherlands). Leiden University Press, Leiden.
nchez Gon ~ i, M.F., Landais, A., Fletcher, W.J., Naughton, F., Desprat, S., Duprat, J., Sa 2008. Contrasting impacts of Dansgaard-Oeshger events over a western European latitudinal transect modulated by orbital parameters. Quat. Sci. Rev. 27, 1136e1151.


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032


n Marco et al. / Quaternary Science Reviews xxx (xxxx) xxx Y. Carrio ~ i, M.F., Bard, E., Landais, A., Rossignol, L., d'Errico, F., 2013. Air-sea S
anchez Gon temperature decoupling in western Europe during the last interglacial-glacial transition. Nat. Geosci. 6, 837e841. ~o, J., 2013. Los conjuntos Sanchis, A., Morales, J., Real, C., Eixea, A., Villaverde, V., Zilha faunísticos del Paleolítico medio del Abrigo de la Quebrada (Chelva, Valencia), I Jornades d'Arqueoozologia. In: Museu de Prehistoria de Valencia. 1 i 2 de desembre de 2011, Valencia, pp. 65e82. Schenck, T., Groom, P., 2018. The aceramic production of Betula pubescens (downy birch) bark tar using simple raised structures. A viable Neanderthal technique? Archaeol. Anthropol. Sci. 10 (1), 19e29. https://doi.org/10.1007/s12520-0160327-y. € lzer, HAUPT. Bern und Schweingruber, F.H., 1990. Anatomie Europ€ aischer Ho Stuttgart.
, C., Rubio-Jara, S., Panera, J., Pe
rez-Gonz
Sese alez, A., 2011. Micromamíferos del Pleistoceno Superior del yacimiento de PRERESA en el valle del Manzanares y su
n a la reconstruccio
n paleoambiental de la cuenca de Madrid contribucio durante el Pleistoceno. Estud. Geol. 67, 471e494. Shillito, L.-M., Almond, M.J., Nicholson, J., Pantos, M., Matthews, W., 2009. Rapid characterisation of archaeological midden components using FT-IR spectroscopy, SEMeEDX and micro-XRD. Spectroc. Acta Pt. A-Molec. Biomolec. Spectr. 73, 133e139. Sievers, C., 2006. Seeds from the Middle stone age layers at sibudu cave. South. Afr. Humanit. 18, 203e222. Sorensen, A.C., 2017. On the relationship between climate and Neandertal fire use during the Last Glacial in south-west France. Quat. Int. 436, 114e128. Sorensen, A.C., Claud, E., Soressi, M.A., 2018. Neandertal fire-making technology inferred from microwear analysis. Sci. Rep. 8, 10065. https://doi.org/10.1038/ s41598-018-28342-9. Stiner, M.C., Munro, N.D., Surovell, T.A., Tchernov, E., Bar-Yosef, O., 1999. Paleolithic population growth pulses evidenced by small animal exploitation. Science 283, 190e194.

ry-Parisot, I., 2001. Economie
olitique. Expe
rimentaThe des combustibles au Pale
ologique, n. tion, taphonomie, anthracologie, Dossier de Documentation Arche
20. CNRS Editions, Paris. Tormo, C., Guillem, P.M., 2015. Los micromamíferos del Paleolítico medio del Abrigo de la Quebrada (Chelva, Valencia): niveles I-VIII. In: Sanchis, A., Pascual, J.L. (Eds.), Preses petites i grups humans en el passat. II Jornades d'Arqueozoologia ria de Vale ncia, pp. 181e208. del Museu de Prehisto Tsartsidou, G., Karkanas, P., Marshall, G., Kyparissi-Apostolika, N., 2015. Palaeoenvironmental reconstruction and flora exploitation at the Palaeolithic cave of Theopetra, central Greece: the evidence from phytolith analysis. Archaeol. Anthropol. Sci. 7 (2), 169e185.
olithique infe
rieur et moyen entre les valle
es de la Dordogne Turq, A., 1992. Le Pale
de Bordeaux, p. 1. et du Lot. PhD Thesis. Universite Uzquiano, P., Arbizu, M., Arsuaga, J.L., Adan, G., Aranburu, A., Iriarte, E., 2008. Datos
n entre 40-32 Ka. BP: antrapaleoflorísticos en la Cuenca media del Nalo coan
alisis de la Cueva del Conde (Santo Adriano, Asturias). Rev. Cuatern. Geomorfol. 22 (3e4), 121e133.
, C., Baena, J., 2012. Uzquiano, P., Yravedra, J., Ruiz Zapata, B., Gil García, M.J., Sese Human behaviour and adaptations to MIS 3 environmental trends (>53e30 ka BP) at Esquilleu cave (Cantabria, northern Spain). Quat. Int. 252, 82e89.

17

Vernet, J.-L., Bazile, E., Evin, J., 1979. Coordination des analyses anthracologiques et des datations absolues sur charbons de bois. In: Bull. Soc. Prehist. Fr. 76-3, pp. 76e79. Vidal-Matutano, P., 2017. Firewood and hearths: Middle palaeolithic woody taxa distribution from el Salt, stratigraphic unit xb (eastern Iberia). Quat. Int. 457, 74e84.
n, B., Mallol, C., 2015. Neanderthal fireVidal-Matutano, P., Hern
andez, C.M., Galva wood management: evidence from Stratigraphic Unit IV of Abric del Pastor (Eastern Iberia). Quat. Sci. Rev. 111, 81e93.
ry-Parisot, I., 2017. Dead wood gathering among Vidal-Matutano, P., Henry, A., The Neanderthal groups: charcoal evidence from Abric del Pastor and El Salt (Eastern Iberia). J. Archaeol. Sci. 80, 109e121.
rez-Jorda , G., Herna
ndez, C.M., Galva
n, B., 2018. MacroVidal-Matutano, P., Pe botanical evidence (wood charcoal and seeds) from the Middle Palaeolithic site of El Salt, Eastern Iberia: palaeoenvironmental data and plant resources catchment areas. J. Archaeol. Sci. Rep. 19, 454e464. ~o, J., 2012. Importancia y valoracio
n de la Villaverde, V., Eixea, A., Rios, J., Zilha
n microlevallois en los niveles II y III del Abrigo de la Quebrada produccio (Chelva, Valencia). Zephyrus LXX, 13e32. Villaverde, V., Martínez-Valle, R., Guillem, P.M., Fumanal, M.F., 1996. Mobility and the role of small game in the Middle paleolithic of the central region of the Spanish mediterranean: a comparison of Cova Negra with other paleolithic deposits. In: Carbonell, E., Vaquero, M. (Eds.), The Last Neandertals, the First Anatomically Modern Humans. Universitat Rovira i Virgili, pp. 267e288.
n a la industria lítica del Abrigo Villaverde, V., Eixea, A., Zilh~ ao, J., 2008. Aproximacio de la Quebrada (Chelva, Valencia). Treballs arqueol. 14, 213e228. ~o, J., Sanchis, A., Real, C., Bergada , M., 2017. Diachronic Villaverde, V., Eixea, A., Zilha variation in the Middle paleolithic settlement of abrigo de la Quebrada (Chelva, Spain). Quat. Int. 435, 164e179. Wang, Y., Hope Jahren, A., Amundson, R., 1997. Potential for 14C dating of biogenic carbonate in hackberry (Celtis) endocarps. Quat. Res. 47, 337e343. Weiner, S., Dove, P.M., 2003. An overview of biomineralization processes and the problem of the vital effect. Rev. Mineral. Geochem. 54 (1), 1e29. ~o, J., 2007. The emergence of ornaments and art: an archaeological perspective Zilha on the origins of behavioural “modernity”. J. Archaeol. Res. 15, 1e54. ~o, J., Angelucci, D., Badal, E., d'Errico, F., Daniel, F., Dayet, L., Douka, K., Zilha
rdez, R., Murcia-Mascaro
s, S., Higham, T., Martínez-Sanchez, M.J., Montes-Berna
rez Sirvent, C., Rolda
n-García, C., Vanhaeren, M., Villaverde, V., Wood, R., Pe Zapata, J., 2010. Symbolic use of marine shells and mineral pigments by Iberian Neandertals. Proc. Natl. Acad. Sci. Unit. States Am. 107 (3), 1023e1028. ~o, J., Ajas, A., Badal, E., Burow, C., Kehl, M., Lo
pez-Sa
ez, J.A., Pimenta, C., Zilha Preece, R.C., Sanchis, A., Sanz, M., Weniger, J.-C., White, D., Wood, R.,
n: a multi-proxy MIS 3 Angelucci, D.E., Villaverde, V., Zapata, J., 2016. Cueva Anto to MIS 5a paleoenvironmental record for SE Iberia. Quat. Sci. Rev. 146, 251e273. ~o, J., Anesin, D., Aubry, T., Badal, E., Cabanes, D., Kehl, M., Klasen, N., Lucena, A., Zilha Martín-Lerma, I., Martínez, S., Matias, H., Susini, D., Steier, P., Wild, E.M., Angelucci, D.E., Villaverde, V., Zapata, J., 2017. Precise dating of the middle-toupper paleolithic transition in murcia (Spain) supports late neandertal persistence in Iberia. Heliyon 3 (11), 2017. https://doi.org/10.1016/j.heliyon.2017. e00435.


n Marco, Y et al., Climate, environment and human behaviour in the Middle Palaeolithic of Abrigo de la Quebrada Please cite this article as: Carrio (Valencia, Spain): The evidence from charred plant and micromammal remains, Quaternary Science Reviews, https://doi.org/10.1016/ j.quascirev.2018.11.032