A context for the last Neandertals of interior Iberia: Los Casares cave revisited
A context for the last Neandertals of interior Iberia: Los Casares cave revisited
Manuel Alcaraz-Castaño 0 1
Javier Alcolea-GonzaÂ lez 1
Martin Kehl 1
Rosa-MarÂõa Albert 1
Javier Baena-Preysler 1 3
Rodrigo de BalbÂõn-Behrmann 1
Felipe Cuartero 1 3
Gloria Cuenca-Besc oÂs 1 2
Fernando JimeÂ nez-Barredo 1
JoseÂ -Antonio LoÂ pez-SaÂ ez 1
Raquel PiqueÂ 1
David RodrÂõguez-AntoÂ n 1
Jos eÂ Yravedra 1
Gerd-Christian Weniger 0 1
0 Neanderthal Museum , Mettmann, Germany , 2 Area of Prehistory, University of AlcalaÂ , Alcal aÂ de Henares, Spain, 3 Institute of Geography, University of Cologne , Cologne, Germany, 4 ERAAUB ( Department of History and Achaeology), University of Barcelona , Barcelona, Spain, 5 ICREA, Barcelona , Spain
1 Editor: Michael D. Petraglia, Max Planck Institute for the Science of Human History , GERMANY
2 Aragosaurus-IUCA, Department of Geosciences, University of Zaragoza , Zaragoza , Spain , 8 CENIEH (National Research Centre on Human Evolution) , Burgos , Spain , 9 Archeobiology Research Group, History Institute , CCHS CSIC, Madrid , Spain , 10 Department of Prehistory, Autonomous University of Barcelona , Barcelona , Spain , 11 Department of Prehistory, Complutense University of Madrid , Madrid , Spain
3 Department of Prehistory and Archeology, Autonomous University of Madrid , Madrid , Spain
Introduction and objectives Although the Iberian Peninsula is a key area for understanding the Middle to Upper Paleolithic transition and the demise of the Neandertals, valuable evidence for these debates remains scarce and problematic in its interior regions. Sparse data supporting a late Neandertal persistence in the Iberian interior have been recently refuted and hence new evidence is needed to build new models on the timing and causes of Neandertal disappearance in inland Iberia and the whole peninsula. In this study we provide new evidence from Los Casares, a cave located in the highlands of the Spanish Meseta, where a Neandertal-associated Middle Paleolithic site was discovered and first excavated in the 1960's. Our main objective is twofold: (1) provide an updated geoarcheological, paleoenvironmental and chronological framework for this site, and (2) discuss obtained results in the context of the time and nature of the last Neandertal presence in Iberia.
Data Availability Statement: All relevant data are
within the paper and its Supporting Information
files. Archeological assemblages from Los Casares
cave are available at the Museo de Guadalajara
(Spain) and Museo ArqueoloÂgico Nacional (Madrid,
Funding: This research was funded by a Marie
Curie Intra European Fellowship within the 7th
European Community Framework Programme
under the project `Testing population hiatuses in
We conducted new fieldwork in an interior chamber of Los Casares cave named `Seno A'.
Our methods included micromorphology, sedimentology, radiocarbon dating, Uranium/
Thorium dating, palinology, microfaunal analysis, anthracology, phytolith analysis, archeo
zoology and lithic technology. Here we present results on site formation processes,
paleoenvironment and the chronological setting of the Neandertal occupation at Los
Casares cave-Seno A.
Results and discussion
The sediment sequence reveals a mostly in situ archeological deposit containing evidence of both Neandertal activity and carnivore action in level c, dated to 44,899±42,175 calendar
the Late Pleistocene of Central Iberia: a
geoarchaeological approach' (Grant number
628179) (MAC, GCW). It was also supported by
funding of the German Research Foundation's
(DFG) project CRC 806 ªOur Way to Europeº
(http://www.dfg.de/en/) (GCW, MK), and the
Spanish Ministry of Economy, Industry and
portal/site/mineco/), project numbers
HAR201348784-C3-3-P (JBP, FC, MAC),
HAR2016-76760C3-2-P (JBP, FC, MAC), HAR2013-43701-P (JALS)
and CGL2012-38434-C03-01 (GCB). MAC
currently holds a post-doc fellowship (Ayuda para
la AtraccioÂn de Talento Investigador 2016-T2/
HUM-1251) awarded by the Comunidad de Madrid
(http://www.madrimasd.org/madrid-cienciatecnologia/). Publication fees were funded by the
FP7 Post-Grant Open Access Pilot (OpenAIRE) of
the European Commission. The funders had no
role in study design, data collection and analysis,
decision to publish, or preparation of the
Competing interests: The authors have declared
that no competing interests exist.
years ago. This occupation occurred during a warm and humid interval of Marine Isotopic
Stage 3, probably correlating with Greenland Interstadial 11, representing one of the latest
occurrences of Neandertals in the Iberian interior. However, overlying layer b records a
deterioration of local environments, thus providing a plausible explanation for the
abandonment of the site, and perhaps for the total disappearance of Neandertals of the highlands of
inland Iberia during subsequent Greenland Stadials 11 or 10, or even Heinrich Stadial 4.
Since layer b provided very few signs of human activity and no reliable chronometric results,
and given the scarce chronostratigrapic evidence recorded so far for this period in interior
Iberia, this can only be taken as a working hypothesis to be tested with future research.
Meanwhile, 42,000 calendar years ago remains the most plausible date for the abandonment of interior Iberia by Neandertals, possibly due to climate deterioration. Currently, a later survival of this human species in Iberia is limited to the southern coasts.
The Iberian Peninsula has long been considered a crucial scenario for the Middle to Upper
Paleolithic transition and the replacement of Neandertals by Modern Humans [1±6]. Since
the late 1980's, a key point on these discussions was the contention that Neandertals persisted
in the center and south of Iberia until at least c. 36.7±34.5 ka cal BP [
], or even as late as c. 32±
28 ka cal BP [
]. This suggested that Neandertals and Modern Humans coexisted for several
millennia, since Modern humans were presumably established in the northern regions of the
peninsula from around 42±40 ka cal BP, or even earlier [
2, 3, 8
]. Here we focus on the interior
lands of Iberia, which are dominated by the highlands of the Northern and Southern Mesetas
divided by the Central System mountain range (Fig 1). Despite these inland territories had
traditionally contributed with some chronometric evidence to the late survival model,
reevaluation of the few sites involved has suggested however that no late Mousterian survival took
place in inland Iberia [6, 9]. Since still few sites from this area have contributed to this
discussion, new evidence is needed to build new models concerning the timing and causes of
Neandertal disappearance in inland Iberia and the whole peninsula. A new interdisciplinary
research project on Los Casares cave is aimed at moving forward in these scientific problems.
Los Casares is a limestone cave located in the interior regions of the Iberian Peninsula
(Spain). Its archeological potential is known since the late 19th century, when first scientific
explorations of the cavity pointed to the presence of bones and fossils in the floor and walls,
and a historical site was discovered outdoors . However, the relevance of this cave for the
Paleolithic field became evident in the 1930's, when its first Upper Paleolithic rock engravings
were described by J. CabreÂ [11±13]. Later, between 1966 and 1968, a team directed by I.
BarandiaraÂn conducted the first systematic excavations in Los Casares, showing archeological
deposits containing Middle Paleolithic assemblages in two different areas [
]. First deposit was
located at the entrance hall of the cave, named Vestíbulo (Vestibule in Spanish), and it
consisted of clayey sediments filling a short gallery at the bottom of this area (Fig 2). As reported
by BarandiaraÂn and recently observed by us, the presence of remnant sediments attached to
the walls at different parts of this vestibule suggests that a now-destroyed larger deposit
probably existed in this area. This is a very plausible hypothesis considering the long history of
occupations and incursions documented both inside and outside the cavity from the Chalcolithic
to Modern times, including its use as a sheep shelter during the 20th century [
2 / 54
Fig 1. Regional setting of Los Casares. Location of Los Casares cave in the Iberian Peninsula (A) and in the Geologic map of the
Guadalajara province (B). C: 3D view of Los Casares cave and the Linares and Valdebuitre valleys (Aerial photography and Digital Terrain
ModelÐPNOAÐfrom Instituto GeograÂfico Nacional de España).
The second site was found in a deeper area of the cave, the so-called Seno A, an interior
chamber where a larger deposit was discovered all along the place (Fig 2). Despite the area
excavated here was of 21 square meters, Mousterian assemblages were scarcer, and recorded
lithic artifacts were less than half in number than those found in the vestibule [
3 / 54
Fig 2. General plan of Los Casares cave showing Vestibule and Seno A areas.
Chalcolithic layer containing ceramics, lithics and faunal remains was also recorded at the top
of the sequence of the Seno A site [
Archeological assemblages recovered at the two areas excavated in Los Casares not only
included faunal and Mousterian lithic assemblages, but also a Neanderthal metacarpal bone
found at the Seno A Middle Paleolithic layers (Fig 3B). This finding, together with the
interesting nature of the lithic assemblages, composed of a high proportion of retouched tools,
especially in the Seno A (Fig 3A), made Los Casares one of the most relevant sites for the study of
the Middle Paleolithic in interior Iberia during the last quarter of the 20th century. The scarcity
of Late Pleistocene sites in these regions at the time, and the high quality of the monographic
publication produced shortly after the excavations [
] were also key points stressing the
relevance of this site for the study of the Iberian Middle Paleolithic. Furthermore, the presence of
Upper Paleolithic rock art in such an interior region, far away from the classic Cantabrian and
Mediterranean clusters, and including a striking proportion of anthropomorph figures ,
was also an indirect factor boosting the importance of Los Casares Middle Paleolithic site.
Despite this relevance, no scientific studies had been published on Los Casares Paleolithic
record since the 1970's, besides some partial analysis of the rock art , and some reviews of
4 / 54
Fig 3. Main findings of the 1960's excavations at Los Casares cave. A: Mousterian artefacts. All come from level c of Seno A except for
numbers 33, 34 and 36 (modified after [
]). B: Neandertal metacarpal found in square 8V' of Seno A (bar is 5 mm) (modified after [
the faunal [
], and lithic assemblages excavated by BarandiaraÂn . Therefore, there was
a significant scarcity of modern data hindering any attempt to integrate Los Casares evidence
in current debates on the Middle Paleolithic settlement of Iberia and southwest Europe. Data
on site formation processes were lacking, chronometric evidence was lacking, and
paleoecological information was virtually absent. In sum, Los Casares Middle Paleolithic record was
behind the times of current Paleolithic research.
In the summer of 2014 we started a new project aimed at the study of population dynamics
and human-environment interactions during the Late Pleistocene in the central region of the
Iberian Peninsula. A main factor driving this project was that record of this area was poorly
known compared to the coastal regions, and in the case of the Middle Paleolithic this was
especially evident concerning occupation of caves [21, 22]. Together with other two sites in the
Guadalajara province (Spain), we selected Los Casares as a case study that could show relevant
data on the Middle Paleolithic settlement of inland Iberia. It was our contention that Los
Casares potential had been inexplicably neglected since the 1970's, and therefore modern
geoarcheological investigations could bring into light new insights for the understanding of
Neandertal adaptations at this once key site of the Iberian Middle Paleolithic.
Overall, our main objective was to gain a better geoarcheological understanding of Los
Casares Middle Paleolithic site in order to contribute to current debates on the Neanderthal
settlement of inland Iberia. Among these debates, the long-claimed Mousterian late survival in
the central and southern areas of the peninsula , and the nature of human adaptations to
the harsh environments of the upland regions of the Spanish plateau [
], were the most
relevant. Both are currently under dispute [4±6, 9, 25±31].
Here we publish results of an interdisciplinary geoarcheological investigation of Los
Casares-Seno A site, where we conducted new field and laboratory works. We undertook
micromorphological, sedimentological and taphonomic analyses aimed at deciphering site
formation processes, we performed radiocarbon and U/Th dating for setting up a chronological
framework, we conducted palynological, anthracological, micromammal, phytolith and
sedimentological analyses for elucidating environmental and climatic settings, and we studied
lithic and faunal assemblages for discussing Neandertal techno-economic behaviours.
5 / 54
Integration of results obtained by all these methods depicts an ecological and chronological
context for the last Neandertals living in this interior area of the Iberian Peninsula.
Regional and local setting
Los Casares is a southwest-oriented cave eroded in a limestone-dolomite cliff corresponding
to the Muschelkalk lithostratigraphic unit (Middle Triassic) (Fig 1B). It is in the environs of La
Riba de Saelices village (Guadalajara Province, Spain), located in the moorlands of SiguÈenza
and Molina de AragoÂn belonging to the Iberian Range, at the northern fringe of the Southern
Meseta at about 1040±1060 m asl (40Ê 56' 22'' N, 2Ê 17' 31'' W, Datum ETRS89) (Fig 1A). The
cave entrance and vestibule are situated at about 40 m above the southward widening valley
floor of the Linares River (Upper Tagus basin), on its left bank (Figs 1C and 4). Los Casares is
a diaclase cave, with few and small lateral galleries, and with a total length of about 264 m from
West to East (Fig 1A). When studying cave art, J. CabreÂ  defined three different concavities
within the main passage that he called, from outside to inside, ªSeno Aº, ªSeno Bº and ªSeno
Cº. Seno A is found after leaving the vestibule and passing a narrow gallery about 20 m long
(Fig 1B). It consists of an east-west trending cavity with a complex topography, about 20 m
long, 10 m wide and up to 4 m high. The archeological deposit object of this study is found all
along the Seno A chamber (Fig 5)
Materials and methods
Permits and repositories
All necessary permits were obtained for the described study, which complied with all relevant
regulations. Field and laboratory works at Los Casares cave were authorized by the Dirección
General de Cultura de la Junta de Comunidades de Castilla–La Mancha (Spain) (Exp.:
14.0955-P1 and Exp.: 14.0955-P3). Study of lithic and faunal remains curated at the Museo
Arqueológico Nacional (Madrid, Spain) was authorized by the Prehistory Department of this
The Los Casares lithic and faunal assemblages excavated in 2014±2015 are housed in the
Museo de Guadalajara (Guadalajara, Spain). Assemblages from the 1960's excavations are
housed at Museo Arqueológico Nacional (Madrid, Spain). Both repositories are accessible for
Fieldwork: Excavation, stratigraphy and sampling
Previous work at the Seno A conducted by I. BarandiaraÂn in the 1960's [
] consisted of the
archeological excavation of 21 square meters (Fig 6). A stratigraphic sequence of grey-greenish
and reddish-brown Holocene and Pleistocene sediments divided in eight sedimentary layers,
from level ªaº to level ªhº, was described in a test pit reaching a total depth of about 1 m
below the modern cave floor. However, in most of the excavated area only the first three layers,
subsequently divided in different sub-levels in some places, were reached, at a total depth of
30±40 cm (Fig 5). Archeological assemblages were found at layer a3, where ceramics, lithics
and faunal remains were assigned to the Chalcolithic and Early Bronze age. While layer b was
described as sterile, Middle Paleolithic assemblages, including a Neandertal metacarpal, were
identified at level c. Below this layer, a flowstone was identified as level d0, followed by a
heavily cemented layer d, very rich in animal bones but lacking any artefacts. Layer e was
identified as a stalagmitic crust, and lower layers f1, f2, g and h were considered archeologically
sterile (Fig 7). Although BarandiaraÂn judged that most of the deposit at the Seno A was
6 / 54
Fig 4. Local setting of Los Casares. A: Los Casares cave and the narrowing of the Linares River downstream of the `Milagros' valley. B:
Entrance to the cavity. C: General view of Los Casares cave from the south. D: View of the Linares River valley from the cave's entrance.
preserved in situ, he acknowledged the presence of post-depositional disturbance at some
areas, mainly related to clandestine excavations (Figs 5 and 6).
The southern profile produced by the above-mentioned test-pit, located in square 1-R', was
still preserved at the site in 2014, hence allowing a direct inspection. We thus identified the
sequence published by BarandiaraÂn, as well as a good example of one of the post-depositional
disturbance features referred by this author: a clandestine pit in the western part of the profile
7 / 54
Fig 5. View of the Seno A chamber prior to our fieldwork. Profile 1R' South produced by the 1960's excavations and adjacent disturbed
area are shown.
(Fig 7). Since this profile offered an excellent platform for starting new excavation works, our
first task consisted on its rejuvenation 25 cm to the south. We thus produced a new profile in
square 3-R', where the old sequence could be contrasted more in detail. In doing so, and in our
general fieldwork at the site, we used grid and datum established by BarandiaraÂn's team in the
1960's (Fig 6). During the two campaigns conducted in 2014 and 2015 we excavated a total
extension of slightly more than 5 square meters, divided in four different areas located at the
perimeter of the 1960's excavation. The first area consisted of the rejuvenation of profile 3-R'
South, the second was square 6-Q', the third included squares 1-O' and 2-O', and the fourth
consisted of squares 6-V', 8-V' and 8W'. All these areas were relatively close and hence
stratigraphic correlations between them were feasible (Fig 6).
Fieldwork methodology followed the excavation of natural levels, which were divided in
artificial layers of 2±3 cm when needed. Both these layers and every archeological object or
feature larger than 2 cm (lithics, bones, charcoal fragments and human-made structures) were
three-dimensionally recorded using a Total Station, and orientation and dip of elongated lithic
and bone products were registered. All excavation layers were digitally photographed before
collecting archeological assemblages, which were also pictured in detail in special cases (i.e.
concentration of objects or relevant lithic artefacts). Stratigraphic profiles and some excavation
plans were also hand-drawn. Every square meter was divided in sectors of 33 sq cm and
sediment was packed accordingly. This sediment was later floated and wet-screened at 2 and 1
mm mesh sieves at the Laboratory of Prehistory of the University of AlcalaÂ, where most of the
lagomorphs, micromammals, charcoals and lithic debris were acquired. Samples for
micromorphology, pollen, phytolith and Uranium/Thorium dating of flowstone were also collected
8 / 54
Fig 6. Plan of the Seno A showing excavated areas in the 1960's and in 2014±2015. For the latter, archeological assemblages from
level c are plotted. However, for the 1960's excavations only lithic artefacts are plotted (after [
]), since no spatial recording of bone or
charcoals were done.
For micromorphology, five sediment monoliths were extracted from the upper part of the
sequence at different excavation areas, covering layers r to c2. Sampling of the heavily
cemented sediments of level d was not successful. Since sediments of the upper sequence were
very brittle, the samples were reinforced with gypsum bandages before extraction. The
monoliths 1 and 2 were taken from the south profile of square 3-R', monoliths 3 and 4 from the
south profile of square 1-O' and monolith 5 from the north profile of 8-W' (Fig 6). While
monolith 3 was kept for reference, the other monoliths were used for preparation of three
uncovered thin sections each (maximum size 60 mm x 80 mm), using methods described by
]. The thin sections were scanned with a flatbed scanner using polarizing foil for
inspection at low magnification. A petrographic microscope was used to study sediment
composition and fabric at magnifications of 12- to 500-fold using plane polarized light (PPL) or
crossed polarizers. In addition, oblique incident light (OIL) was used in some cases. The
9 / 54
Fig 7. Stratigraphic sequence excavated in the 1960's. A: Stratigraphy described by BarandiaraÂn in profile 1-R' South (modified after
]). B: Uncleaned profile 1-R' South prior to our fieldwork. Post-depositional disturbance at the upper western part was easily recognized.
description of thin sections followed terminology suggested by Stoops . All lab works were
conducted at the Institute of Geography of the University of Cologne.
Speleothem samples from level d0 were extracted with hammer and chisel from different areas
of the excavation. Two samples were extracted from square 3-R', one from square 6-Q' and
two more from square 2-O' (Fig 6). All samples were flowstones presenting a good degree of
crystallisation except for that collected at square 6-Q'. The latter was better defined as a calcite
incrusted layer or stalagmitic crust (Fig 8).
Three of these samples, S1, S2 and S3, were processed in the Laboratory of Uranium Series
at the CENIEH (Burgos, Spain). S1 was taken at square 3-R', while S2 and S3 were collected at
2-O'. Despite lateral variation of stratigraphic sequence all along the deposit, ªzº values
(deepness) of these samples were very similar (Figs 9 and 10). At the lab, smaller samples were
extracted using a hand drill and 0.8mm tungsten carbide drill bits. From S1 two sub-samples
were taken: S1a and S1b (Fig 8). After weighing, around 50mg precisely measured samples
were dissolved in HNO3 and digested in several steps involving HNO3, H2O2 and HCl
treatments. Uranium and thorium were then separated and their solutions purified by using two
ion exchange resin column steps (AG1X8 and UTEVA) following [34 and 35] sample
Uranium and thorium isotope measurements were performed using a Multicollector
Inductively Coupled Plasma Mass Spectrometer (MC-ICPMS, Thermo NEPTUNE) with a
membrane desolvator Aridus 2, X-cones and Jet interface pumping to increase the signal.
Nebulizer consisted on a PFA microtip calibrated for 50μl/min. Measurement protocols followed
10 / 54
Fig 8. Thin section scans of the flowstone of d0 at profile 3R under plain polarized light (A) and crossed polarizers (B). C shows a
scan of a calcite incrusted layer of d0 extracted from square 6-Q'. The lower image shows the three speleothem samples selected for
Uranium/Thorium dating (D). Note the small sampling areas.
can be found in [34 and 35]. Spike working solution used was made of a 1M ultrapure HNO3
gravimetrically produced solutions of NIST 4328c for 229Th, and IRMM3636a for 236U.
Standard solutions of those reference materials were used for standard sample bracketing. Both
standard solutions and samples were checked for U, Th elemental concentration by ICPOES
(PerkinElmer 5300DV) or ICPHRMS (ELEMENT XR).
230Th/U ages were obtained by solving numerically or graphically the corresponding
equation [36, 37]. Corrected ages were calculated assuming an initial isotope ratio
230Th/232Th = 4.4 x 10−6. Decay values considered for the equation were those found in .
Sample S1 showed some layers where oncolite accumulation is predominant. Since this is
usually related to bioturbations and could indicate open system conditions, attempts to date
those and surrounding areas would lead to obtain wrong U series dates. However, although
the lower layer sampled here (S1a) showed a high concentration of 232Th (151ng g-1),
230Th/232Th ratio was high and hence U/Th dating was performed. On the other hand, sample
S1b was collected from a series of well laminated layers of calcite formed after the upper
oncolite deposits at the top part of the speleothem (Fig 8). This area was wide enough to obtain
enough sample amounts suitable for U series dating over the brown-reddish narrow layer.
Concentration of 232Th (151ng g-1) was lower in this sample.
11 / 54
Fig 9. Stratigraphic sequence and vertical distribution of items recorded in profile 3-R'. Stratigraphic position of samples for pollen,
phytolith, micromorphology and Uranium/Thorium dating is shown.
From level b, two charcoal fragments and two faunal bones were selected for radiocarbon
dating. From level c, one charcoal fragment and three bones were selected. All these samples
12 / 54
Fig 10. Stratigraphic sequence and vertical distribution of items recorded in profile 2-O' West. Stratigraphic position of samples for
radiocarbon and Uranium/Thorium dating is shown.
(n = 8) were submitted to the CologneAMS centre at the University of Cologne. None of the
bones were anthropogenically modified, since no clear cut-marks or any other anthropogenic
sign were identified in the faunal assemblages collected at the 2014±2015 excavations.
Measurement of the five bones failed due to no collagen yield. Therefore, only three radiocarbon
measurements on charcoals could be obtained out of the eight samples tried.
Charcoal samples were first identified to taxon and then AAA (Acid-Alkali-Acid
extraction) processed according to sample preparation described by Rethemeyer et al. . Dating
results will be presented below in conventional radiocarbon years and as calibrated ages BP
using OxCal 4.3  and IntCal13 .
Six sediment samples of 5 square cm were extracted from profile 3-R' (Fig 9) during the 2014
excavation season following standardized techniques for archeological sites as described in [42
and 43]. Five of them (excluding that corresponding to the Chalcolithic a3 layer) were
prepared for pollen analysis at the CSIC labs (Madrid) using standard HCl sieving, HF and
density separation techniques (solution density 1.9) outlined in Burjachs et al. . After
processing, the samples were suspended in glycerin prior to being mounted on slides.
Counting was undertaken using a Nikon Elipse 50i light microscope at x400 magnification until a
sum of 300 total land pollen (TLP) had been achieved, excluding Aster type, Cardueae and
Cichorioideae with possible zoophily . Pollens were identified with the aid of the key in
Moore et al. , photographs in Reille  and modern reference material. Pinus nigra-type
pollen grains were categorised following measurements in Desprat et al. . Pollen diagrams
13 / 54
were constructed using TILIA 2.0 and TGView software  and percentages were based
upon the TLP.
CONISS  was used to assist with the biostratigraphic zonation of the pollen diagram in
Local Pollen Assemblage Zones (LPAZs) according to the dissimilarity matrix of Euclidean
distances and squared root transformed of percentage data. Ordination by principal
components analysis (PCA) was performed using CANOCO 4.5 software, as a linear interpretation
method for the fossil dataset since a previously detrended correspondence analysis (DCA)
pointed to a linear response of species data to environmental gradients . Samples were
square-root transformed for a better comparability. In the PCA scatter plot, pollen taxa are
shown as distance biplot arrows, and the direction of the arrow indicates the direction in
which the values of the corresponding taxa increases.
Fossils of small vertebrates were collected by hand during fieldwork seasons of 2014 and 2015,
and by wet-screening of sediments at 2 and 1 mm mesh sieves at the Laboratory of Prehistory
of the University of AlcalaÂ during 2015 and 2016. Microfaunal remains were found all through
the excavated areas in the Seno A, and in all stratigraphic layers besides a2, a3 and a4.
A total of 109 plastic bags filled with sediments were wet-screened at the laboratory. The
resulting concentrates were examined by naked eye as well as by optical microscopes.
Microfauna and other small fragments of large fossils were separated by picking up the elements.
The resulted 109 collections of fossils were then sent to the Department of Earth Sciences of
the University of Zaragoza, where assemblages were examined, photographed and stored.
A total of 102 out of the 109 analyzed samples showed fossil remains that were classified to
the species level. Additional washing with micro-mesh techniques and 10% HCl, and/or H2O2
was used when the surfaces of the molars, especially the enamel-dentine junction on the
occlusal surface, was covered with particles of sediment that impeded the visual analysis. This
anatomical region is needed pristine for the good classification and the morphometric analysis of
small mammals. Drawings were made after photographs taken with an Olympus SZ61
microscope with a camera attached to it. Images and measurements were taken with the camera and
the LCMicro software provided for the Olympus equipment.
Classification of small mammals into species was based on the morphology and biometry
of the occlusal surface of the molars, following general criteria of systematic paleontology
[50±54]. In each sample, we counted the number of skeletal elements, mainly dentition, and
calculated the minimum number of individuals (MNI). The input for computing the MNI are
the diagnostic skeletal parts that represents one individual of the species in a sample; i.e. two
left lower first molars (m1) of a given species represents two individuals, whereas two m1, one
left and one right represents one individual.
Charcoal remains were sampled by hand during fieldwork and by flotation in the Laboratory
of Prehistory of the University of AlcalaÂ. A total of 44 fragments of carbonized wood from
levels b, c and a5 have been studied in the Archeological Analysis Service of the Autonomous
University of Barcelona. Identification of taxa was carried out following standard procedures.
The anatomical patterns of each wood species were observed along three sections (transversal,
longitudinal tangential, and longitudinal radial) using a reflected light microscope equipped
with light field/dark field and objectives of 4x, 10x, 20x and 40x. Archeological samples have
been compared with modern woods as well as with wood anatomy atlases [
14 / 54
Three sediment samples from profile 3-R' were collected during the 2014 excavation season
for phytolith analyses. Samples were processed at the Laboratory for Palaeoecology and Plant
Palaeoeconomy±BioGeoLab (IMF±CSIC, Barcelona). The samples corresponded to sub-layers
c1, c2 and c3 of level c (Fig 9).
The extraction process follows the methods proposed by Madella et al., . Samples were
oven-dried at 80ÊC for 24 hours and screened with a 1 mm mesh to remove sands larger than
1000 μm. 15 millilitres of a 5% solution of hydrochloric acid (HCl) was added for 3 hours to
dissolve carbonate minerals and after the reaction ceased the acid was removed by centrifuging
the sample at 2000 RPM in 50 ml tubes. Samples were then de-flocculated with a 5% weight
solution of sodium hexametaphosphate ((NaPO3)6) for 36 hours, in order to disaggregate and
remove the clays, and centrifuged at 2000 rpm with distillate water. 30% solution of hydrogen
peroxide was added to samples for 3 hours to remove organic material. The resulting sediment,
what is the Acid Insoluble Fraction (AIF) , was then oven-dried at 60ÊC. A 10 ml of
sodium polytungstate solution (SPT) (Na6(H2W12O40)H2O) with a density of 2.35 g/cm3 was
added to separate siliceous minerals by density, vortexed and centrifuged for 3 min at 2000
rpm. The supernatant, where phytoliths are, was recovered with a Pasteur pipette discarding
the heaviest fraction and oven-dried at 40ÊC for 72 hours. 5μg per sample were finally placed
on microscope slides, mixed with Entellan and covered with 20 x 20 mm cover slips for
examination under the petrographic microscope (Olympus BX43) at 400x. The analysis of phytoliths
was conducted during 2016 in the Archeological Analysis Service of the Autonomous
University of Barcelona.
Morphological and taxonomical identification of phytoliths was based on standard
literature [58±62], including the PhytCore online data base . Other references from
paleoclimatic and ecological analogues, such as the areas from the Mediterranean-Alpine climatic
zones , as well as references from sedimentological and paleoenvironmental contexts from
the Iberian Peninsula albeit from different chronologies (MIS 4±5), were also consulted .
Special attention was paid to the presence of weathered phytoliths due to postdepositional
processes [66, 67].
Archeozoology and taphonomy
52 faunal remains from level b, 1,318 from level c and 85 from level d were subject to
zooarcheological and taphonomic analyses at the Prehistory Department of the Complutense
University of Madrid. No human remains where identified despite close inspection by
paleoanthropologists. While all remains from levels b and d corresponded to the 2014±2015
fieldworks, assemblages from level c included 515 remains from the 1960's excavations
], housed at the Museo ArqueoloÂgico Nacional (Madrid), and 803 from the recent ones.
Since it was hypothesized that an archeological selection of bone fragments could have be
done during the 1960's fieldworks [
], most probably due to a lack of wet-screening, data
from the two assemblages were recorded separately in a first stage of research.
Studied remains included both identifiable and unidentifiable fragments and the
taxonomical identification was based on reference materials. When the identification was not feasible,
epiphyses, axial and shaft fragments were assigned to three animal weight/size classes: 1)
small-sized carcasses, <100 kg (e.g. Capra pyrenaica, Rupicapra rupicapra), 2) medium-sized
carcasses, >100±300 kg (e.g. Cervus elaphus) and 3) large-sized carcasses, >300 kg (e.g. Equus
ferus, Bos primigenius).
The estimation of NISP (Number of Identified Specimens) and MNI (Minimum Number
of Individuals) was used to determine the most appropriate features of the faunal taxonomic
15 / 54
distribution. NISP follows Lyman's synthesis  and MNI is based on Brain's model  that
includes bone laterality -right/left- and animal age. Furthermore, skeletal profiles and MNI
consider shaft thickness, section shape and medullar surface properties . In this way, bones
were divided into four anatomical regions: 1) cranial (antlers-horn, skull, mandible and
dentition), 2) axial (vertebrae, ribs, pelvis and scapula, sensu ), 3) upper appendicular limbs
(humerus, radius, ulna, femur, patella and tibia) and 4) lower appendicular limbs (metapodial,
carpals, tarsals, phalanges and sesamoideal).
Mortality patterns were divided into (1) infants (individuals dead before the first six months
of life, as shown by the absence of the first permanent molar), (2) juvenile-prime adults
(individuals showing the second permanent molar and deciduous p4) and (3) adults (those with all
permanent teeth). Age profiles were estimated from tooth crown wear and the emergence of
the teeth according to Stelle  for deer, PeÂrez Ripoll  for ibex and Levine  for Equus.
A systematic observation of bone surfaces to explore the presence of cut, percussion and
tooth marks was also carried out with 10X-20X hand lenses and different lighting . Our
diagnostic criteria for cut, tooth and percussion marks are the ones defined respectively by
] and Potts and Shipman , Blumenschine , and Blumenschine and Selvaggio
. For comparative purposes, observation of bone surfaces includes the observation of
epiphysis and shafts . Modifications of bone surfaces were also quantified by types of
fragments and bone sections [80, 81] based on NISP values. The presence of tooth, percussion and
cut marks was recorded for the whole assemblages, and percentages of tooth, percussion and
cut marks included only bones with a good surface preservation. Weathering stages were also
observed following Behrensmeyer  to estimate the bone subaerial time exposure. Water
effects on bone surfaces were estimated according to the presence of abrasion, polishing,
rounded bones, and carbonates following Parson and Brett , CaÂceres  and Yravedra
. Signs of polishing, rounding or abrasion are to be expected in transported assemblages,
but also in non-transported assemblages exposed to circulating water and mobile sediments,
such as those encased in sand strata . Biochemical alterations were estimated according to
DomÂõnguez-Rodrigo & Barba . To differentiate between green and dry fractures on long
bones we analyzed shafts larger than 30 mm following Villa & Mahieu's  criteria.
Besides one single flint flake collected in layer b, all lithic artefacts recovered at Los
CasaresSeno A come from level c. In consonance with those recorded by BarandiaraÂn [
] in the same
level during the 1960's excavations, they represent a scarce sample. In the circa 5 square meters
excavated, we collected just 7 lithic artefacts, including two debris recovered after wet-sieving.
Assemblages recovered at the Seno A during the 1960', summing up to 38 lithic artefacts, were
studied at the Museo Arqueológico Nacional (Madrid), where they are currently curated.
However, we considered only 37 items, since one blade, found in mostly disturbed square 2-Q' as
reported by this scholar (Fig 6), must be conceived as a very likely intrusion from above [
Therefore, we analyzed a total of 44 lithic artefacts, 32 of them produced on flint (72.7%), and
12 on quartzite (27.3%). A spatial distribution of the whole assemblage was possible due to the
spatial recording of objects during the old excavations [
]. Plotting of artefacts, both
horizontal and vertical, is shown in Figs 6 and 9±15. Density of artefacts is very low, only reaching 1.76
artefacts per square meter.
Lithic artefacts were studied at the Prehistory Laboratory of the University of AlcalaÂ under
the chaÃõne opeÂratoire or `operational sequence' approach as described by Inizan et al.  and
discussed in Bar-Yosef and Van Peer . We assigned each lithic artefact to one of the three
chaÃõne opeÂratoire stages commonly recognized in the literature . Thus, cortical flakes,
16 / 54
Fig 11. Stratigraphic sequence and vertical distribution of items recorded in profile 1-O' South and 1-O' West. Since some items
could not be plotted in both views due to stratigraphic dip, topographic numbering of items has been included.
preparation products and tested cores were assigned to the initialization stage or phase I, raw
blanks, core maintenance by-products and rejuvenation flakes to the exploitation stage or
phase II, and retouched blanks and exhausted cores to the consumption and abandonment
stage or phase III. We also analyzed artefacts in terms of formal recycling and reuse processes
as discussed by Amick  and Baena et al. [
Stratigraphic sequence and micromorphology
Stratigraphic sequence documented in the Seno A deposit during 2014±2015, preliminarily
published in [
], was first described after rejuvenation of profile 1-R' South previously
excavated by BarandiaraÂn in the 1960's. Main stratigraphic layers and sub-layers recognized at the
deposit were first identified at this profile, corresponding to the northern sector of square 3-R'
(hereafter, just `profile 3-R'). Also, this is the only area where we excavated level d, while in
other test pits we only reached the top of level d0. As previously mentioned, profile 3-R'
showed a similar stratigraphy as documented in the 1960's, but it was less affected by
bioturbation, and disturbance reflected in a pit infilling at the western side of the profile was less severe
than in the original profile 1-R' [
] (Fig 9). The top 2 to 3 cm of 3R', denominated as "r",
consisted of compacted, dark brown and black sandy loam, containing weathered rock debris,
pieces of charcoal and few artefacts. It represents a reworked surface layer probably compacted
by trampling in the recent past. Layer "a2", defined by BarandiaraÂn as a thin discontinuous
black band with concentrations of charcoal, although partially visible in some areas, was
17 / 54
Fig 12. Stratigraphic sequence and vertical distribution of items recorded in profile 6-Q' North.
integrated by us in layer r. Layer "a3" (about 5cm) consists of a light grey to greenish-yellow,
densely packed silty clay loam, with inclusions of charcoal, weathered rock fragments and few
Chalcolithic ceramics. At the base of this layer, another thin black band (a4) is found, very rich
in charcoal and Chalcolithic pottery. Layers a3 and a4 are rich in fine silt and clay displaying a
textural contrast to the underlying reddish brown sandy loams of layers b0 and b. The latter
layer contains few gravel and shows some yellowish patches and several discontinuous black
laminae, the lowermost one forming the lower boundary of layer b (about 4 cm thick). In
contrast to the stratigraphy of BarandiaraÂn (Fig 7A), we identified three sublevels for the sandy
loams of layer c (up to 10 cm thick) as based on differences in colour and degree of
compaction. Sublevel c3 was characterized by an orange colour and was only locally preserved. Below
layer c, flowstone of level d0, up to 3 cm thick, was found. It covered heavily cemented reddish
loams of level d, the latter being very rich in animal bone, but lacking any artefacts. Excavation
at 3-R' stopped after reaching another stalagmitic crust correlating with layer e as defined by
BarandiaraÂn (Fig 7A).
18 / 54
Fig 13. Stratigraphic sequence and vertical distribution of items recorded in profile 8-W' North.
Although presenting some minor differences concerning sub-layering of stratigraphic
levels, sequence described in 3-R' was also recognized in the other areas excavated in the Seno A
(Figs 9±15). An important difference was however registered in square 6-V', where a
prehistoric pit, most probably Neolithic or Chalcolithic, and recorded as layers a5 and a5.1,
penetrated the clays of layer b (Fig 14).
Detailed micromorphological descriptions of three different profiles are provided in tables
A to C (S1 Appendix) and illustrations of the thin sections including microstratigraphic
subdivision of archeological levels are shown in Figs 16±18. In some cases, archeological levels
defined in the field included several sediment layers in thin section, which have been
characterised separately, where appropriate. Monoliths 1, 2 and 5 cover most of the sequence
including layers r to c, whereas monolith 4 covers the lower part of the sequence only, starting with
a4. The profiles show slight differences in stratigraphy as indicated below.
Archeological level r consists of sandy loams rich in charcoal and bone fragments. The
sediment layers detected in thin section are densely compacted. In profile 8-W', level r can be
subdivided based on color and composition of coarse materials. At the top, an orange microlayer
is visible, which owes its color to phosphate accumulation probably from bat guano. Level a3 is
rich in silt and clay and contains many partly disintegrating limestone fragments in its upper
part, forming a light grey layer, which is found in all profiles sampled. This layer includes
admixtures of charcoal and siliceous fines, best visible in thin section 5.1, and, in profile 3-R',
it can be subdivided into two sublayers, the lower of which containing considerably less rock
Level a3 overlies the dark coloured level a4. In profile 3-R', a4 represents a black microlayer,
few mm in thickness, while in the other two profiles sampled it is about 1 cm thick. Level a4 is
mainly composed of fine pieces of charcoal and amorphous burned organic materials,
19 / 54
Fig 14. Stratigraphic sequence and vertical distribution of items recorded in profile 6-V'. Stratigraphic position of two dated charcoals
concentrated in two different microlayers in profile 1-O'. In profile 8-W', a4 is particularly
rich in small phosphatic coprolites of bat guano.
The underlying sediments have a more reddish or orange brown groundmass. Level b0 is
about 1 cm to 1.5 cm thick, shows a high degree of compaction and is moderately to strongly
enriched in phosphate. In profiles 1-O' and 8-W', level b0 has a well-developed
subhorizontally-laminated fabric (Fig 19E and 19F). Underlying layers of level b (or b1 in profile 8-W')
are much less compacted and less enriched in phosphate (e.g. Fig 19A and 19B). In profile
8-W', this level was subdivided into two layers: b1 consists of sandy loam with bones and
coprolites, while b2 represents an intercalation of several clayey or sandy to gravelly microlayers
20 / 54
Fig 25. Microphotographs of phytoliths identified at level c Los Casares cave-Seno A. Pictures were
taken at 400x. A) Rondel short cell; characteristic of Pooideae grass subfamily; B) Saddle short cell
characteristic of the Chlorodoideae grass subfamily; C) Crenate; phytolith characteristic of Pooideae grass
subfamily (Gramineae); D) Elongate echinate from inflorescence of Poaceae (Gramineae).
sedge (Cyperaceae) phytoliths were noted and represented between 1.2 and 2.4% of the total
phytolith counting [59, 60, 95].
Phytoliths from arboreal woody taxa, most probably dicotyledonous plants, were identified
in both samples (16.2% in c1 and 20.5% in c2). Nevertheless we cannot disregard the presence
of conifers, since some morphotypes such as blocky and tracheids may be found in both
groups  (Table 6).
In general terms, both phytolith assemblages are very similar and show a predominance of
grasses, especially of the Pooideae grass subfamily (C3) and to a lesser extent of the
Chloridoideae grass subfamily (C4). Morphological identification also shows the presence of plants
belonging to the Cyperaceae family, as well as woody plants (Dicotyledonous and possibly
Coniferae). This arboreal presence is even more significant considering that phytolith
production in grasses is substantially higher than in woody taxa (up to twenty times higher), especially
in Mediterranean-Alpine environments [61, 96, 97].
A detailed taxonomic and taphonomic analysis of macrofaunal remains has only been possible
for level c. Level b has yielded just 52 remains, of which only 8 could be identified as
Oryctolagus cuniculus. Remains from level d, only excavated in square 3-R', have been also included in
this study, although they represent a very uninformative sample composed of few remains of
33 / 54
Attribution of phytolith morphotypes to plant taxa, plant parts and types of vegetation is shown.
herbivores and carnivores (Tables 7 and 8), where bear is the most represented species both in
NISP and MNI.
At level c, the study of bone assemblages excavated in the 2014 and 2015 seasons has
confirmed previous suggestions [
] pointing out a bias in the recording of bone fragments during
the 1960's excavations. The low presence of undetermined material in the latter compared to
the new assemblages (Table 7) suggest that an artificial selection was made during the
excavation process, most probably due to an absence of sediment wet-screening. However, the higher
diversity of herbivores in the 1960's assemblage (Table 7) is best explained by the larger size of
the excavated area. Concerning carnivores, both assemblages show an equivalent abundant
sample (Table 7). Together with a higher presence of cranial remains in the old assemblage
(see below), this evidence suggests that data from the two assemblages should be presented
separately (Table 7 and Table F in S1 Appendix). However, since it is clear that both
assemblages come from the same stratigraphic context and they both present similar taxonomic and
taphonomic profiles, these limited recording biases did not prevent us of considering results
Considering both assemblages from the 1960's and recent excavations, level c has yielded
more than 1,300 faunal remains and a minimum number of 48 individuals. It shows a high
taxonomic diversity, with Iberian ibex as the most represented herbivore species, but also
including large bovids, horses, wild asses, deers, roe deers, chamois and wild boars. Among
carnivores, the most abundant groups are hyenids and ursids, but canids and felids are also
well represented (Table 7). Considering MNI, animals typical of rocky environments, such as
Iberian ibex and chamois, are the most relevant, followed by deer. As for carnivores, bear is
the most represented with 4 individuals (Table 8). Mortality patterns show that adult
34 / 54
Faunal assemblages from level c include remains from both old (1960's) and recent (2014±2015) excavations. ª% partialº refers to total of carnivores or
individuals dominate the faunal assemblage, both for carnivores and herbivores. Infant or
juvenile-prime adults individuals have only been identified for Iberian ibex, chamois, deer and
large bovid (Table 8).
Skeletal profiles for level c are biased by the low number of remains recorded for most
animal species, with only lagomorphs being above 100 remains. Cranial elements are the most
represented, ant teeth sum up to more than 50% of the sample for all taxa (Tables D and E in
S1 Appendix). This preeminence of cranial elements, including teeth, is more striking when
considering only the 1960's assemblage (Table F in S1 Appendix), and hence could be related
to a recording bias during the old excavations, as described above.
Among lagomorphs, all anatomical portions are found, but metapodials and phalanges
account for 50% of remains (Table D in S1 Appendix). Concerning level d, skeletal profiles,
dominated by teeth, are not considered representative due to the low amount of available
faunal remains (Table G in S1 Appendix).
35 / 54
Faunal assemblages from level c include remains from both 1960's and recent excavations. ª% partialº refers to total of carnivores or herbivores. A: Adult, J:
Juvenile and prime adult; I: Infant.
Taphonomic analysis of level c depicts a well-preserved assemblage, but showing an
important skeletal bias towards the denser bones, especially cranial remains (Tables D and E in
S1 Appendix), probably related to the bias recording of the 1960's fieldworks (Table F in
S1 Appendix). Weathering is only slightly recorded in bone surfaces, and the incidence of
biochemical alteration is documented in less than 15% of the bones. Trampling affects to 2.5% of
the bones, while hydric modification as showed by polishing, abrasion or carbonates, has been
documented in less than 5%. Regarding types of breakage, only 10% of bones larger than 30
mm shows dry pattern, while green fractures have been recorded in 20%. The remaining 70%
of bones show indeterminate breakage patterns. Therefore, impact of these processes on the
faunal representation is not relevant.
Carnivore action has been also recorded in level c, but not in a prominent way considering
that bones recording tooth marks are scarce in most taxa (Table 9). However, it is very likely
that carnivores were responsible for the disappearance of several osseous portions as showed
by (1) the mentioned teeth marks, (2) the presence of corrosion marks caused by digestive
36 / 54
CM: Cut marks, TM: tooth Marks.
processes in some lagomorph bones, (3) the relative high amount of carnivores in the
assemblage and (4) the absence of axial bones, such as ribs and vertebrae, coupled with the
predominance of dense bones, such as teeth or lower appendicular limb bones. In the bone assemblage
of layer d no carnivore alterations have been observed.
Concerning human activity, although no percussion marks have been observed in layer c, a
limited number of cut marks has been recorded on remains corresponding to Iberian ibex,
rabbit and wild ass (Table 9). While these marks denote some kind of human action on some
animal species, they are not abundant enough to propose any conclusion in terms of economic
behavior or subsistence strategies. No evidence of human action has been recorded on the
faunal assemblage of layer d.
In sum, faunal assemblages of Los Casares-Seno A can be considered as produced basically
by carnivore action, and only sporadically by humans in layer c.
In Table 10 we show all technological categories described for the lithic assemblage of Los
Casares-Seno A level c, and in Fig 26 we present the chaîne opératoire stages identified. Despite
the low quantity of artefacts found at the site, it is noteworthy the high proportion of products
corresponding to the consumption and abandonment stage (68.2%), being the rest assigned to
the exploitation phase (31.8%). No elements have been related to the initialization phase. This
predominance of consumption products is even higher when considering only artefacts made
on flint (71.8%), while in the case of quartzite is significantly lower but still high (58.3%).
Furthermore, most of these products are not simply retouched flakes, but formal tools, especially
sidescrapers. All blanks are flakes except for one sidescraper on blade and one raw blade, both
produced on flint.
Retouched products, highly dominated by sidescrapers (Table 10), are mostly configured
on blanks produced by recurrent centripetal Levallois methods as shown by centripetal scars
on dorsal surfaces and facetted platforms (Fig 27). Some of these tools present evidences of
recycling processes, such as exploitation of ventral surfaces, thus generating `core on tools'
pieces (Fig 27.7). A low number of small rejuvenation flakes produced during the resharpening
of sidescrapers' edges has been also found.
37 / 54
All these features suggest that the lithic assemblage preserved in level c of Los Casares-Seno
A, undoubtedly reflecting a typical Middle Paleolithic technology, is mostly related to
consumption activities. No knapping processes, besides some recycling or maintenance tasks,
were developed at this part of the cave, which was perhaps focused on specialized activities as
shown by the high presence of sidescrapers and other domestic tools.
Fig 26. Mousterian lithic artefacts recorded in level c of Los Casares cave-Seno A according to the
chaÃõne opeÂratoire stages. Note that stage I (Initialization) is totally absent.
38 / 54
Fig 27. Mousterian lithic artefacts from level c of Los Casares cave-Seno A. Sidescrapers (1,3,4 6±7), denticulate (2) and point (5). All
artefacts come from the 1960's excavations (curated in the Museo ArqueoloÂgico Nacional, Madrid) except 3, which was recovered in our
recent excavations. Item 7 is a sidescraper recycled into a core.
Concerning level b, only one flint flake was recovered during fieldworks. Although this
finding suggests that previous contentions that this layer was sterile [
] were probably wrong,
it does not suffice to make any chrono-cultural assignment for it.
Site formation processes
Micromorphological analyses have shown compelling evidence for site formation processes of
the Seno A deposit. Level d0 represents differentially crystallised stalagmitic crusts and
flowstones several cm thick and accumulated by chemical precipitation over a long period of time.
Although the lateral continuity of the well-crystallised crust in profile 3-R' is limited, level d0
represents a good stratigraphic marker for the unconsolidated overlying deposits.
Sediment composition and fabric of levels c and b suggest that they originate from an
interplay of different transport and deposition processes within Seno A. Subsurface flow in the
vadose zone of the cave system and possibly infiltration of fines through cracks has
accumulated diverse carbonate and siliclastic mineral grains including well-rounded limestone
boulders, siliceous gravel or phyllosilicate clay. In-situ remnants of water-laid deposits are
39 / 54
preserved as intercalation of sand/fine gravel with silt/clay in level b2 of profile 8-W'. Although
local concentration of well-rounded gravel, such as in layers b and cs4 of profile 1-O', also
reflect changes in flow velocity, these gravel do not appear in extended pockets or beds. Hence,
in all layers except of b2, subaqueous deposition is not clearly indicated.
The contribution of roof-fall during accumulation of levels c to r was probably limited,
because few angular to subangular limestone fragments were found. In the grey layers of level
a3 limited roof-fall is included. The small rock fragments disintegrate leaving a clayey loam
with reprecipitated calcite grains behind. This weathering product probably formed in
waterfilled basins of the cave floor.
During sediment accumulation of levels c, b0 and r, considerable zoogenic inputs of bat
guano and carnivore coprolites occurred. In addition, bone constitutes a major component of
most levels, but its origin may be related to both animals and humans. Charcoal is related to
anthropogenic input, while the low numbers of charcoal in sediments from sediments below
level a4 may at least partly be related to microbial degradation.
Postdepositional processes include corrosion and mechanical disintegration of limestone
fragments and calcite grains. Carbonate leaching is indicated by local presence of
undifferentiated b-fabrics, as well. Locally, heavily corroded carbonate grains are found in sand-size
pores. Calcite pedofeatures including infillings and coatings indicate precipitation of
secondary carbonates in several layers. This shows, that both partial leaching of carbonates and
precipitation of secondary carbonates occurred, with stronger intensities in the lower part of the
sequence, probably related to a longer period of time encompassed with sediments of level c.
Besides calcitic pedofeatures, few other types of pedofeatures were detected. Locally, iron
hydroxide or manganese oxide nodules, such as in level b0 of profile 8-W' were found. Overall,
few indicators of post-depositional mixing by sediment dwelling animals were detected in thin
A prominent feature in the studied thin section is the preservation of sediment boundaries
showing accumulation of small charcoal fragments or manganese precipitates at the former
surface or remnants of small sedimentary crusts of fine materials. The often high degree of
compaction directly underneath these sediment interfaces or in whole layers clearly point to
trampling effects during or after the accumulation of the layers. This is very obvious in
sediments from the current cave floor down to level c1. The sequence thus clearly shows good
preservation of layering, except of in its lower part (archeological level c) which neither shows
clear evidence of mixing such as burrows nor of preservation of former surfaces, compacted
parts or primary deposition by running water.
In sum, micromorphological features support the field distinction between dark or light
grey sediments of levels r to a4, and the reddish-brown deposits of levels b0 to c. The sediment
sequence in Seno A is well-stratified, particularly in the upper part down to above level c
where remnants of several former trampled cave floors are preserved as indicated by
characteristic sediment features. Mixing across boundaries between archeological levels was therefore
very limited and hence the deposit can be considered as mostly in situ, at least in analyzed
samples. The intensity of post-depositional changes including carbonate leaching and precipitation
as well as precipitation of phosphate is higher in level c and b0 than in the upper levels,
probably related to a longer time of exposure to this kind of diagenetic changes.
Chronological and paleoenvironmental framework
Despite problems experienced with collagen-depleted bones, two independent chronometric
methods place the Neandertal occupation of the Seno A within the middle-advanced stages of
MIS 3. As the U-series ages obtained for layer d0 flowstone provide a terminus post quem of c.
40 / 54
48 ka BP (sample S3) for layer c, radiocarbon date of 44.9±42.2 ka cal BP obtained in this layer
can be taken as a reliable approach to its age. This chronology, which is also consistent with
biostratigraphic data provided by micromammal analysis, places the Middle Paleolithic
occupation of Los Casares cave-Seno A within the final stages of the Neandertal presence in interior
Iberia as currently documented. Although no reliable chronometric evidence is available for
layer b, and its archeological content is uncertain and non-diagnostic, paleoenvironmental
data gathered at this layer provide useful insights into its potential age and implications, as it
will be discussed below.
Paleobotanical and microfaunal evidence presented in this study has substantially improved
previous knowledge of the environmental and climatic framework where Los Casares'
Neandertals lived. Taken together, pollen, microvertebrates, charcoal and phyotlith data firmly
point to a relatively temperate and humid interval within MIS 3 for level c. The presence of
taxa such as Acer, Tilia, Salix, Alnus, Pistacia terebinthus, Fraxinus and deciduous Quercus in
pollen samples collected in this level indicates a relatively forested Pyrenean oak landscape
enriched in mesophilous trees and shrubs with some black pines . The contention that
central Iberia contained deciduous oak populations during glacial stages  is supported by our
results, at least for MIS 3. In this sense, the study area can be considered as a glacial refuge for
deciduous oaks and other Late Pleistocene temperate taxa, probably associated with higher
water availability along river valleys, as has been reported for other nearby sites during MIS 2
Concerning the microvertebrates, the presence of forest-dwelling taxa, such as Sciurus
vulgaris and Apodemus, Mediterranean species such as Eliomys quercinus and Hystrix sp., as well
as species adapted to humid habitats such as Castor Fiber, Arvicola sapidus and Iberomys
cabrerae, also suggest a warm and humid environment for level c [54, 101±102]. The absence of
cold-indicator taxa in this level, such as the snow and tundra voles, is also of relevance here.
Evidence shown by proxies reflecting a more anthropogenic input into the site is in
agreement with the pollen and microvertebrate results. Phytolith data gathered at level c also point
to humid and warm environments, as shown by the high presence of Pooideae and
Chloridoideae grass subfamilies and woody plants such as dicotyledonous, which are indicative of
woodland landscapes and grassland or shrubs areas [59±60, 65, 96]. Although charcoal data have
been limited to the presence of Pinus t. sylvestris-nigra, this is best described as evidence
reflecting the trees supplying the fuel collected by Neandertals around the cave, as documented
in many sites in Iberia during MIS 3 [103, 104].
Despite the scarcity of archeological or paleontological sites yielding paleoenvironmental
data assigned to MIS 3 in interior Iberia, a good parallel for the paleoecological framework
reconstructed at Los Casares can be found at Zarzamora Cave (Segovia). This site, very close to
the northern foothills of the Central System range, is also dominated by Quercus and presents
a micromammal assemblage reflecting temperate and humid conditions . In the southern
part of the Central System range, the MIS 5 site of Camino, in Pinilla del Valle [52, 106] also
shows similar micromammal assemblages, albeit including some cold-indicators taxa which
are absent in Los Casares-Seno A level c. Beyond the Meseta, but still in an interior region of
AndalucÂõa, microfaunal evidence from Carihuela cave (Granada) correlates well with Los
Casares-Seno A assemblage, as reflected in the presence of Allocricetus bursae, the arvicolines
Iberomys cabrerae, Pliomys lenki and the water vole Arvicola sapidus .
Taking together paleoenvironmental and chronometric evidence, layer c of Los Casares
cave-Seno A is most probably correlated with Greenland Interstadial 11, starting at 43.3 ka BP
on the NGRIP δ18O timescale  (Fig 28). However, overlying layer b shows a very different
paleoenvironment composition, pointing to a later phase probably correlated with subsequent
stadial phases. LPAZ CS2, corresponding to level b, reflects a cold and arid climatic period
41 / 54
Fig 28. Correlation of radiocarbon calibrated date COL 4208.1.1 with Greenland Interstadials against
the NGRIP δ18O record [111±112]. U/Th sample 3 is shown as a terminus post quem for the Middle
dominated by Pinus nigra, evergreen Quercus, Helianthemum, Juniperus, Cytisus/Genista,
Poaceae and Artemisia. It thus demonstrates the climatic variability within the MIS 3 in inland
Iberia, and suggests the existence of relatively open black pine woodlands with some holm oak
stands, grasslands and an abundant shrub cover of broom communities and juniper [109,
110]. This is consistent with microfaunal evidence, as seen in the reduction in the number of
taxa identified in level b with respect to c. Also, the disappearance of forest-dwelling taxa that
were present in level c, such as Sciurus vulgaris and Apodemus species, and of Mediterranean
indicators such as the dormice, Eliomys quercinus, or the wood mouse, most probably record
an increasingly colder climate in layer b. In this context, survival of species such as Arvicola
sapidus and Iberomys cabrerae, both related to humid habitats , is best explained
considering that they were probably less affected by climatic stress than the
Woodland-Mediterranean indicators [54, 102].
Despite four radiocarbon attempts, there is no reliable chronometric evidence for layer b,
which yielded only 52 macrofaunal remains and a single non-diagnostic flake. However, given
chronometric results for levels c and d0, coupled with the paleoenvironmental and
geochemical evidence described above, it is very likely that level b corresponds to a stadial phase
following GI 11 as recorded in layer c. On this matter, the presence in layer b of the rodent species
Pliomys lenki, which went extinct during the Late Pleistocene, points to a MIS 3 chronology.
While the last appearance datum (LAD) of this species in northern Iberia is around 14 ka cal
BP , its presence in the central and southern regions of the peninsula, although not
radiometrically dated at the sites of Cova Negra  and Carihuela , suggest a LAD within
MIS 3 . Therefore, the most parsimonious interpretation is that layer b is of MIS 3 age,
and most probably not much more recent than layer c, as also supported by sedimentological
and geochemical data. Thus, GS 11, GS 10 or even Heinrich Stadial 4 (H4), spanning from c.
42 to 38 ka years ago [108, 116], are the most plausible correlations for layer b, and hence for a
tentative phase of occupation at Los Casares reflecting a very scarce presence in the cave, at
least for the interior area of Seno A. A subsequent hypothesis is that layer b indeed reflects the
final stages of Neandertal presence in Los Casares, occurring sometime between c. 42 and 38
ka cal BP, and perhaps also the very last occupation of this high area of the Iberian interior due
to climatic deterioration. However, given the scarce archeological content of layer b, some
further reflections on this hypothesis will be made below.
42 / 54
The last Neandertals of interior Iberia
Since the late 1980's, the center and south of the Iberian Peninsula has been considered a sort
of refuge where the last Neandertals persisted long after the first Modern Humans arrived to
the north of the Peninsula and the rest of Europe [2, 23, 107, 117±124]. More recently, some
authors have argued for a Neandertal survival south of the Ebro basin until at least c. 36.7±34.5
ka cal BP [
], while others propose dates of c. 32±28 ka cal BP for the extreme southern
regions of Iberia [
]. Considering that dates for the appearance of the Proto-Aurignacian in
the north of Iberia are well established around 42 ka cal BP [8, 125], a millennial coexistence
between Neandertals and Modern Humans at the peninsular scale was accepted by most
researchers until recently. However, in the very last years, new research focusing on the
chronometric evidence [
], and especially on new radiocarbon-dating projects based on
ultrafiltration pretreatment of bone samples [
], have questioned the late Neandertal survival
model, thus supporting previous criticisms already raised by some scholars [126±128]. After
refuting previously accepted late chronologies at the sites of Zarafarraya (MaÂlaga) and Jarama
VI (Guadalajara), and questioning the dates obtained in Gorham's cave (Gibraltar), Carihuela
(Granada), Gruta da Oliveira (Portugal) and Sima de las Palomas (Murcia), Wood et al. 
have proposed a new probable scenario whereby Neandertal and Modern Human populations
in Iberia did not co-exist and Middle Paleolithic sites do not occur after 42 ka cal BP. This is a
relevant proposal, since it contradicts decades of acceptance of the late Neandertal survival
hypothesis as the paradigmatic model.
However, the hypothesis of a not-so-late Neandertal population breakdown south of the
Ebro basin has already received some criticism [26±27]. Both in the Mediterranean and
Atlantic southern coasts of Iberia, some sites still suggest a post-42 ka cal BP chronology for the last
Neandertal presence at the peninsula. Gorham's cave (Gibraltar) [
], Oliveira (Portugal)
, Carihuela (Granada) [
], Sima de las Palomas (Murcia)  and Cueva AntoÂn
(Murcia)  provide both chronometric and paleoecological data suggesting a persistence of
Mousterian contexts after 42 ka cal BP. Despite the cases of Gorham and Carihuela have
received strong criticism [6, 23, 131], dates obtained for Oliveira, Cueva AntoÂn and Sima de
las Palomas, although not without problems , remain unchallenged by means of new
chronometric results. If these late survival cases are accepted, it would imply that Neandertals were
present in the southern Iberian coasts at least until c. 37 ka cal BP, correlating with Greenland
Interstadial 8. Since this chronology contradicts current trend suggested by the last
chronometric investigations, research on this topic should be kept in the realm of hypothesis and
theory building for now.
Considering the Iberian interior territories, the strongest evidences supporting a late
Neandertal survival have been unquestionably refuted. At La Ermita cave (Burgos), dates obtained
by Aminoacid Racemization and Uranium/Thorium techniques have reassigned level 5a to
MIS 5 , previously radiocarbon dated in the range of c. 36.6±34.7 ka cal BP . At
Jarama VI rockshelter (Guadalajara), the latest Mousterian occupation, previously radiocarbon
dated between c. 41 and 30 ka cal BP , have been re-dated by new chronometric analyses,
including radiocarbon measurements of bone samples pre-treated with ultrafiltration  and
luminescence dating (post-IR IRSL) of associated sediments , to between c. 60 and 50 ka cal
BP. Other interior Middle Paleolithic sites having yielded reliable chronometric dates within
MIS 3 are Abrigo del Molino [134, 135], Prado Vargas , Hotel California ,
Valdegoba , Peña Cabra , La Mina  and Hundidero [
]. Since none of these sites
have provided any date younger than 42 ka cal BP (Fig 29), the hypothesis of a not-so-late
breakdown of Neandertal populations in Iberia remains unchallenged in the interior regions
of the peninsula.
43 / 54
Fig 29. Middle Paleolithic sites in interior Iberia dated to MIS 3. Sites having yielded reliable chronometric dates are shown in black.
Sites with uncertain results are numbered in red. For complete dating results and methods see [6, 9, 30, 132, 134±141, 145, 146].
Radiocarbon dates were calibrated using OxCal 4.3  and IntCal13 . OSL: Optically Stimulated Luminiscence. AAR: Aminoacid
Racemization. AMS: Accelerator mass spectrometry.
However, an important shortcoming faced by any study dealing with population dynamics
in the Late Pleistocene of interior Iberia is the poor quantity and quality of the
geoarcheological, paleoenvironmental and chronometric data available. This issue has been acknowledged in
recent chronometric research [6, 9], and is most probably due to (1) a lack of research projects
in interior Iberia compared to the coastal regions, and (2) the difficulties of locating open-air
sites, potentially much more common than cave archives in the Spanish Meseta [16, 142±144].
In fact, there are three sites that could still suggest a post-42 ka cal BP chronology for Middle
Paleolithic contexts in the Spanish plateau. In Cueva MillaÂn (Burgos), two radiocarbon dates
on bone obtained in the 1980's ranged from c. 41 to 43 ka cal BP [
]. However, these
measurements were obtained by the conventional radiocarbon method, and hence a new
chronometric program is required before the proposed dates can be considered to be reliable. In
the Madrid basin, open-air sites of 12 de Octubre and Cañaveral-AÂ rea 3 have produced
luminescence dates younger than 40 ka BP. In the 12 de Octubre deposit, a typical Mousterian
assemblage is associated to a series of OSL dates between 40 and 33 ka BP. However, the
excavators of this site cast doubt on these results suggesting that the proposed dates, which
contradict geomorphological data, are most probably underestimates [
]. As for Cañaveral-AÂ rea 3,
44 / 54
a TL date of 33 + 4.0/-3.5 ka BP was obtained at the top of a layer containing Levallois
]. However, in addition to the high standard deviation of this measurement, and
the fact that the date must be considered a terminus ante quem for human activity, a full
discussion of methods and results of chronometric research conducted at this site is still to be
In short, although some uncertainties must be acknowledged when dealing with the Iberian
interior territories, no strong chronometric evidence supporting a post-42 ka cal BP survival
can be currently attested in them. In fact, Neandertals occupying the deep interior of Los
Casares cave at c. 44.9±42.2 ka cal BP, must be considered among the last of their kind living in
the interior lands of the Iberian Peninsula prior to their final disappearance (Fig 29). This
evidence does not support a late survival of Neandertals in the Iberian interior, but rather suggests
a not-so-late disappearance of this human group from these territories, roughly coincident
with the proposed chronology for this process in northern Iberia [
6, 8, 125
However, although limited to a single flake and 52 faunal remains with no signs of human
action, evidence gathered from layer b of Los Casares-Seno A must be also considered in this
discussion. Paleoenvironmental data recorded in this layer show a cold and arid environment
most probably correlating with GS 11, GS 10 or H4 (c. 42±38 ka years ago), thus suggesting a
possible late and scarce presence of Neandertals at the cave. Since this could be interpreted as
reflecting the near-abandonment of this high area (>1,000 m asl) of the Iberian interior due to
climatic deterioration, it could even be hypothesized that this layer also correlates with the
final disappearance of this human species from inland Iberia due to climatic stress. Yet, since
this is a hypothesis based on scarce empirical evidence, it cannot be used to support a late
survival of Neandertals in interior Iberia. Given the still poor record gathered at this layer, and in
general the scarce data available for discussing human-environment interactions in the Iberian
interior during the Late Pleistocene, these reflections should be taken as working hypotheses
to be tested with future research.
In any case, independent of whether layer b represents a late Neandertal presence at Los
Casares or just an arid and cold episode devoid of human occupation (a question that remains
open given that only one flake was recorded at this layer), the current record in interior Iberia
shows a pattern in which little or no evidence for a Middle Paleolithic presence is registered
after 42 ka cal BP. If we accept the late persistence of Neandertals in at least some of the
southern coastal sites that are currently claimed to reflect Middle Paleolithic occupations until at
least c. 37 ka years ago, a parsimonious corollary is that populations living in the highlands of
the Spanish Meseta abandoned these potentially risky environments [
] and moved to the
coastal areas of southern Iberia during some of the cold stadials following GI 11. The exact
timing of this potential population movement is a question that needs further research.
Notwithstanding, since no Upper Paleolithic occupations have been attested in inland Iberia until
c. 25.5 ka cal BP [
], no action by Modern Humans could be invoked as triggering or even
affecting this process. The breakdown of Neandertal populations in the Iberian interior is best
explained as an abandonment of the area due to climatic deterioration or some other internal
factor. This suggest that climate change could have been an important factor contributing to
the final demise of the Neandertals [147, 149±152].
Los Casares cave is a classic site for the study of the Middle Paleolithic settlement of inland
Iberia. Despite its relevance in the last quarter of the 20th century, data on this site was of little use
for current research due to a prolonged period of scientific inactivity. New stratigraphic,
micromorphological, chronometric, paleoenvironmental, archeozoological and technological
45 / 54
data provided in this study have changed this situation. Los Casares cave has emerged as a
relevant multi-proxy archive for studying human-environment interactions and population
dynamics at the end of the Middle Paleolithic in the Iberian interior. Evidence discussed in
this paper supports a breakdown of the Neandertal settlement system in inland Iberia around
42 ka cal BP or slightly later, and suggests that this could be related to an abandonment of the
interior highlands of the Meseta due to climate deterioration. The last Neandertals of Iberia
are thus only found in the southern coastal areas of the peninsula, where a post-42 ka cal BP
survival of Middle Paleolithic contexts has not been falsified. Although evidence discussed in
this paper represents a significant advance on these topics, the geoarcheological,
paleoecological and chronometric record in the Iberian interior are still too weak to allow for theory
building at the regional level, despite significant progress in the recent past. It is our contention that
further fieldwork on the under-investigated interior regions of the Iberian Peninsula will
substantially change±again±models on population dynamics in Iberia and southwest Europe
during this critical period of human prehistory. Until then, unbiased data gathering and
hypothesis testing remain crucial.
On epistemic grounds, far from the classic Kuhnian scenario of rapid and definitive
paradigmatic shift±which is rarely verified±, it is our contention that the current scientific
situation on the problem of Neandertal disappearance in Iberia should be best considered as a
not-so-fast process of data accumulation and hypotheses proposal that should eventually lead
to a new big picture on the issue. Whether this picture will be totally different to previously
accepted one, slightly different, or even in consonance, is a question that remains open
despite great advances in the last years. Only more fieldwork (including excavation of new
sites), data gathering (not only chronometric, but also stratigraphic, paleoenvironmental and
archeological), and problem-oriented research, will eventually lead to still not definitive,
but increasingly better, scientific answers. Ongoing investigations in a handful of sites in
the interior regions of Iberia, albeit limited to the foothills of the Central System range, the
Madrid basin and the Atapuerca area, will hopefully contribute to that end [
135, 139, 144,
S1 Appendix. Supporting tables on micromorphology and archeozoology and taphonomy.
The staff from the Dirección General de Cultura de la Junta de Comunidades de Castilla-La
Mancha and the Museo Comarcal de Molina de Aragón greatly facilitated our fieldwork in Los
Casares cave. We especially acknowledge friendly and efficient work of Juan Manuel
Monasterio, JoseÂ Antonio MartÂõnez and Marca Perruca. Dr. Carmen Cacho Quesada provided access
to the lithic and faunal assemblages curated at the Museo Arqueológico Nacional (Madrid). Drs.
Ignacio MartÂõnez, AdriaÂn Pablos and Nohemi Sala checked the bone assemblages in search of
human remains. Dr. Ignacio BarandiaraÂn gave support to our work. Dr. Ariane Burke kindly
revised some parts of the text. We gratefully acknowledge contribution made by the excavation
and laboratory team of Los Casares cave, mainly composed of students from the University of
AlcalaÂ. Photos B and C of Fig 4 were authored by Alfonso DaÂvila. Adara LoÂpez-LoÂpez
contributed with the 3D view of Fig 1C. We thank two anonymous reviewers and the editor for their
46 / 54
Conceptualization: Manuel Alcaraz-Castaño, Javier Alcolea-GonzaÂlez, Rodrigo de
Behrmann, Gerd-Christian Weniger.
Formal analysis: Martin Kehl.
Funding acquisition: Manuel Alcaraz-Castaño, Martin Kehl, Javier Baena-Preysler,
Antonio LoÂpez-SaÂez, Gerd-Christian Weniger.
Investigation: Manuel Alcaraz-Castaño, Javier Alcolea-GonzaÂlez, Martin Kehl, Rosa-MarÂõa
Albert, Javier Baena-Preysler, Rodrigo de BalbÂõn-Behrmann, Felipe Cuartero, Gloria
Cuenca-BescoÂs, JoseÂ-Antonio LoÂpez-SaÂez, Raquel PiqueÂ, David RodrÂõguez-AntoÂn, JoseÂ
Yravedra, Gerd-Christian Weniger.
Methodology: Manuel Alcaraz-Castaño, Javier Alcolea-GonzaÂlez, Martin Kehl, Rosa-MarÂõa
Albert, Felipe Cuartero, Gloria Cuenca-BescoÂs, Fernando JimeÂnez-Barredo, JoseÂ-Antonio
LoÂpez-SaÂez, Raquel PiqueÂ, David RodrÂõguez-AntoÂn, JoseÂ Yravedra.
Project administration: Manuel Alcaraz-Castaño, Javier Alcolea-GonzaÂlez, Gerd-Christian
Resources: Javier Baena-Preysler.
Writing ± original draft: Manuel Alcaraz-Castaño, Javier Alcolea-GonzaÂlez, Martin Kehl,
Rosa-MarÂõa Albert, Gloria Cuenca-BescoÂs, JoseÂ-Antonio LoÂpez-SaÂez, Raquel PiqueÂ, David
RodrÂõguez-AntoÂn, JoseÂ Yravedra.
Writing ± review & editing: Manuel Alcaraz-Castaño, Javier Alcolea-GonzaÂlez, Martin Kehl,
Rosa-MarÂõa Albert, Javier Baena-Preysler, Rodrigo de BalbÂõn-Behrmann, Felipe Cuartero,
Gloria Cuenca-BescoÂs, Fernando JimeÂnez-Barredo, JoseÂ-Antonio LoÂpez-SaÂez, Raquel
PiqueÂ, David RodrÂõguez-AntoÂn, JoseÂ Yravedra, Gerd-Christian Weniger.
47 / 54
9. Kehl M, Burow C, Hilgers A, Navazo M, Pastoors A, Weniger GC et al. Late Neanderthals at Jarama
VI (central Iberia)? Quat. Res. 2013; 80: 218±234.
10. Puig y Larraz C. Cavernas y simas de España. BoletÂõn de la ComisioÂn del Mapa GeoloÂgico de España.
Tomo II, segunda serie. Madrid; 1894.
11. CabreÂ J. Las cuevas de los Casares y de la Hoz. Archivo Español de Arte y ArqueologÂõa 1934; X:
12. CabreÂ J. Figuras antropomorfas en la cueva de Los Casares (Guadalajara). Archivo Español de
ArqueologÂõa 1940; XIV (41): 81±96.
16. BalbÂõn-Behrmann R, Alcolea-GonzaÂlez JJ. La grotte de Los Casares et lÂArt PaleÂolithique de la Meseta
espagnole. LÂAnthropologie. 1992; 96 (2±3): 397±452.
21. Alcaraz-Castaño M, Weniger GC, Alcolea JJ, Kehl M, Baena J, Yravedra J et al. DinaÂmicas
poblacionales en el centro de la PenÂõnsula IbeÂrica durante el Pleistoceno superior: un nuevo proyecto
geoarqueoloÂgico. In: Galve JP, AzañoÂn JM, PeÂrez JV. Ruano P., editors. Una visioÂn global del Cuaternario.
El hombre como condicionante de procesos geoloÂgicos. XIV ReunioÂn Nacional de Cuaternario,
Granada (España). pp. 42±45.
22. Alcaraz-Castaño M, Weniger GC. Testing population hiatuses in the Late Pleistocene of Central Iberia:
a geoarchaeological approach. In Hugo Obermaier-Gesellschaft fuÈr Erforschung des Eiszeitalters und
der Steinzeit e.V. 57th Annual Meeting in Heidenheim, 7th± 11th of April, 2015. Erlangen: Hugo
Obermaier-Gesellschaft; 2015. pp.: 14±15.
23. Zilhão J. Chronostratigraphy of the Middle-to-Upper Paleolithic Transition in the Iberian Peninsula.
Pyrenae. 2006; 37: 7±84.
48 / 54
31. Zilhão J, Ajas A, Badal E, Burow C, Kehl M, LoÂpez-SaÂez JA et al. Cueva AntoÂn: A multi-proxy MIS 3 to
MIS 5a paleoenvironmental record for SE Iberia. Quat Sci Rev. 2016; 146: 251±273. http://dx.doi.org/
33. Stoops G. Guidelines for the Analysis and Description of Soil and Regolith Thin Sections. Madison
(WI): Soil Science Society of America; 2003.
34. Hoffmann DL, Prytulak J, Richards DA, Elliott TR, Coath CD, Smart PL et al. Procedures for accurate
U and Th isotope measurements by high precision MC-ICPMS. International Journal of Mass
Spectrometry. 2007; 264: 97±109.
35. Hoffmann DL. 230Th isotope measurements of femtogram quantities for U-series dating using multi ion
counting (MIC) MC-ICPMS. International Journal of Mass Spectrometry. 2008; 275: 75±79.
36. Ivanovich M, Harmon RS. Uranium-series disequilibrium: applications to Earth, Marine and
Environmental Sciences. Oxford: Oxford University Press; 1992.
37. Scholz D, Hoffmann D. 230Th/U-datind of fossil corals and speleothems. Quaternary Science Journal.
2008; 57 (1±2): 52±76.
38. Cheng H, Lawrence R, Shen CC, Polyak VJ, Asmerom Y, Woodhead J et al. Improvements in 230Th
dating, 230Th and 234U half-life values, and U-Th isotopic measurements by multi-collector inductively
coupled plasma mass spectrometry. Earth Planet Sci Lett. 2013; 371±372: 82±91.
39. Rethemeyer J, FuÈloÈp RH, HoÈfle S, Wacker L, Heinze S, Hajdas I et al. Status report on sample
preparation facilities for 14C analysis at the new CologneAMS center. Nucl Instrum Methods Phys Res B.
2013; 294: 168±72.
40. Bronk Ramsey C, Lee S. Recent and planned developments of the program OxCal. Radiocarbon.
2013; 55: 720±730.
41. Reimer PJ, Bard E, Bayliss A, Beck JW, Blackwell PG, Bronk Ramsey C et al. IntCal13 and
MARINE13 radiocarbon age calibration curves 0±50000 years calBP. Radiocarbon. 2013; 55:
42. Burjachs F, LoÂpez-SaÂez JA, Iriarte MJ. MetodologÂõa ArqueopalinoloÂgica. In: BuxoÂ R, PiqueÂ R, editors.
La recogida de muestras en ArqueobotaÂnica: objetivos y propuestas metodoloÂgicas. La gestioÂn de los
recursos vegetales y la transformacioÂn del paleopaisaje en el MediterraÂneo occidental. Barcelona:
Museu d'Arqueologia de Catalunya; 2003. pp. 11±18
43. LoÂpez-SaÂez JA, LoÂpez-GarcÂõa P, Burjachs F. ArqueopalinologÂõa: SÂõntesis crÂõtica. Polen. 2003; 12:
44. Moore PD, Webb JA, Collinson ME. Pollen analysis. 2nd ed. Oxford: Blackwell Scientific; 1991.
45. Reille M. Pollen et spores d'Europe et d'Afrique Nord. Marseille: Laboratoire de Botanique historique
et Palynologie, EÂditions CNRS; 1992.
46. Desprat S, DÂõaz-FernaÂndez P, Coulon T, Ezzat L, Pessarossi-Langlois J, Gil. Pinus nigra (Spanish
black pine) as the dominant species of the last glacial pinewoods in south-western to central Iberia: a
morphological study of modern and fossil pollen. J Biogeog. 2015; 42: 1998±2009.
47. Grimm EC. TILIA: a program for analysis and display. 1st ed. Springfield: Illinois State Museum;
48. Grimm EC. Coniss: a Fortran 77 program for stratigraphically constrained cluster analysis by the
method of incremental sum of squares. Computer & Geosciences. 1987; 13: 13±35.
49. ter Braak CJF, SÏ milauer P. CANOCO Reference manual and Canodraw for Windows user's guide:
software for canonical community ordination (version 4.5.). NY, USA: Biometris; 2002.
50. LoÂpez-GarcÂõa JM. Los micromamÂõferos del Pleistoceno Superior en la PenÂõnsula IbeÂrica. EvoluioÂn de
la diversidad taxonoÂmica y cambios paleoambientales y paleoclimaÂticos. 1st ed. Editorial AcadeÂmica
51. SeseÂ C, Rubio-Jara, Panera J, PeÂrez-GonzaÂlez A. MicromamÂõferos del Pleistoceno Superior del
yacimiento de PRERESA en el valle del Manzanares y su contribucioÂn a la reconstruccioÂn paleoambiental
de la cuenca de Madrid durante el Pleistoceno. Estudios GeoloÂgicos. 2011; 67 (2): 471±494. http://dx.
52. Laplana C, Blain HA, Sevilla P, Arsuaga JL, Baquedano E, PeÂrez-GonzaÂlez A. Un assemblage de
petits verteÂbreÂs hautement diversifieÂ de la fin du MIS5 dans un environnement montagnard au centre
49 / 54
de l'Espagne (Cueva del Camino, Pinilla del Valle, CommunauteÂ Autonome de Madrid). Quaternaire.
2013; 24 (2): 207±216. http://dx.doi.org/10.4000/quaternaire.6617
53. LoÂpez-GarcÂõa JM, Cuenca-BescoÂs G. EÂ volution climatique durant le PleÂistocène SupeÂrieur en
Catalogne (Nord-est de l'Espagne) d'après l'eÂtude des micromammifères. Quaternaire. 2010; 21:
54. LoÂpez-GarcÂõa JM, Blain HA, Bennàsar M, FernaÂndez-GarcÂõa M. Environmental and climatic context of
Neanderthal occupation in southwestern Europe during MIS3 inferred from the small-vertebrate
assemblages. Quat Int. 2014: 326±327: 319±328.
56. Madella M, Powers-Jones AH, Jones MK. A Simple Method of Extraction of Opal Phytoliths from
Sediments Using a Non-Toxic Heavy Liquid. J Archaeol Sci. 1998; 25: 801±803.
57. Albert RM, Tsatskin A, Ronen A, Lavi O, Estroff L, Lev-Yadun S et al. Mode of occupation of Tabun
Cave, Mt. Carmel Israel, during the Mousterian period: A study of the sediments and the phytoliths. J
Archaeol Sci. 1999; 26: 1249±1260.
58. Madella M, Alexandre A, Ball T. International code for phytolith nomenclature 1.0. Ann Bot. 2005; 96:
253±260 https://doi.org/10.1093/aob/mci172 PMID: 15944178
59. Piperno DR. Phytoliths: a comprehensive guide for archaeologists and paleoecologists Lanham, MD:
AltaMira Press; 2006.
60. Barboni D, Bremond L. Phytoliths of East African grasses: An assessment of their environmental and
taxonomic significance based on floristic data. Rev Palaeobot Palynol. 2009; 158: 29±41.
61. Tsartsidou G, Lev-Yadun S, Albert RM, Miller-Rosen A, Efstratiou N, Weiner S. The Phytolith
Archaeological Record: Strengths and Weaknesses Based on a Quantitative Modern Reference Collection
from Greece. J Archaeol Sci. 2007; 34 (8): 1262±1275.
62. Ball TA, Davis AL, Evett RR, Ladwig JL, Tromp M, Out WA, et al. Morphometric analysis of phytoliths:
recommendations towards standardization from the International Committee for Phytolith
Morphometrics. J Archaeol Sci. 2016; 68: 106±111.
63. Albert RM, RuÂõz JA, Sans A. PhytCore ODB: A new tool to improve efficiency in the management and
exchange of information on phytoliths. J Archaeol Sci. 2016; 68: 98±105.
64. Carnelli AL, Theurillat JP, Madella M. Phytoliths types and type-frequencies in subalpine-alpin plant
species of the European Alps. Rev Palaeobot Palynol. 2004; 129: 39±65.
65. Esteban I, Albert RM, Zilhão J, Villaverde V. Neanderthal use of plants and past vegetation
reconstruction at the Middle Paleolithic site of Abrigo de la Quebrada (Chelva, Valencia, Spain). Archaeol
Anthropol Sci. 2015; 9: 265±278.
66. Albert RM, Bamford MK, Cabanes D. Taphonomy of phytoliths and macroplants in different soils from
Olduvai Gorge (Tanzania) and the application to Plio-Pleistocene palaeoanthropological samples.
Quat Int. 2006; 148: 78±94.
67. Madella M, Lancelotti C.Taphonomy and phytoliths: A user manual. Quat Int. 2012; 275: 76±83.
68. Altuna J. Fauna de MamÂõferos del yacimiento prehistoÂrico de Los Casares (Guadalajara). In:
BarandiaraÂn I, editor. La cueva de Los Casares (en Riba de Saelices, Guadalajara). Excavaciones
ArqueoloÂgicas en España 76. Madrid: Ministertio de EducacioÂn y Ciencia; 1973. pp. 97±116.
69. Lyman RL. Relative abundance of skeletal specimens and taphonomic analysis of vertebrate remains.
Palaios. 1994; 9: 288±298.
70. Brain CK. The contribution of Namib Desert Hottentot to understanding of Australopithecus bone
accumulations. Scientific Papers in Namibian Desert Research Station. 1969; 32: 1±11.
71. Barba R, DomÂõnguez-Rodrigo M. The Taphonomic Relevance of the Analysis of Bovid Long Limb
Bone Shaft Features and Their Application to Element Identification. Study of Bone Thickness and
Morphology of the Medullar Cavity. Journal of Taphonomy. 2005; 3: 29±42.
72. Yravedra J, DomÂõnguez-Rodrigo M. The shaft-based methodological approach to the quantification of
long limb bones and its relevance to understanding hominid subsistence in the Pleistocene: application
to four Palaeolithic sites. J Quat Sci. 2009; 24: 85±96.
73. Steele TE. Red deer: their ecology and how they were hunted by Late Pleistocene hominids in western
Europe. Ph.D. dissertation, Stanford University, California, 2002.
74. PeÂrez-Ripoll M. Estudio de la secuencia del desgaste de los molares de Capra pyrenaica de los
yacimientos prehistoÂricos. Archivo de Prehistoria Levantina. 1988; 18: 83±128.
75. Levine M A. The use of crown height measurements and eruption-wear sequence to age horse teeth.
In Wilson B, Grigson C, Payne S, editors. Aging and sexing from archaeological sites. Oxford: BAR
British Series 109;1982, pp. 223±250.
50 / 54
76. Blumenschine R. Percussion marks, tooth marks and the experimental determinations of the timing of
hominid and carnivore access to long bones at FLK Zinjanthropus, Olduvai Gorge, Tanzania. J Hum
Evol. 1995; 29: 21±51.
78. Potts R, Shipman P. Cut marks made by stone tools from Olduvai Gorge, Tanzania. Nature. 1981;
79. Blumenschine R, Selvaggio MM. Percussion marks on bone surfaces as a new diagnostic of hominid
behaviour. Nature. 1988; 333: 763±765.
80. DomÂõnguez-Rodrigo M. Meat eating by early homids at FLK Zinj 22 Site, Olduvay Gorge Tanzania: An
experimental approach using cut-mark data. Journal of Human Evolution. 1997; 33: 669±690. https://
doi.org/10.1006/jhev.1997.0161 PMID: 9467775
81. DomÂõnguez-Rodrigo M, Barba R. A study of cut marks on small-sized carcasses and its application to
the study of cut marked bones from small mammals at the FLK Zinj site. Journal of Taphonomy. 2005;
82. Behrensmeyer AK. Taphonomic and ecologic information from bone weathering. Paleobiology. 1978;
83. Parson MK, Brett E. Taphonomic processes and biases in modern marine environments: an actualistic
perspective on fossil assemblage preservation. In Donovan SK, editor. The processes of fossilization.
1st ed. New York: Columbia University Press; 1989. pp. 22±65.
84. CaÂceres I. TafonomÂõa de yacimientos antroÂpicos en karst. Complejo GalerÂõa (Sierra de Atapuerca,
Burgos), Vanguard Cave (Gibraltar) y Abric RomanÂõ (Capalledes, Barcelona). Ph.D. dissertation,
Universidad Rovira i Virgili, 2002.
85. Yravedra J. TafonomÂõa aplicada a ZooarqueologÂõa. Aula Abierta-UNED, 2006.
86. Thompson CE. Ball S, Thompson TJU, Gowland R. The abrasion of modern and archaeological bones
by mobile sediments: the importance of transport modes. J Arch Sci. 2011; 38: 784±793.
87. DomÂõnguez-Rodrigo M, Barba R. New estimates of tooth mark and percussion mark frequencies at the
FLK ZINJ site: The carnivore-hominid-carnivore hypothesis falsified. J Hum Evol. 2006; 50: 170±194
https://doi.org/10.1016/j.jhevol.2005.09.005 PMID: 16413934
88. Villa P, Mahieu E. Breakage patterns of human long bones. J Hum Evol. 1991; 21: 27±48.
89. Inizan ML, Reduron-Ballinger M, Roche H, Tixier J. Technology and Terminology of Knapped Stone.
PreÂhistoire de la Pierre TailleÂe, tome 5. Nanterre: CREP; 1999.
90. Bar-Yosef O, Van Peer P. The ChaÃõne OpeÂratoire Approach in Middle Paleolithic Archaeology. Curr
Anthropol. 2009; 50 (1): 103±131.
91. Amick DS. Reflection on the origins of recycling: a Paleolithic perspective. Lithic Technology. 2014;
39 (1): 64±69.
92. Gibert L, Scott GR, Scholz D, Budsky A, Ferràndez C, Ribot F et al. Chronology for the Cueva Victoria
fossil site (SE Spain): Evidence for Early Pleistocene Afro-Iberian dispersals. J Hum Evol. 2016; 90:
183±197. https://doi.org/10.1016/j.jhevol.2015.08.002 PMID: 26581114
93. Cuenca-BescoÂs G. AnaÂlisis filogeneÂtico de Allocricetus del Pleistoceno (Cricetidae, Rodentia,
Mammalia). Coloquios de PaleontologÂõa. 2003; Extra (1): 95±113.
94. Cuenca-BescoÂs G, Straus LG, GonzaÂlez-Morales M, GarcÂõa-Pimienta JC. The reconstruction of past
environments through small mammals: from the Mousterian to Bronze Age in El MiroÂn cave. J
Archaeol Sci. 2009; 36: 947±955.
95. Twiss PC. Predicted World Distribution of C3 and C4 grass phytoliths. In: Rapp GJR, Mulholland SE,
editors. Phytolith systematics. Emerging issues. Advances in Archaeological and Museum Science,
Vol. 1. London: Plenum Press; 1992. pp. 113±128.
96. Albert RM, Weiner S. Study of phytoliths in prehistoric ash layers using a quantitative approach, In:
Meunier JD, Coline F, editors. Phytoliths: Applications in Earth Sciences and Human History, Lisse:
A.A. Balkema Publishers; 2001. pp. 251±266.
97. Carnelli AL, Madella M, Theurillat JP. Biogenic silica production in selected alpine plants species and
plant communities. Ann. Bot. 2001; 87: 425±434.
98. LoÂpez-SaÂez JA, Alba-SaÂnchez F, SaÂnchez-Mata D, Abel-Schaad D, GavilaÂn RG, PeÂrez-DÂõaz S. A
palynological approach to the study of Quercus pyrenaica forest communities in the Spanish Central
System. Phytocoenologia. 2015; 45: 107±124.
99. CarrioÂn JS, Scott L, Arribas A, Fuentes N, Gil-Romera G, Montoya E. Pleistocene landscapes in
central Iberia inferred from pollen analysis of hyena coprolites. J Quat Sci. 2007; 22: 191±202.
51 / 54
100. Torre I, Albert RM, AllueÂ E, AÂ lvarez-FernaÂndez E, Aparicio MT, Arroyo A et al. Chronological and
palaeoenvironmental context of human occupations at the BuendÂõa rockshelter (Central Spain) during
the late Upper Pleistocene in inland Iberia. J Quat Sci. 2015; 30: 376±390.
101. Cuenca-BescoÂs G, LoÂpez-GarcÂõa JM, Galindo-Pellicena MA, GarcÂõa-Perea R, Gisbert J, Rofes J et al.
The pleistocene history of Iberomys, an endangered endemic rodent from South Western Europe.
Integrative Zoology. 2014; 9, 481±497. https://doi.org/10.1111/1749-4877.12053 PMID: 25236417
102. LoÂpez-GarcÂõa JM, Blain HA, Cuenca-BescoÂs G, Arsuaga JL. Chronological, environmental and
climatic precisions on the Neanderthal site of the Cova del Gegant (Sitges, Barcelona, Spain). J Hum
Evol. 2008; 55: 1151±1155. https://doi.org/10.1016/j.jhevol.2008.08.001 PMID: 18789810
103. Uzquiano P. El registro antracoloÂgico durante la transicioÂn Musteriense-PaleolÂõtico Superior Inicial en
la RegioÂn CantaÂbrica: VegetacioÂn, paleoambiente y modos de vida alrededor del fuego. Museo de
Altamira. MonografÂõas. 2005; 20: 255±274.
104. Badal E, Villaverde V, Zilhão J. Middle Paleolithic wood charcoal from three southern Iberian sites:
biogeographic implications. Saguntum. 2012; Extra 13: 13±24.
105. Sala MTH, Arsuaga JL, Laplana C, Ruiz-Zapata B, Gil-GarcÂõa MJ, GarcÂõa N et al. Un paisaje de la
Meseta durante el Pleistoceno Superior. Aspectos paleontoloÂgicos de la Cueva de la Zarzamora
(Segovia, España). Bol R Soc Esp Hist Nat Secc Geol. 2011; 105: 67±85.
106. Arsuaga JL, Baquedano E, PeÂrez-GonzaÂlez A, Sala N, Quam, RodrÂõguez L et al. Understanding the
ancient habitats of the last-interglacial (late MIS 5) Neanderthals of central Iberia: Paleoenvironmental
and taphonomic evidence from the Cueva del Camino (Spain) site. Quat Int. 2012; 275; 55±75.
107. Vega-Toscano LG, Hoyos M, Ruiz-Bustos A, Laville H. La seÂquence de la grotte de la Carihuela (Piña,
Grenade): Chronostratigraphie et paleÂoeÂcologie du PleÂistocene SupeÂrieur du Sud de la PeÂninsule
IbeÂrique. In Otte M, editor, L'Homme de Neandertal. Vol 2. L'environnement. Liège: Univ. de Liège;
1988. pp. 169±180.
108. Rasmussen SO, Bigler M, Blockley SP, Blunier T, Buchardt SL, Clausen HB et al. 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.
2014; 106: 14±28.
109. LoÂpez-SaÂez JA, Alba-SaÂnchez F, LoÂpez-Merino L, PeÂrez-DÂõaz S. Modern pollen analysis: a reliable
tool for discriminating Quercus rotundifolia communities in Central Spain. Phytocoenologia. 2010; 40:
110. LoÂpez-SaÂez JA, SaÂnchez-Mata D, Alba-SaÂnchez F, Abel-Schaad D, GavilaÂn RG, PeÂrez-DÂõaz S.
Discrimination of Scots pine forests in the Iberian Central System (Pinus sylvestris var. iberica) by means
of pollen analysis. Phytosociological considerations. Lazaroa. 2013; 34: 191±208.
111. Andersen KK, Svensson A, Johnsen S, Rasmussen SO, Bigler M, RoÈthlisberger R et al. The
Greenland ice core chronology 2005, 15±42 ka. Part 1: Constructing the time scale. Quat Scie Rev. 2006;
25 (23±24): 3246±3256.
112. Svensson A, Andersen KK, Bigler M, Clausen HB, Dahl-Jensen D, Davies SM et al. The Greenland ice
core chronology 2005, 15±42 ka. Part 2: Comparison to other records. Quat Scie Rev. 2006;
25 (23±24): 3258±3267.
113. Cuenca-Bescos G, Straus LG, Garcia-Pimienta JC, GonzaÂlez-Morales MR, Lopez-Garcia JM. Late
Quaternary small mammal turnover in the Cantabrian Region: The extinction of Pliomys lenki
(Rodentia, Mammalia). Quat Int. 2010; 212 (2): 129±136.
114. Guillem P. PaleontologÂõa continental: microfauna. In. RosselloÂ i Verger. Editor, El Cuaternario del PaÂõs
Valenciano. Valencia: AEQUA & Universitat de Valencia; 1995. pp. 227±233.
115. Ruiz-Bustos A, GarcÂõa-SaÂnchez M. Las condiciones ecoloÂgicas del Musteriense en las depresiones
granadinas. La fauna de micromamÂõferos en la cueva de La CariguÈela (Piñar, Granada). Cuadernos
de Prehistoria de la Universidad de Granada. 1997; 2: 7±1.
116. SaÂnchez-Goñi MF, Harrison SP. Millennial-scale climate variability and vegetation changes during the
Last Glacial: Concepts and terminology. Quat Sci Rev. 2010; 29 (21±22): 2823±2827.
117. Vega Toscano LG. El traÂnsito del PaleolÂõtico Medio al PaleolÂõtico Superior en el Sur de la PenÂõnsula
IbeÂrica. In Cabrera V, editor. El origen del hombre moderno en el suroeste de Europa. Madrid:
UNED; 1993. pp. 147±170.
118. Villaverde V. Fumanal MP. Relations entre le PaleÂolithique Moyen et le PaleÂolithique SupeÂrieur dans
le versant meÂditerraneÂen espagnol. In Farizy C, editor. Paleolithique moyen recent et Paleolithique
superieur ancien en Europe. Nemours: MeÂmoires du MuseÂe de PreÂhistoire d'Ile-de-France 3;
1990. pp. 177±183.
119. Iturbe G, Fumanal M, CarrioÂn J, Cortell E, MartÂõnez R, Guillem P et al. Cova Beneito (Muro, Alicante):
una perspectiva interdisciplinar. Recerques del Museo d'Alcoi. 1990; 2: 23±88.
52 / 54
120. Straus LG, Bischoff J, Carbonell E. A review of the Middle to Upper Paleolithic transition in Iberia.
PreÂhistoire EuropeÂenne. 1993; 3: 11±27.
121. Zilhão J. Le passage du PaleÂolithique moyen au PaleÂolithique supeÂrieur dans le Portugal. In Cabrera
V, editor. El origen del hombre moderno en el suroeste de Europa. Madrid: UNED; 1993. pp.
122. Hublin JJ, Spoor F, Braun M, Zonneveld F, Condemi S. A late Neanderthal associated with Upper
Palaeolithic artefacts. Nature. 1996; 381: 224±226. https://doi.org/10.1038/381224a0 PMID: 8622762
123. Raposo L, Cardoso J. Las industrias lÂõticas de la Gruta Nova de Columbeira en el contexto del
Musteriense final de la PenÂõnsula IbeÂrica. Trabajos de Prehistoria. 1998; 55: 39±62.
124. JordaÂ J. Radiocarbono y cronologÂõa del Jarama (Sistema Central, España) durante el Pleistoceno
Superior y Holoceno. In Mata E, editor. Cuaternario y ArqueologÂõa. Homenaje a Francisco Giles
Pacheco. CaÂdiz: DiputacioÂn Provincial de CaÂdiz; 2010. pp. 101±110.
125. Wood R, Arrizabalaga A, Camps M, Fallon S, Iriarte-Chiappuso MJ, Jones J et al. The chronology of
the earliest Upper Palaeolithic in northern Iberia: New insights from L'Arbreda, Labeko Koba and La
Viña. J Hum Evol. 2014; 69: 91±109. http://dx.doi.org/10.1016/j.jhevol.2013.12.017 PMID: 24636733
126. JoÈris O, AÂ lvarez-FernaÂndez A, Weninger B. Radiocarbon evidence of the Middle to Upper Palaeolithic
transition in Southwestern Europe. Trabajos de Prehistoria. 2003; 60 (2):15±38.
127. Vaquero M. El traÂnsito PaleolÂõtico Medio/Superior en la PenÂõnsula IbeÂrica y la Frontera del Ebro.
Comentario a Zilhão. Pyrenae. 2006; 37(2): 107±129.
128. JoÈris O, Street M. At the end of the 14C time scaleÐThe Middle to Upper Paleolithic record of western
Eurasia. J Hum Evol. 2008; 55 (5):782±802. https://doi.org/10.1016/j.jhevol.2008.04.002 PMID:
129. FernaÂndez S, Fuentes N, CarrioÂn J, GonzaÂlez-SampeÂriz P, Montoya E, Gil G et al. The Holocene and
Upper Pleistocene pollen sequence of Carihuela Cave, southern Spain. Geobios. 2007; 40 (1):
130. Walker MJ, LoÂpez-MartÂõnez MV, Ortega-RodrigaÂñez J, Haber-Uriarte M, LoÂpez-JimeÂnez A,
AvileÂsFernaÂndez A et al. The excavation of buried articulated Neanderthal skeletons at Sima de las Palomas
(Murcia, SE Spain). Quat Int. 2012; 259 (9):7±21. http://dx.doi.org/10.1016/j.quaint.2011.03.034
131. Zilhão J, Pettitt P. On the new dates for Gorham's Cave and the late survival of Iberian Neanderthals.
Before Farming. 2006; 3: 3. http://dx.doi.org/10.3828/bfarm.2006.3.3
132. SaÂnchez-Yustos P, DÂõez F. Dancing to the rhythms of the Pleistocene? Early Middle Paleolithic
population dynamics in NW Iberia (Duero Basin and Cantabrian Region). Quat Sci Rev. 2015; 121: 75±88.
133. Moure A, Delibes G, Castanedo I, Hoyos M, Cañaveras JC, Housley RA et al. Revision y nuevos datos
sobre el musteriense de la cueva de La Ermita (HortiguÈela, Burgos). In: de BalbÂõn R, Bueno P, editors.
II Congreso de ArqueologÂõa Peninsular. Tomo IÐPaleolÂõtico y EpipaleolÂõtico. Zamora: Fundacion Rei
Alfonso Henriques; 1997. pp. 67±83.
134. AÂlvarez-Alonso D, AndreÂs-Herrero M, DÂõez-Herrero A, Medialdea A, Rojo-HernaÂndez J. Neanderthal
settlement in central Iberia: Geo-archaeological research in the Abrigo del Molino site, MIS 3 (Segovia,
Iberian Peninsula). Quat Int. 2016 in press; http://dx.doi.org/10.1016/j.quaint.2016.05.027
135. Kehl M, AÂ lvarez-Alonso D, AndreÂs-Herrero M, Herrero-DÂõez A, Klasen N, Rojo-HernaÂndez J et al.
Dating the last Neanderthals in Central IberiaÐNew evidence from Abrigo del Molino, Segovia, Spain.
Geophysical Research Abstracts; 2017; 19 (EGU2017): 6402.
136. Navazo M, DÂõez C, Torres T, Colina A, Ortiz JE. La cueva de Prado Vargas. Un yacimiento del
PaleolÂõtico Medio en el sur de la Cordillera CantaÂbrica. Museo de Altamira. MonografÂõas. 2005; 20: 151±166.
137. Arnold L, Demuro M, Navazo M, Benito-Calvo A, PeÂrez-GonzaÂlez A. OSL dating of the Middle
Palaeolithic Hotel California site, Sierra de Atapuerca, north-central Spain. Boreas. 2013;
138. DÂõez C, JordaÂ J, Arceredillo D. El yacimiento paleolÂõtico de Valdegoba (HueÂrmeces, Burgos). In Sala
R, editor. Los cazadors recolectores del Pleisteceno y del Holoeno en Iberia y el Estrecho de Gibraltar.
Burgos: Universidad de Burgos-FundacioÂn Atapuerca; 2014, pp. 608±610.
139. Alcaraz-Castaño M, Alcolea-GonzaÂlez JJ, Weniger GC, Baena-Preysler J, BalbÂõn-Behrmann R,
Cuartero F et al. Neandertal adaptations in Central Iberia: a multi-proxy investigation of the Middle
Paleolithic site of Peña Cabra, Guadalajara, Spain. In Proceedings of the European Society for the study of
Human Evolution 5. AlcalaÂ de Henares (Madrid): European Society for the Study of Human Evolution.
2016; p. 31.
140. DÂõez C, Alonso R, Bengoechea A, Colina A, JordaÂ JF, Navazo M et al. El PaleolÂõtico Medio en el valle
del Arlanza (Burgos). Los sitios de La Ermita, MillaÂn y La Mina. Cuaternario & GeomorfologÂõa. 2008;
22 (3±4): 135±157.
53 / 54
1. d'Errico F , Zilhão J , Baffier D , Julien M , Pelegrin J . Neandertal acculturation in Western Europe? A critical review of the evidence and its interpretation . Curr Anthropol . 1998 ; 39 : S1±S44 . https://doi.org/10. 1086/204689
2. Straus LG . A mosaic of change: the Middle±Upper Paleolithic transition as viewed from New Mexico and Iberia . Quaternary International. 2005 ; 137 : 47 ± 67 . https://doi.org/10.1016/j.quaint. 2004 . 11 .019
3. Vaquero M , Maroto J , Arrizabalga A , Baena J , Baquedano E , CarrioÂn E et al. The Neandertal-modern human meeting in Iberia: a critical view of the cultural, geographical and chronological data . In Conard NJ, editor. When Neanderthals and Modern Humans Met . TuÈbingen: Kerns Verlag; 2006 . pp. 419 ± 439 .
4. Finlayson C , Fa D , JimeÂnez-Espejo F , CarrioÂn J , Finlayson G , Giles-Pacheco G et al. Gorham's Cave, GibraltarÐThe persistence of a Neanderthal population . Quat Int . 2008 ; 181 ( 1 ): 64 ± 71 . https://doi. org/10.1016/j.quaint. 2007 . 11 .016
5. Zilhão J , Davis SJM , Duarte C , Soares A , Steier P , Wild E. Pego do Diabo (Loures, Portugal): Dating the emergence of anatomical modernity in westernmost Eurasia . PLoS ONE; 2010 5 ( 1 ): e8880. https://doi.org/10.1371/journal.pone. 0008880 PMID: 20111705
Acad. Sci. USA . 2013 ; 110 ( 8 ): 2781 ± 2786 . www.pnas.org/cgi/doi/10.1073/pnas.1207656110
7. Finlayson C , Giles-Pacheco F , RodrÂõguez-Vidal J , Fa D , GutieÂrrez JM , Santiago A et al. Late survival of Neanderthals at the southernmost extreme of Europe . Nature . 2006 ; 443 ( 7113 ): 850 ± 853 . https:// doi.org/10.1038/nature05195 PMID: 16971951
8. Maroto J , Vaquero M , Arrizabalaga A , Baena J , Baquedano E , JordaÂ JF et al. Current issues in late Middle Palaeolithic chronology: New assessments from Northern Iberia . Quat Int . 2012 ; 247 : 15 ± 25 . https://doi.org/10.1016/j.quaint. 2011 . 07 .007
13. CabreÂ J , Herreros ME . La cueva de los Casares, Riba de Saelices, Guadalajara. Actes du XVI CongreÂs International dÂAnthropologie (Bruxelles , 1935 ). 1936 ; I: 402 ± 416 .
14. BarandiaraÂn I. La cueva de Los Casares (Riba de Saelices , Guadalajara). Excavaciones ArqueoloÂgicas en España , 76 . Madrid: Ministerio de EducacioÂn y Ciencia; 1973 .
15. Alcaraz-Castaño M , Weniger GC , Alcolea JJ , AndreÂs-Herrero M , Baena J , BalbÂõn R et al. Regreso a la Cueva de Los Casares (Guadalajara). Un nuevo proyecto de investigacioÂn para el yacimiento del Seno A. ARPI . 2015 ; 02 : 68 ± 89 .
17. Basabe JM . Metacarpiano humano de la cueva de Los Casares (Guadalajara). In BarandiaraÂn I, editor. La cueva de Los Casares (Riba de Saelices, Guadalajara). Excavaciones ArqueoloÂgicas en España , 76 . Madrid: Ministerio de EducacioÂn y Ciencia; 1973 . pp. 117 . 122 .
18. Arribas A , JordaÂ JF . Los mamÂõferos del Cuaternario kaÂrstico de Guadalajara (Castilla-La Mancha , España). In: Aguirre E, RaÂbano I , editors. La Huella del Pasado . FoÂsiles de Castilla-La Mancha. Patrimonio HistoÂrico. ArqueologÂõa Castilla-La Mancha ; 1999 . pp. 327 ± 353 .
19. Yravedra J . Aproximaciones tafonoÂmicas a los cazadores de la segunda mitad del Pleistoceno superior de la mitad norte del interior de la PenÂõnsula IbeÂrica . Arqueoweb. 2007 ; 9 ( 1 ).
20. Mingo A , Barba J , GarcÂõa MA , Berzosa R . El yacimiento prehistoÂrico de Los Casares (Riba de Saelices, Guadalajara): revisioÂn del material lÂõtico y ceraÂmico depositado en el Museo ArqueoloÂgico Nacional y sus implicaciones crono-culturales . Quad. Preh. Arq. Cast . 2014 ; 32 : 21 ± 42 .
24. Domingo R , Peña-MonneÂ JL , Torres T , Ortiz E , Utrilla P . Neanderthal highlanders: Las Callejuelas (Monteagudo del Castillo, Teruel, Spain), a high-altitude site occupied during MIS 5 . Quat Int . 2017 ; 435/A: 129 ± 143 . http://dx.doi.org/10.1016/j.quaint. 2015 . 09 .088
25. Hoffmann DL , Pike AWG , Wainer K , Zilhão J . New U-series results for the speleogenesis and the Palaeolithic archaeology of the Almonda karstic system (Torres Novas, Portugal) . Quat Int . 2013 ; 294 : 168 ± 182 . http://dx.doi.org/10.1016/j.quaint. 2012 . 05 .027
26. Zilhão J . Neandertal-Modern Human Contact in Western Eurasia: Issues of Dating, Taxonomy, and Cultural Associations . In: Akazawa T., Nishiaki Y. , Aoki K ., editors. Dynamics of Learning in Neanderthals and Modern Humans Volume 1 . Cultural Perspectives . Springer Japan; 2013 . pp. 21 ± 57 .
27. Peña P. The transition in southern Iberia: Insights from paleoclimatology and the Early Upper Palaeolithic . Proc Natl Acad Sci USA . 2013 ; 110 ( 23 ): E2086. https://doi.org/10.1073/pnas.1303596110 PMID: 23633575
28. GalvaÂn B , HernaÂndez C , Mallol C , Mercier N , Sistiaga A , Soler V . New evidence of early Neanderthal disappearance in the Iberian Peninsula . J. Hum. Evol . 2014 ; 75 : 16 ± 27 . http://dx.doi.org/10.1016/j. jhevol. 2014 . 06 .002 PMID: 25016565
29. Higham T , Douka K , Wood R , Bronk Ramsey C , Brock F , Basell L et al. The timing and spatiotemporal patterning of Neanderthal disappearance . Nature . 2014 ; 512 ( 7514 ): 306 ± 309 . https://doi.org/10.1038/ nature13621 PMID: 25143113
30. Baena-Preysler J , Ortiz I , Torres C , BaÂrez S. Recycling in abundance: Re-use and recycling processes in the Lower and Middle Paleolithic contexts of the central Iberian Peninsula . Quat Int . 2015 ; 361 : 142 ± 154 . http://dx.doi.org/10.1016/j.quaint. 2014 . 07 .007
32. Beckmann T. PraÈparation bodenkundlicher DuÈnnschliffe fuÈr mikromorphologische Untersuchungen . Hohenheimer Bodenkundliche Hefte . 1997 ; 40 : 89 ± 103
55. Schweingruber FH . European wood anatomy . 1st ed. Bern: Paul Haupt; 1990 .
77. Bunn HT . Meat-eating and human evolution: studies on the diet and subsistence patterns of PlioPleistocene hominins in East Africa . Ph. Dissertation . Berkeley: University of California. 1982 .
141. Navazo M , Carbonell E. Neanderthal settlement patterns during MIS 4±3 in Sierra de Atapuerca (Burgos, Spain) . Quat Int . 2014 ; 331 : 267 ± 277 . http://dx.doi.org/10.1016/j.quaint. 2014 . 03 .032
142. Cacho C , Martos J , JordaÂ J , Yravedra J , Avezuela B , Valdivia J et al. El PaleolÂõtico superior en el interior de la PenÂõnsula IbeÂrica . RevisioÂn crÂõtica y perspectivas de futuro. In: Mangado X. Editor, El PaleolÂõ- tico superior peninsular. Novedades del Siglo XXI . Barcelona: Universitat de Barcelona; 2010 . pp. 115 ± 136 .
143. Alcaraz-Castaño M. Central Iberia around the Last Glacial Maximum: Hopes and Prospects . Journal of Anthropological Research . 2015 ; 71 ( 4 ): 565 ± 578 .
144. Arsuaga JL , GoÂmez-Olivencia A , BonmatÂõ A , Pablos A , MartÂõnez-Pillado V , Lira J et al. Neandertals at Atapuerca: the MIS3 GalerÂõa de las Estatuas site . In Proceedings of the European Society for the study of Human Evolution 5 . AlcalaÂ de Henares (Madrid): European Society for the Study of Human Evolution . 2016 ; p. 36 .
145. Moure A , GarcÂõa-Soto E . Radiocarbon dating of the Mousterian in Cueva MillaÂn (HortiguÈela , Burgos, Spain). Curr Anthropol . 1983 ; 19 : 155 ± 157 .
146. Silva P , LoÂpez-Recio M , Cuartero F , Baena J , Tapias , Manzano I et al. Contexto geomorfoloÂgico y principales rasgos tecnoloÂgicos de nuevos yacimientos del Pleistoceno Medio y Superior en el Valle Inferior del Manzanares (Madrid, España). Estudios GeoloÂgicos . 2012 ; 68 ( 1 ): 57 ± 89 .
147. Schmidt I , BradtmoÈller M , Kehl M , Pastoors A , Tafelmaier Y , Weninger B et al. Rapid climate change and variability of settlement patterns in Iberia during the Late Pleistocene . Quat Int . 2012 ; 179 ± 204 .
148. Alcaraz-Castaño M , Alcolea J , de BalbÂõn R , GarcÂõa-Valero MA , Yravedra J , Baena J . Los orÂõgenes del Solutrense y la ocupacioÂn pleniglaciar del interior de la PenÂõnsula IbeÂrica: implicaciones del nivel 3 de Peña CapoÂn (valle del Sorbe, Guadalajara) . Trabajos de Prehistoria . 2013 ; 70 ( 1 ): 28 ± 53 . http://dx. doi.org/10.3989/tp. 2013 .12101
149. Sepulchre P , Ramstein G , Kageyama M , Vanhaeren M , Krinner G , SaÂnchez- Goñi MF et al. H4 abrupt event and late Neanderthal presence in Iberia . Earth Planet. Sci. Lett . 2007 ; 258 : 283 ± 292 .
150. Finlayson C , CarrioÂn JS . Rapid ecological turnover and its impact on Neanderthal and other human populations . Trends Ecol. Evol . 2007 ; 22 : 213 ± 222 . https://doi.org/10.1016/j.tree. 2007 . 02 .001 PMID: 17300854
151. BradtmoÈller M , Pastoors A , Weninger B , Weniger GC . The repeated replacement model±Rapid climate change and population dynamics in Late Pleistocene Europe . Quat Int . 2012 ; 247 : 38 ± 49 . https://doi.org/10.1016/j.quaint. 2010 . 10 .015
152. Hodgkins J , Marean CW , Turq A , Sandgathe D , McPherron SJP , Dibble H . Climate-mediated shifts in Neandertal subsistence behaviors at Pech de l'AzeÂ IV and Roc de Marsal (Dordogne Valley , France). J Hum Evol . 2016 ; 96 : 1± 18 . https://doi.org/10.1016/j.jhevol. 2016 . 03 .009 PMID: 27343769
153. Baquedano E , Arsuaga JL , PeÂrez-GonzaÂlez A , MaÂrquez B , Laplana C , Ortega MC et al. The DesCubierta Cave (Pinilla del Valle , Comunidad de Madrid, Spain): a Neanderthal site with a likely funerary/ritualistic connection . In Proceedings of the European Society for the study of Human Evolution 5 . AlcalaÂ de Henares (Madrid): European Society for the Study of Human Evolution . 2016 ; p. 41 .
154. Klasen N , Kehl M , Alcaraz-Castaño M , AÂlvarez-Alonso D , AndreÂs-Herrero M , Zilhão J et al. Application of luminescence dating of archaeological sequences±examples from Europe and North Africa . Geophysical Research Abstracts; 2017 ; 19 ( EGU2017 ): 6919 .