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As the south-westernmost region of Europe, the Iberian Peninsula stands as a key area for understanding the process of modern human dispersal into Eurasia. However, the precise timing, ecological setting and cultural context of this process remains controversial concerning its spatiotemporal distribution within the different regions of the peninsula. While traditional models assumed that the whole Iberian hinterland was avoided by modern humans due to ecological factors until the retreat of the Last Glacial Maximum, recent research has demonstrated that hunter-gatherers entered the Iberian interior at least during Solutrean times. We provide a multi-proxy geoarchaeological, chronometric and paleoecological study on human–environment interactions based on the key site of Peña Capón (Guadalajara, Spain). Results show (1) that this site hosts the oldest modern human presence recorded to date in central Iberia, associated to pre-Solutrean cultural traditions around 26,000 years ago, and (2) that this presence occurred during Heinrich Stadial 2 within harsh environmental conditions. These findings demonstrate that this area of the Iberian hinterland was recurrently occupied regardless of climate and environmental variability, thus challenging the widely accepted hypothesis that ecological risk hampered the human settlement of the Iberian interior highlands since the first arrival of modern humans to Southwest Europe.
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
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First modern human settlement
recorded in the Iberian hinterland
occurred during Heinrich Stadial
2 within harsh environmental
conditions
M. Alcaraz‑Castaño 1*, J. J. Alcolea‑González1, M. de Andrés‑Herrero1, S. Castillo‑Jiménez1,
F. Cuartero2, G. Cuenca‑Bescós3, M. Kehl4, J. A. López‑Sáez5, L. Luque1, S. Pérez‑Díaz6,
R. Piqué7, M. Ruiz‑Alonso5, G.‑C. Weniger8 & J. Yravedra9
As the south‑westernmost region of Europe, the Iberian Peninsula stands as a key area for
understanding the process of modern human dispersal into Eurasia. However, the precise timing,
ecological setting and cultural context of this process remains controversial concerning its
spatiotemporal distribution within the dierent regions of the peninsula. While traditional models
assumed that the whole Iberian hinterland was avoided by modern humans due to ecological factors
until the retreat of the Last Glacial Maximum, recent research has demonstrated that hunter‑
gatherers entered the Iberian interior at least during Solutrean times. We provide a multi‑proxy
geoarchaeological, chronometric and paleoecological study on human–environment interactions
based on the key site of Peña Capón (Guadalajara, Spain). Results show (1) that this site hosts the
oldest modern human presence recorded to date in central Iberia, associated to pre‑Solutrean
cultural traditions around 26,000 years ago, and (2) that this presence occurred during Heinrich
Stadial 2 within harsh environmental conditions. These ndings demonstrate that this area of the
Iberian hinterland was recurrently occupied regardless of climate and environmental variability, thus
challenging the widely accepted hypothesis that ecological risk hampered the human settlement of
the Iberian interior highlands since the rst arrival of modern humans to Southwest Europe.
The rst modern human settlement of southwest Europe and the role of the Iberian hinter‑
land. e rst appearance of anatomically modern humans in a given region of the world is always a con-
tentious topic. In Western Europe, the Iberian Peninsula stands as the last region of the process of modern
human dispersal into Eurasia, and hence is considered of key importance for understanding its cultural and
natural constraints. However, the precise timing, ecological setting and cultural context of this process remains
especially controversial when considering its spatiotemporal distribution within the dierent regions of the pen-
insula. Bearing aside the controversy on the makers of the Chatelperronian and other so-called transitional
technocomplexes13, if we accept the Proto-Aurignacian as the rst proxy for modern humans in Western
Europe, these people were present in the Cantabrian and northern Mediterranean regions of Iberia at 42kacal
BP4,5 (Fig.1A), or even 43kacal BP6. is is a roughly similar time as recorded in other regions of Western and
Central Europe7,8, although signicantly younger than in Eastern Europe according to recent data9,10. However,
in light of prevailing evidence, it was much later when Aurignacian cultures spread to the southern parts of
OPEN
Prehistory Area, University of Alcalá, Alcalá de Henares, Spain. Atapuerca Foundation, Burgos,
Spain.           
          
             
           
Spain.          
 *
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Iberia, reaching the southwesternmost regions of Europe at around 36.5kacal BP11, probably matching the time
of the Neandertals’ nal demise1214 (Fig.1B). Signicantly earlier occurrences of Proto-Aurignacian have been
recently claimed for the sites of Bajondillo (43.0–40.8kacal BP)15 and Lapa do Picareiro (41.1–38.1kacal
BP)16, in southern Spain and central Portugal respectively. However, published archaeological and chronostrati-
graphic data in the case of Bajondillo are disputable and have been strongly contested14,1719.
e vast hinterland territories of the Iberian Peninsula, an upland plateau divided into the Northern and
Southern Mesetas by the Central System mountain range (Figs.1 and 2), have been traditionally considered irrel-
evant in this process, as they have been regarded as a virtually “no-man’s land” until the Late Upper Paleolithic20.
At present, there is some sparse data between 33 and 28kacal BP in the western and northern borders of the
Northern Meseta14,2124 (Fig.1C). However, to date, eective presence of modern human occupations in the
central regions of Iberia is not found until 25.5kacal BP, as shown by preliminary evidence gathered at one
single site: the Peña Capón rock shelter25,26 (Fig.1C).
Reasons behind this odd population pattern have revolved around a potentially late-persisting Neandertal
presence in the center and south of Iberia12 and, more prominently, the potentially harsh climatic and environ-
mental conditions of the interior and upland regions of Iberia as opposed to the more favored environments
Figure1. Process of peopling of the Iberian Peninsula by modern humans during the Upper Paleolithic. A:
42 – 38kacal BP, B: 38 – 30kacal BP, C: 30–25kacal BP, D: 25–20kacal BP (see Supplementary Text S3 for
discussion and Supplementary Datasets 1–4 for full data). Maps generated with ArcGIS (ArcMap 10.3.1.)
(https:// www. arcgis. com/ index. html) using ASTER Global Digital Elevation Model V0032019, distributed by
NASA EOSDIS Land Processes DAAC, (10.5067/ASTER/ASTGTM.003).
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of the peninsular coastal regions, long considered as refugia for ora, animals and humans [see20,27]. Recent
reviews have roughly supported this latter picture, as they have limited the pre-Magdalenian human presence
in most of inland Iberia to short-term or sporadic incursions during temperate intervals starting only in Solu-
trean times (i.e. between 25 and 20kacal BP)11,27,28. Furthermore, habitat suitability models based mainly on
paleoclimate simulations2933 have provided further support to the traditional model. ese works describe the
Iberian interior as an ecologically risky area for human settlement, especially during the Last Glacial Maximum
(LGM) sensu stricto (i.e. 23–19ka BP)34, due to its relatively high elevation, degree of climate variability and
resource unpredictability (but see3537 for signicant dierences concerning habitat suitability of the Iberian
interior, due to the high number of variables and methods involved in modeling building). Other studies have
recently pointed to arid and cold environmental conditions in central Iberia during 40–30kacal BP. ey are
based both on paleoecological38,39 and sedimentological40,41 data and provide further support to the idea that
climate and environmental conditions somehow hampered the human occupation of these regions during the
beginning of the Upper Paleolithic.
However, the idea of inland Iberia as either a desolate landscape or a mere crossing-area where human groups
based elsewhere entered only sporadically during most of the Upper Paleolithic has been under attack in the
last years14,20,42. Among the growing number of evidence suggesting a more relevant settlement of the Iberian
interior not only during the Solutrean, but also before20,43, data recorded at the Peña Capón rock shelter has
revealed crucial25,26 (Supplementary Text S1). Here we report results from new eldwork and laboratory analyses,
including geomorphology, sedimentology, micromorphology, radiocarbon dating, palynology, anthracology, zoo-
archeology, microvertebrate paleontology and analysis of lithic technology, with the main aim of providing new
evidence on human–environment–climate interactions during the rst settlement of the Iberian central regions
by modern humans recorded to date. Under a theoretical framework that conceives cultural change, population
dynamics and adaptive traits of hunter-gatherers as multifactorial responses to uctuating social and natural
parameters4446, including climate and environmental change4749, these results show relevant patterns concerning
the timing, nature and ecological setting of this process. More specically, considered in the context of recent
research on the relations between population dynamics, settlement patterns and techno-cultural change in the
Late Pleistocene of Iberia on one side, and rapid climate and environmental change on the other27,2933,5056, our
results allow us to test the hypothesis that the rst modern human settlement of inland Iberia occurred earlier
than previously thought, and was not impeded by ecological variability.
Figure2. Geological maps showing the location of the Peña Capón rock shelter in the Iberian Peninsula
and the Tagus basin (Guadalajara, Spain) (A), the Sorbe River basin (B) and at the shore of the Beleña water
reservoir (C). Maps generated using QGIS Open Source Geographic Information System v. 3.4 (Madeira)
(https:// www. qgis. org/ en/ site/ about/ index. html) combined with Digital Terrain Models and slope maps from
the Spanish National Centre for Geographic Information (CNIG) (https:// www. ign. es/ web/ ign/ portal/ qsm- cnig)
and geological maps from the Spanish Geological Survey (IGME) (https:// www. igme. es/ zarag oza/ ingles/ inicio.
htm).
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The Peña Capón rock shelter and its regional setting. e Peña Capón rock shelter (Guadalajara
province, Spain) is located near the le bank of the Sorbe River, which ows into the Henares, tributary of the
Tagus, the main Iberian watercourse, crossing the Spanish Southern Meseta from E to W (Fig.2). e Sorbe
has its source in the highest part of Sierra de Pela, a mountain range located in the eastern limit of the Central
System Range at 1,500m above mean sea level (amsl). e river runs southwards, rst crossing gentle Paleozoic
reliefs (schist, quartzite and slate) and marine Mesozoic carbonates before meeting the alluvial terrains of the
Tertiary sediment inll of the Tagus basin, and nally joining the Henares River at a height of 710m amsl. e
archaeological site is located at an altitude of 826m amsl and 11 to 13.5m above the current riverbed, under a
dolostone rock cli, in an area where the Sorbe valley widens and the Quaternary uvial and alluvial deposits
become more frequent (Figs.2 and 3B). e Quaternary deposits are described in the geological maps of the
region57,58 and mainly consist of uvial terraces, alluvial cones and slope deposits (Fig.3A). According to these
geological maps, there are fourteen Quaternary uvial terraces, from + 6 to + 180–190m above the current river-
bed. ose below + 20m are generally considered Upper Pleistocene and Holocene in other nearby areas of the
Tagus basin and Duero basins59,60.
e Peña Capón site is located under an east–west oriented, 42m high rock cli, formed by Upper Cretaceous
marine dolostone layers dipping to the south. e dolostone outcrops as part of a long hogback relief oriented to
the NW–SE that surrounds the Paleozoic shales, schists and quartzites, as well as the Lower Triassic Buntsandstein
facies, located to the north and west. e archaeological site is located 80m away from the current riverbed,
close to a narrowing of the valley excavated in the dolostone relief. Due to its location, the site is ooded by the
Beleña reservoir waters for most of the year since a dam was constructed in 1982 (Supplementary Figs.S1–S2,
S4–S7 and Supplementary Video 1).
Results
Stratigraphy, sedimentology and micromorphology. e geomorphological analysis of the site
indicates that the accommodation space where the Upper Pleistocene sediments were deposited, at the foot
of the rock wall, was created due to (1) the high slope angle of the dolostone running west to east and crossing
this sector of the valley and (2) the dierential erosion of a less compact marly and nely laminated layer at the
base of the compact dolostone formation. Vertical dolostone layers parallel to the rock wall and several large
gravitational blocks could have generated a sort of corridor where mostly ne-grained deposits accumulated and
were protected from erosion, as it is also observed on the opposite riverbank (Supplementary Figs.S5–S6). e
Figure3. (A) Geomorphological map of the study area showing the position of Peña Capon at the foot of a
dolomite cretaceous relief, and the distribution of the Quaternary deposits located in the area. (B) General view
of the site from above. Map generated as explained in Fig.2.
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geomorphological map shows that both uvial oods of Sorbe River and surface runo on nearby alluvial fans
could have played a role in the formation of the deposits (Fig.3A).
e Peña Capon archaeological site covers a 30m long and 5 to 8m wide area of about 150 sq m along the foot
of the dolostone rock wall (Fig.3B; Supplementary Fig.S5). e deposit slopes 4.5° to the west, perpendicular
to the direction of the valley. e maximum thickness of the deposit recorded in the archaeological excavations
is 0.95m.
From a sedimentological point of view, the archaeological deposit consists of ne sand and silt intermixed
with rock fragments showing a mixture of natural and anthropogenic sediment components, including varying
amounts of lithics, charcoal and bones. Rock fragments consist of subangular dolostone derived from the rock
shelter wall and sub-rounded to rounded quartzite and ne siltstone gravel. Few small angular pieces of chert,
rock crystal and quartzite represent by-products of tool production61. e stratigraphic sequence is composed
of six dierent sedimentological units containing archaeological remains (Levels 1 to 6), dened mainly on the
basis of sediment color variation (Fig.4). An overlying sedimentary unit of heterogeneous dark grey sandy loam,
containing mixed archaeological remains including few pottery sherds, unconformably covers the Pleistocene
archaeological deposit. is layer has been named R and can be roughly subdivided into a darker (Munsell Color
Code 0,7Y 5/2), more coarse-grained lower one (R1) and the lighter-colored (0,4Y 5/2), ner-grained upper one
(R0). e layers of the archaeological deposit are quite homogenous, sharing many common sedimentological
and micromorphological features but varying in color due to their dierent content in burned components and
carbonate. Levels 1, 3, 5 and 6 are light orange-brown to reddish-brown (9YR 5/3), while levels 2 and 4 are darker
grey-brown (10YR 5/2, 10YR 5/3). Levels 1 and 3 are very similar, corresponding to mainly homogeneous ne
sand and silt layers. Level 1, thicker, shows some lamination in its lower part. Level 3 thins eastward and has a
sharp contact with the overlying level 2 and diuse transition to the underlying level 4. Level 2 is divided into two
sub-units, 2a and 2b. 2a on top is grey-brown and contains charcoal-rich lenses with abundant bone fragments; 2b
below, is darker and shows higher contents of charcoal and charred organic matter. Its lower contact is irregular.
Level 2b includes some small pits and a possible micromammal burrowing. e color of level 4 ranges from
reddish brown to a more intense reddish tone (10YR 5/2 to 5YR 5/3), suggesting rubefaction processes. Levels
5 and 6, in the lower part of the sequence, are lighter in color (10YR 7/3) and richer in secondary carbonate.
e granulometric analyses of the sediment sequence display a homogeneous textural composition dominated
by poorly sorted very ne sand and coarse silt, with a low proportion of clay-size particles (maximum of 10%)
(Fig.S19). e carbonate content ranges from 10 to 32% with an increase in the lower levels. Total Organic Car-
bon (TOC) values range from 0.3 to 2.9% and match with the color change, indicating that this is mainly related
to variation in organic matter content. in sections show that charcoal with well-preserved cell structures is
common in dark-colored levels, where it occurs together with amorphous charred organic matter of unknown
origin (Fig.5). High values in magnetic susceptibility, χlf, also occur in the dark-colored, organic matter-rich
levels R1, 2b and 4, i.e. layers with high amounts of charred organics. Sediment generally has a low degree of
compaction, related to presence of abundant pores consisting of biogenic channels and burrows formed by roots
or soil-dwelling mesofauna62 creating bioturbation on a microscale. Geometry and thickness of the stratigraphic
levels are represented in Fig.4, and detailed results of sedimentology and micromorphology are included in
Supplementary Texts S4 and S5 and Supplementary TableS2.
Radiocarbon dating and Bayesian modeling. e chronological setting of the Peña Capón sequence
is based on radiocarbon dating. Selected samples are faunal remains with anthropogenic modications (mostly
cut marks) (n = 21) and charcoal fragments (n = 12) recovered from secure stratigraphic contexts and covering
the whole sequence excavated thus far. ese samples were rst identied to taxon (when possible) and then sent
to two dierent laboratories for cross-checking results: 15 samples were sent to the CologneAMS centre at the
University of Cologne and 18 samples to the Oxford Radiocarbon Accelerator Unit (ORAU) at the University
of Oxford. Out of 33 samples we obtained 22 AMS reliable determinations and found no signicant dierences
between results provided by both labs. One charcoal sample showed a modern age, while eight bones and two
charcoals, mostly from levels 4 to 6, failed due to low, very low or no yield (Supplementary TableS3).
With the aim of building a strong probabilistic framework for the sequence of human occupations recorded at
Peña Capón we constructed a Bayesian model based on obtained radiocarbon determinations. As lengthy applied
and discussed in the last years, if properly devised, Bayesian modeling is an accurate and informative way to
integrate radiometric data with stratigraphically recorded archaeological evidence to create reliable chronological
models, and thus improve chronometric precision, mitigating uncertainties and eliminating outliers6366. A pre-
liminary model (Model 1), containing all radiocarbon dates, showed an Amodel of 53.2, thus pointing to potential
problems in the relation between the prior and posterior distributions (i.e. between the unmodeled dates and
their location within the sequence on the one hand, and the modeled calibrated results on the other). One date
from level 3 (COL4217.1.1) and one from level 5 (OxA-39749) showed agreement indexes < 60% and posterior
probabilities of being outliers of 81% and 14% respectively (Supplementary TableS4 and Fig.S21). Hence, these
dates were not included in the nal model (Model 2). is model, composed of 19 radiocarbon dates, is presented
in Fig.6 and Supplementary TableS5, and shows individual agreement values ranging between 139.3 and 74.7,
an Amodel of 107.6, and posterior probabilities of ≤ 5% for all determinations. e consistency of Model 2 is very
high, as it shows an excellent degree of agreement between prior information, radiocarbon determinations and
posterior probabilities, thus conrming geoarchaeological interpretation on the site’s stratigraphic integrity based
on sedimentology and micromorphology.
e model results allow us to place the start boundary of the Peña Capón sequence excavated thus far
(base of level 6) between 26.3 and 25.7kacal BP at 95.4% probability, and the end boundary (top of the known
sequence) at 24.1–23.6kacal BP. However, when considering the results of applying the ‘date’ command in OxCal,
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the Probability Distribution Function (PDF) for the time span of human occupation at the site is constrained
between 26.1 and 23.9kacal BP. e calendar age estimates for each level are shown in Table1 and their associ-
ated PDFs are provided in Supplementary Fig.S22 (complete data and boundaries between levels are shown in
Supplemaentary TableS5).
ese results conrm the high sedimentation rate of the deposit25,26, where few more than 2,000years are
recorded in 95cm. is explains the overlap between some dates (both unmodeled and modeled) obtained in
adjacent levels, which is hence not related to post-depositional mixing—as also shown by sedimentology and
micromorphology—but to the standard deviations of radiocarbon dates.
Figure4. (A) Stratigraphic units dened in the Peña Capón site. Note that the main dierences between layers
are due to variations in organic matter and secondary carbonate content. (B) Stratigraphic sequence recorded
in the western prole of square 2B showing sample location for micromorphology (B1–B5) and sedimentology
(dashed-lines rectangles). (C) Views of the excavation proles at Peña Capón showing the dierent distribution
and geometry of the stratigraphic units dened in the site.
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Figure5. Flatbed scans (1200 dpi) of selected thin sections from the sediment sequence at Peña Capón. (A) Erosional
contact between level R1 (LR1) characterized by strong compaction and a platy microstructure and level 1 (L1) showing less
compaction and a channel microstructure. At the interface, a sediment lens rich in charcoal and bone is present (thin section
PCPN 1.2). e dashed line is the boundary between the two levels. (B) in section PCPN 6.1 with the interface between
levels L1 and L2a. At the sampling location, the upper part of L2a consists of a 2cm thick band rich in charcoal and bone
fragments. (C) in section PNCP 3.2 showing the gradual transition between levels 2a and 2b. (D) Same as C but captured
under XPL. Note abundant calcite hypocoatings around biopores. (E) in section PNCP 4.1 from level L2b with several
large biopores partly relled with granules. (F) e interface between levels L3 and L4, which is delineated by a thin layer of
ne gravel. e light-colored L3 shows diuse impregnation with secondary calcite. (G) Same as E, but captured under XPL.
Secondary carbonate is indicated by high birefringence. (H) in section PNCP 3.2 showing the transition from level 4 to
level 5.
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Concerning the cultural sequence, the Solutrean at Peña Capón is rst recorded at 25.3ka BP and lasts
until 23.8ka BP. In turn, the rst modern human presence recorded thus far, associated in level 6 to pre-
Solutrean assemblages (either Gravettian or Proto-Solutrean), is detected at 26.1ka BP. Both episodes started
and developed during Greenland Stadial 3 (GS 3), a stadial phase within the LGM69 and, more signicantly, both
were triggered during the rapid cooling of Heinrich Stadial 2 (HS 2) (Fig.6). In fact, the whole pre-Solutrean
Figure6. Bayesian Final Model (2) for the Peña Capón sequence showing Probability Distribution Functions
(PDFs) for all radiocarbon determinations and boundaries between archaeological levels. Results are plotted
against the δ18O record of the NGRIP ice core, indicating Greenland Interstadials 3 and 2 (GI 2 & GI 3),
Greenland Stadial 3 (GS 3)67, and the chronology of Heinrich Stadial 2 (blue bar)68. 14C dates are shown
in parentheses, and Agreement indexes and Outliers’ prior and posterior probabilities are shown in square
brackets. Calibration of dates and Bayesian modeling were calculated using OxCal 4.4 online soware (https://
c14. arch. ox. ac. uk/ oxcal. html).
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occupation, and most of the Solutrean one, were developed during HS 2. is bears relevant implications for
understanding human-climate-environment interactions during the rst settlement of the Iberian interior by
modern humans, as will be discussed below.
Pollen. For the Peña Capón sequence, 9 pollen spectra were analyzed and 28 taxa were identied. To facilitate
description and interpretation of the pollen diagram with respect to vegetational changes, three Local Pollen
Assemblage Zones (LPAZs) were established (Fig.7). ese zones denote signicant changes in the pollen com-
position and represent major changes in vegetation. LPAZ-1 is largely dominated by Pinus nigra (31.5–35.2%)
indicating a Spanish black pine forest in the vicinity7072, which represents trees that, nowadays, grow in the
higher mountains of Mediterranean central Iberia under supramediterranean climatic conditions (mean annual
temperature of 8.13°C and 400–1000mm of annual rainfall)73,74. Quercus ilex/coccifera (9.4–13.1%) and Juni-
perus (7.4–9.9%) show continuous high values, concomitant with those of Berberis vulgaris (1.1–2.2%), Linum
(1.1–1.7%) and Rhamnus (2.3–3.3%), suggesting the regional presence of holm oak (Quercus ilex subp. rotun-
difolia) and juniper (Juniperus thurifera) woodlands75,76. Mesophilous trees such as Quercus pyrenaica/faginea,
Acer, Alnus, Betula, Fraxinus, Salix and Tilia are usually low (< 5%) and beech (Fagus) is absent. is zone shows
the highest percentages of herbs (30.1–32.6%), mainly represented by cryoxerophytic taxa (Artemisia 5.5–8.5%,
Chenopodiaceae 7.4–8.5%) and heliophilous/cryophilous herbs (Poaceae 5.9–9.4%), probably indicating cold
and dry conditions, as also shown by cryoxerophytic shrubs, such as Helianthemum (2.8–4.6%). Anthropogenic-
zoophilous pollen taxa (Asterioideae, Carduoideae, Cichorioideae) present relatively low values that do not
exceed 12%.
LPAZ-2 is characterized by synchronous increases of mesophilous taxa such as Quercus pyrenaica/faginea
(23.8–25%), Acer (5–5.5%), Alnus (9.3–9.8%), Betula (2.7–3%), Fraxinus (5.3–7.9%), Salix (4–5%) and Tilia
(3.7–6%), and the rst appearance in the pollen diagram of Fagus sylvatica (2.7–3.7%). ese results suggest (i)
Table 1. Calendar age estimates for each archaeological level at Peña Capón, based on the Bayesian Model
2 (Fig.6) as calculated by the ‘date’ command in OxCal 4.4 online soware (https:// c14. arch. ox. ac. uk/ oxcal.
html). Associated PDFs are shown in Fig.S22.
Level (phase)
Duration (cal BP)
68.2%
probability 95.4%
probability
Peña Capón 25,770 24,110 25,954 23,874
1 (Solutrean) 24,064 23,868 24,190 23,784
2a (Solutrean) 24,330 24,092 24,460 23,968
2b (Solutrean) 24,792 24,370 25,046 24,282
3 (Solutrean) 25,210 24,970 25,264 24,668
4 (Pre-Solutrean) 25,280 25,140 25,406 25,068
5 (Pre-Solutrean) 25,616 25,260 25,798 25,192
6 (Pre-Solutrean) 25,962 25,722 26,130 25,448
Figure7. Percentage pollen diagram from the Peña Capón sequence.
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denser deciduous oak woodlands (trees and shrubs values between 88.4 and 90%), and (ii) the rst arrival of
beech near the site as a result of climatic warming and increased humidity77. us, the Iberian Central System
is conrmed as a refuge zone for this taxon during much of the Late Pleistocene, including the LGM7880, as has
been also documented in other mountains in northern Iberia81,82. is nding represents the rst record of beech
from the LGM in the Iberian Central System. Pinus nigra, Quercus ilex/coccifera and Juniperus decrease, as well as
cryoxerophytic (Artemisia, Chenopodiaceae) and heliophilous/cryophilous (Poaceae) taxa, while Helianthemum,
Berberis vulgaris, Linum and Rhamnus disappear. Shrub and herb taxa percentages related to deciduous oak
woods (Arctostaphylos uva-ursi 1.4–2.4%, Cistus laurifolius 3.3–4.3%, Cytisus/Genista 3.3–3.7%, Geum 1.2–1.7%,
Lavandula stoechas 1.3–1.8%) experience an increasing tendency, while anthropogenic-zoophilous pollen taxa
(Asterioideae 14.7–22.1%, Carduoideae 6–7.9%, Cichorioideae 26–26.4%) are much more abundant.
Finally, LPAZ-3 shows a similar pattern to that of LPAZ-1. Pinus nigra is at its maximum in this pollen
sequence (37.3%). Other components increase such as Quercus ilex/coccifera (10.1–11.6%), Juniperus (7.7–11.6%),
Berberis vulgaris (1.8–2.2%), Rhamnus (1.8–2.2%), Artemisia (6.1–7.7%), Chenopodiaceae (6.1–7.7%), Helian-
themum (4.1–5%), Linum (0.6–1.1%) and Poaceae (8.8–8.9%), while some tree or shrub taxa (Acer, Alnus, Betula,
Cistus laurifolius, Cytisus/Genista, Fraxinus, Quercus pyrenaica/faginea, Tilia) abruptly decrease and some even
disappear (Fagus, Salix), as do Arctostaphylos uva-ursi, Lavandula stoechas and Geum. Anthropogenic-zoophilous
taxa (Asterioideae, Carduoideae, Cichorioideae) also decrease.
Wood charcoal. Charcoal remains were mostly scattered throughout the excavated area, besides some con-
centrations, including a replace in level 2a (Supplementary Figs.S12–S13). In general terms, carbonized wood
found at Peña Capón is scarce in relation to the number of fragments recovered. Even in the otation samples,
where all charcoal remains present in the sediment are recovered, the number of identied fragments is low, lim-
ited in many cases to just one fragment, being most of them smaller than 2cm. ese samples mostly included
black and ash-gray sediment devoid of charcoal input.
Among the 154 identied wood fragments, taxonomic diversity is low (Table2). Most of the fragments have
been assigned to either Salix sp. or Juniperus sp., being the rest taxa (Alnus sp., Cistus sp., Fraxinus sp., Legumi-
nosae, Rhamnus/Phillyrea, Rosaceae) represented by a low number of fragments. Moreover, a large number of
remains could only be identied as angiosperms dicotyledons or gimnosperms (conifers) due to poor preserva-
tion or small size. A total number of 52 remains were unidentiable.
Unlike other proxies studied at the site –especially pollen– wood charcoal analysis do not show signicant
environmental dierences throughout the sequence. us, charcoal results point to a recurrent pattern of wood
procurement from level 5 to 1, being level 6 the only episode where a dierent behavior is attested (Table2).
is level has shown the higher number of Juniperus sp. (juniper/sabina) fragments, together with one remain
of Rhamnus/Phillyrea and one of Rosaceae (Maloideae type), while Salix is totally absent. In contrast, in levels 5
and 4 Salix wood is the most represented, and Juniperus is represented only by 4 fragments in level 5.
It is noteworthy that thicker and more extensively excavated levels (1, 2a, 2b and 3) (Fig.4) present relatively
lower amounts of charcoal remains compared to levels excavated only in the 1 sq meter test pit (4, 5 and 6).
us, in these levels, corresponding to the Solutrean human occupations, the number of identied fragments
is scarce, and a relevant number of them have been assigned to unidentiable angiosperms, mostly due to their
small size and their deformed and altered conditions. However, levels 1, 2a and 2b show the greatest diversity
within the sequence, including taxa such as Alnus, Cistus, Fraxinus or Leguminosae (although limited in most
cases to isolated fragments) together with Salix and Juniperus (Table2).
Although the low number of identied charcoals calls for some caution concerning the palaeoenvironmental
interpretation of these results, some conclusions can be drawn from them. Juniperus sp, present in levels 1, 2b, 5
and especially 6, is a little tree or shrub typical of open landscapes, and some species within this genus, such as
Table 2. Wood charcoal identied in Peña Capón.
Taxon/level 1 2a 2b 3 4 5 6
Alnus sp 1
Cistussp 1 1
cf. Cistus 2 2
Fraxinussp 1 4
Juniperussp 2 5 4 25
Leguminosae 1
Rhammus/Phillyrea 1
Rosacea tpmaloidea 1
cf. Rosaceae (tp maloidea) 1
Salixsp 7 1 3 11 31
cf. Salix 3
Ang. no id 13 6 9 2 1 1
Gim. no id 2 1 1 1
Tot al 26 11 25 2 12 40 29
Unidentif. 5 3 15 1 1 27
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sabina, cope well with low temperatures and rocky oors. e rest of identied taxa are mainly riverbank species,
including Salix, Alnus and Fraxinus, which could point to a greater availability of these species in the surrounding
area. Lastly, Cistus, Leguminosae and Juniperus point to shrub-dominated landscapes.
Microvertebrates. e small vertebrate assemblage identied at Peña Capón accounts for one of the few
microfossil collections associated to the LGM time span recorded to date in Iberia8385, being the only one in the
whole Iberian hinterland. A total of 226 samples, collected throughout the whole sequence of human occupa-
tion at the site, have been analyzed. Of them, 205 contained identiable remains, among which we determined
shes, amphibians, snakes, birds, insectivores, lagomorphs and rodents. Only lagomorphs and rodents have
been determined to the species level (Table3 and Fig.8). Among the lagomorphs, the species Oryctolagus cunic-
ulus, the eld rabbit or European rabbit, dominates (see also Macrovertebrates section below). e dominant
species of rodents is Microtus arvalis, the common vole, while other species are poorly represented. ese are
Eliomys quercinus, the dormouse; indeterminate species of the genera Apodemus, the eld mouse; Terricola, the
“microtus” group of species with pitymyan rhombus; and Microtus agrestis, the eld vole or short-tailed vole. e
complete microvertebate taxonomic association per stratigraphic level is presented in Table3 according to the
NISP (number of identied specimens).
Microtus arvalis is an Arvicolinae (Cricetidae) that currently lives in temperate Europe and some regions of
western Asia. It is a strictly herbivorous species inhabiting open meadows, and it is well adapted to cold condi-
tions, as shown by its survival to the climatic changes of the LGM. Local extinctions of M. arvalis have been
recorded in northern and central Europe, as well as in Britain, due to post-LGM reforestation, which replaced
the dominant open grassland areas present during the LGM86. erefore, the signicant presence of M. arvalis
in levels 2a, 2b and 5 of Peña Capón points to environmental conditions that favored open and humid meadows,
the species’ favorite habitat.
More broadly, although samples from levels 4 to 6 have been only gathered in a 1 sq-meter test pit, thus biasing
the sampling throughout the sequence (Figs.S24 & S25), the distribution of taxa and the biodiversity recorded in
levels 1 to 6 at Peña Capón, points to levels 1, 2a and 3 as those presenting better climatic and habitat conditions
for Mediterranean species and open spaces, such as wet meadows. In turn, the lower biodiversity found in levels
4 to 6, as measured by the low number of taxa (Table3), and in general the lesser representation of micromam-
mals in these layers, could point to harsher environments, as shown by the presence of Microtus agrestis in level 6.
Macrovertebrates. Most of the 17,275 analyzed bone fragments throughout the Peña Capón sequence pre-
sent a high degree of fragmentation, and hence only 2.85% of them were identied to taxon (Table4). However,
although 97.15% of remains are thus indeterminate, 11.9% of them could be assigned to either small, medium
or large-sized animals (Table4).
Among the identied species, rabbit is the most abundant throughout the sequence, except in levels 5 and 6,
where horse is equally represented (Tables4 and 5). Concerning macrovertebrates, horse is the best-represented
taxon according to the MNI, and it also dominates the NISP in levels 2a, 2b, 5 and 6. Iberian ibex is the most
represented with respect to NISP in level 1, while red deer dominates level 3 and shows similar values to horse
in level 4 (Table5). Other identied species are the large bovid in levels 2a and 2b, roe deer in level 1, chamois
in levels 1, 2a and 2b, and badger and wildcat in level 2a (Table5). e three best-represented macroverte-
brates –horse, red deer and Iberian ibex– are related to dierent landscapes, being horse mostly adapted to
open environments, deer pointing to woods and ibex to rocky areas. However, the three of them are generalist
animals, capable of adapting to a range of climatic conditions and are not typical of either cold or temperate
Table 3. Microfossil remains (NISP) identied in Peña Capón. “#Samples” refers to the number of plastic bags
collected for the study. “#Taxa (S)” is the number of taxa identied in the total number of samples for each
level.
Taxon/level 1 2a 2b 3 4 5 6
Fishes 512
Amphibians 1 5 1
Serpentes 1 1
Birds 11 14 2 3 2
Insectivores 2
Rodentia indet 2 5 4
Eliomys quercinus 1
Apodemus 2
Microtus 2 8 1 1
Microtus arvalis 6 8 2
Microtus agrestis 2111 2
Terricola 7
#Taxa (S) 7 9 5 6 1 2 2
#Samples 50 72 45 20 9 20 10
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environments. us, in the absence of dental wear and stable isotope analyses (both to be conducted soon),
macromammal evidence is still uninformative in terms of reconstructing surrounding environments, mainly
due to the low number of remains still collected for layers 3 to 6. However, when considered together with pol-
len and micromammal results, macrovertebrate evidence is consistent with the interpretation of level 2a and
2b as showing temperate and humid environments. is is supported by the presence of roe deer, wildcat and
badger in these levels, as these species are adapted to wooded environments. Likewise, although available data
Figure8. Photographs of the occlusal surface of molars from selected species of small mammals collected at
Peña Capón levels 1–6. (a) le m1, m2 of Microtus arvalis (level 1); (b) M1, M2 of Microtus arvalis-M. agrestis
(level 2a); (c) upper M1 of Arvicolinae (level 2a); (d) right m1, m2 of Microtus agrestis (level 2a); (e) right m1,
m2 of Microtus agrestis (level 2a); (f) m1d of Microtus arvalis (level 2a); (g) m1d of Microtus agrestis, juvenile
(level 2a); (h) Microtus arvalis (level 2b); (i) Microtus arvalis (level 3); (j)–(n) lower molars of lagomorphs (jl
level 1; m level 3; n level 5).
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for levels 5 and 6 is still sparse, the relatively higher presence of horse, together with the decrease in the number
of rabbits, is consistent with the existence of dry environments and open landscapes as shown by the pollen and
micromammal analyses for these levels.
Overall, these patterns are consistent with those indicated by the study of faunal assemblages from level II
(Solutrean) and level III (Proto-Solutrean) as dened in the 1972 excavation (Supplementary Text S1), where
the preferred hunted animals were also horses, red deer and ibex25,87. Stable isotopes obtained from herbivore
teeth from those levels pointed to warm climate and temperate environments around 24kacal BP87, which
also ts current palaeoenvironmental evidence for levels 2a and 2b, but not for level 1 as especially shown by
pollen remains.
Mortality patterns and taphonomic analysis of the macrofaunal assemblages are provided in the Supplemen-
tary Information (Supplementary Text S7, TablesS6–S9, Fig.S26).
Table 4. Taxonomic representation of macrovertebrates remains at the Peña Capón sequence.
Taxon/level 1 % 2a % 2b % 3 % 4 % 5 % 6 % Tota l
Bos/Bison 0.0 2 0.0 1 0.0 0.0 0.0 0.0 0.0 3
Equus caballus 7 0.2 39 0.6 20 0.4 11 0.8 2 1.3 13 0.9 4 6.8 96
Cervus elaphus 2 0.1 20 0.3 5 0.1 12 0.9 2 1.3 0.0 0.0 41
Capreolus capreolus 1 0.0 0.0 0 0.0 0.0 0.0 0.0 1
Capra pyrenaica 9 0.3 17 0.3 5 0.1 3 0.2 0.0 1 0.1 0.0 35
Rupicapra pyrenaica 2 0.1 8 0.1 1 0 0.0 0.0 0.0 0.0 11
Meles meles 0.0 4 0.1 0 0.0 0.0 0.0 0.0 4
Felix silvestris 0.0 1 0.0 0 0.0 0.0 0.0 0.0 1
Carnivore indet 0.0 1 0.0 0 0.0 0.0 0.0 0.0 1
Oryctolagus cuniculus 128 3.6 97 1.6 28 0.6 51 3.7 4 2.6 13 0.9 4 6.8 325
Indet. large size 43 1.2 59 1.0 35 0.8 24 1.8 16 10.6 4 0.3 0.0 181
Indet. medium size 89 2.5 144 2.3 77 1.7 70 5.1 7 4.6 13 0.9 1 1.7 401
Indet. small size 382 10.7 551 8.9 291 6.5 213 15.6 7 4.6 34 2.3 1 1.7 1479
Indetermined 2899 81.3 5218 84.6 4033 89.7 984 71.9 113 74.8 1400 94.7 49 83.1 14,696
Tot al 3562 100.0 6161 100.0 4496 100.0 1368 100.0 151 100.0 1478 100.0 59 100.0 17,275
Table 5. Taxonomic representation of macrovertebrates remains at the Peña Capón sequence according to
NISP and MNI.
NISP/level 1 % 2a % 2b % 3 % 4 % 5 % 6 % Tota l
Bos/Bison 0.0 2 1.0 1 1.8 0.0 0.0 0.0 0.0 3
Equus caballus 7 4.7 39 20.1 20 36.4 11 14.3 2 25.0 13 48.1 4 50.0 96
Cervus elaphus 2 1.3 20 10.3 5 9.1 12 15.6 2 25.0 0.0 0.0 41
Capreolus capreolus 1 0.7 0.0 0.0 0.0 0.0 0.0 0.0 1
Capra pyrenaica 9 6.0 19 9.8 3 5.5 3 3.9 0.0 1 3.7 0.0 35
Rupicapra pyrenaica 2 1.3 8 4.1 1 1.8 0.0 0.0 0.0 0.0 11
Meles meles 0.0 4 2.1 0.0 0.0 0.0 0.0 0.0 4
Felix silvestris 0.0 1 0.5 0.0 0.0 0.0 0.0 0.0 1
Carnivore indet 0.0 1 0.5 0.0 0.0 0.0 0.0 0.0 1
Oryctolagus cuniculus 123 85.9 97 51.5 25 45.5 51 66.2 4 50.0 13 48.1 4 50.0 325
Tot al 144 100.0 191 100.0 55 100.0 77 100.0 8 100.0 27 100.0 8 100.0 518
MNI 1 % 2a % 2b % 3 % 4 % 5 % 6 % Tot al
Bos/Bison 0.0 1 4.8 1 9.1 0.0 0.0 0.0 0.0 2
Equus caballus 1 7.7 3 14.3 3 27.3 2 33.3 1 33.3 1 33.3 4 80.0 15
Cervus elaphus 1 7.7 2 9.5 1 9.1 1 16.7 1 33.3 0.0 0.0 6
Capreolus capreolus 1 7.7 0.0 0.0 0.0 0.0 0.0 0.0 1
Capra pyrenaica 1 7.7 1 4.8 2 18.2 1 16.7 0.0 1 33.3 0.0 6
Rupicapra pyrenaica 1 7.7 4 19.0 1 9.1 0.0 0.0 0.0 0.0 6
Meles meles 0.0 1 4.8 0.0 0.0 0.0 0.0 0.0 1
Felix silvestris 0.0 1 4.8 0.0 0.0 0.0 0.0 0.0 1
Oryctolagus cuniculus 8 61.5 8 38.1 3 27.3 2 33.3 1 33.3 1 33.3 1 20.0 22
Tot al 13 100.0 21 100.0 11 100.0 6 100.0 3 100.0 3 100.0 4 100.0 60
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Archaeological assemblages. As previously reported20,25,26,88, Peña Capón hosts a sequence of recurrent
occupations of hunter-gatherers bearing Solutrean and pre-Solutrean technocomplexes which, to date, has no
parallel in the whole Iberian Meseta43. Together with anthropized faunal remains, a limited number of bone
tools, ocher fragments and at least two replaces in levels 2a and 2b, lithics account for the most abundant
archaeological material (Supplementary Figs.S8–S18). Foliate lithic armatures obtained through bifacial and
unifacial invasive at retouch are found throughout levels 1 to 3 (Fig.9). Considering the classic typology-based
chronological framework of the Solutrean50,8993, the presence in all these levels of bifacial laurel leaf points and
the absence of typically Upper Solutrean types, such as shouldered or barbed-and-tanged points (only recorded
at Peña Capón as surface nds), enable us to place them in the Middle Solutrean (Supplementary Dataset 5). e
dating of levels 1, 2a and 2b between 23.8 and 25.0kacal BP is roughly coherent with a Middle Solutrean attri-
bution. However, level 3, dated between 24.7 and 25.3kacal BP, best ts the initial phase of the Solutrean. is
partial incoherence supports the questioning of the traditional type-fossils as precise temporal markers64,94,95.
Anyhow, typically Lower Solutrean types, such as pointes à face plane, are not found in the Peña Capón sequence,
a circumstance that parallels the nearly absence of this phase/facies in central and southern Portugal, where the
Proto-Solutrean is followed by the Middle Solutrean64,93,96.
Since levels 4–6, dated between 25.1 and 26.1kacal BP, have been excavated in only 1 sq meter, their archaeo-
logical assemblages are still scarce. Although no index fossils or signicant technological strategies have been rec-
ognized in these levels, the absence of foliate armatures, the higher presence of quartz and bladelets as compared
to levels 1–3 (Supplementary Dataset 5), and their chrono-stratigraphic position, allow us to securely relate them
to pre-Solutrean human occupations (Fig.10). Based on the assemblages from the 1972 excavation at the site
(Supplementary Text S1), a Proto-Solutrean component with Vale Comprido points (level III) (Supplementary
Fig.S3), and a potentially Gravettian occupation (level IV) were described20,25. According to new radiocarbon
determinations, these two levels (III and IV as dened in 1972) are currently dated to 25.6–24.9kacal BP and
26.3–25.7kacal BP (Supplementary TableS1), thus fairly mirroring levels 4–5 and level 6 of the new excava-
tions respectively (Table1 and Fig.6). However, a ne-grained correlation between levels from the old and new
excavations has not been possible to date, since no clear Proto-Solutrean or Gravettian traits have been identied
in either level 4, 5 or 6, or a combination of them. erefore, we prefer to provisionally describe archaeologi-
cal assemblages of levels 4–6 just as “pre-Solutrean. Yet, the presence of a Proto-Solutrean component at Peña
Capón remains without doubt, and is now reinforced by obtained radiocarbon dates in levels 4–6 matching
the Proto-Solutrean time span, as recorded in Portugal, between 26 and 25kacal BP53,64,96. Furthermore, this
dating supports the triggering of the Proto-Solutrean as related to the rapid climate and environmental changes
caused by HS 2, and especially to a decrease in vegetation cover and forest diversity5254, as also reinforced by
palaeoecological data obtained at Peña Capón.
Discussion
Site formation processes. e site has been previously interpreted as a result of the contribution of uvial
sedimentation and fallen blocks from the roof26. Fluvial origin is conrmed because of the homogeneity in grain
size of the ne sediments, which suggests the lack of inuence of debris ows coming from closer alluvial fans
or slope deposits. e few sub-rounded to rounded quartzite and ne siltstone gravel found are consistent with
the accumulation by uvial transport. However, the downslope geometry of the deposit suggests that it is not
that much a classic at ood plain terrace deposit but the deposition during oods on the hill slope. e insitu
preservation of the deposits can be explained by protection from reworking and erosion by the corridor formed
between the rock wall and verticalized dolostone layers and big gravitational blocks in the external side of the
site. e sediment low heterogeneity observed is just due to variations in organic matter and secondary carbon-
ate content and not to changes in the sedimentary environment. e variation in χlf over some of the layers
makes it more likely that the higher magnetic signal of the darker ones has been induced by burning. Besides
uvial deposition, physical disintegration of the rock shelter wall and gravitational processes have provided
limited amounts of nes and frost-shattered debris contributing to accumulation of the archaeological sequence.
Apart from the erosion of the uppermost levels of the archaeological sequence, micromammal burrowing and
local sediment disturbing in the top of level 3 and the lower part of level 2b, the main post-depositional processes
are just the dissolution of primary carbonate grains and the intergranular precipitation of secondary carbonate,
sometimes forming small nodules. ese calcitic pedofeatures testify the lixiviation or carbonate leaching of
larger blocks and ne sediments. e dominating channel and burrow microstructure testies to intensive rooting
and burrowing activity of mesofauna but only on a microscale. is bioturbation may have destroyed the primary
depositional fabric of the nes but no signs of mixing between levels have been recorded. Overall, levels 5 to 1
rapidly accumulated over a period of about 2,000years and represent an excellently preserved sediment sequence.
Human–environment interactions and population dynamics during the HS 2 in the Iberian
hinterland. Although the whole sequence of human occupation recorded to date at Peña Capón occurred
during a stadial phase (GS-3), it included periods of both cold and relatively warm climates, corresponding
to arid and more humid environments respectively. e warmest period corresponds to the bulk of the Solu-
trean occupations between 25.3 and 24.0kacal BP (levels 2a, 2b and 3), and is characterized by deciduous oak
groves enriched with numerous mesophilous trees, including beech (LPAZ-2), and the presence of animals well-
adapted to wooded environments, such as roe deer, wildcat and badger. is picture partially supports previous
data from stable isotopes pointing to warm climate and temperate conditions at the site87, but it is now clear that
this period was limited to part of the Solutrean (and not the Proto-Solutrean) and occurred well within GS-3
(and not during GI-2), including part of HS 2.
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e Peña Capón site hosts the oldest Upper Paleolithic presence recorded so far in central Iberia, starting at
26.1kacal BP as dated in level 6. is presence, currently accounting for the rst peopling of the deep interior
of the peninsula by modern humans, occurred during the central moments of HS 2, and hence during a cold
and arid global period68. is harsh climate/environment is associated with the pre-Solutrean occupations of
Peña Capón (levels 4, 5 and 6), dated between 26.1 and 25.0kacal BP. At these levels, data point to the presence
of pine forests at higher altitudes and evergreen oak and juniper woodlands at lower ones, as well as to shrub
and herb communities dominated by cryoxerophytic and heliophilous/cryophilous elements (LPAZ-1). Faunal
associations, dominated by horse, show a generalized decrease in biodiversity as compared to levels 2a, 2b and 3,
and include the presence of the cold-adapted Microtus agrestis. is paleoecological framework is consistent with
the general context of increasing aridity and decrease in vegetation cover for the HS 2 in Iberia, as recorded both
in continental97,98 and coastal archives99101 (Fig.11). Furthermore, it poses a marked contrast with the warm/
Figure9. Solutrean lithic assemblages. Level 1: 1–3. Level 2a: 4–6. Level 2b: 7–9. Level 3: 10–12.
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humid conditions recorded in the Solutrean levels, except for level 1, which shows again a cold/arid landscape
(LPAZ-3) between 24.1 and 23.8kacal BP.
Additionally, regional climatic and environmental proxies documented in inland Iberia point to harsh con-
ditions during the time in which Peña Capón was occupied by hunter-gatherers. A pronounced period of loess
deposition recorded in the Upper Tagus basin between 25.9 ± 2.4ka and 23.2 ± 1.6ka BP, interpreted as evidence
of arid conditions40,41, matches the time span of the human presence at Peña Capón (Fig.11). Also matching the
rst occupation of the site by humans, the maximum extent of glaciers recorded in the Iberian Central System
range, dated at 26.1 ± 1.3ka BP and backed by a local speleothem record, points to high precipitation rates within
a cold period102,103 (Fig.11). Further south, pollen data from the Fuentillejo maar lacustrine record and the TD
core at Tablas de Daimiel National Park point to a cold period at 25–23ka BP, characterized by Juniperus and
xeric vegetation dominance104,105. All these data demonstrate that the rst human settlement of this area of central
Iberia occurred during a cold and mostly arid period.
ese results confront traditional views and recent models positing that harsh climatic and environmental
conditions hampered the human occupation of inland Iberia during most of the Upper Paleolithic. Geoarchaeo-
logical and paleoecological evidence gathered at Peña Capón demonstrate that this area of the Iberian hinterland
was recurrently occupied during both temperate/humid and cold/arid periods, and thus regardless of climatic
and environmental variability. Furthermore, although no other Solutrean or pre-Solutrean sites have been clearly
identied thus far in the surrounding area of the site, the nearby caves of El Reno, El Cojo (both 9km away),
and Los Casares (76km away) (Fig.1D and Supplementary Fig.S1), were probably occupied by humans at the
same time as Peña Capón. ese sites host pre-Magdalenian rock art depictions, including cold-adapted fauna,
which have been related to Solutrean or Gravettian times based on stylistic grounds and superimpositions with
other images43,106. is, together with the nearby (but still poorly dated) Solutrean cluster of the Madrid basin107
strongly suggests that the human occupation sequence recorded at Peña Capón was not the product of isolated
occasional visits to this region. Rather, it is increasingly clear that it was part of an organized settlement, estab-
lished perhaps throughout the Tagus basin, and covering a prolonged sequence of time during the LGM, which
in Peña Capón started at least in the HS 2 (Fig.1C,D). Results of lithic raw material sourcing showing mobility
between some Gravettian sites of the Côa valley (inner northeastern Portugal) and inner areas of the Northern
and Southern Mesetas23,108,109, reinforces this hypothesis.
However, besides Peña Capón, pre-Solutrean occupations in inland Iberia remain absent for the central area
of the peninsula, and they are very sparse in the rest of the plateau. Only at the western limit of the Northern
Meseta there is sound evidence of Upper Paleolithic prior to 26kacal BP. Here, several Gravettian sites at the
Côa valley are dated around 28kacal BP23, and the single site of Cardina-Salto do Boi have recently yielded an
Figure10. Pre-Solutrean lithic assemblages. Level 4: 1–3. Level 5: 4–5. Level 6: 6–7.
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OSL date as old as 33.6 ± 2.0kacal BP associated to Aurignacian assemblages14 (Fig.1B,C). Although these dates
shorten the gap of human occupation aer the Neandertal disappearance for the western margins of the plateau,
in the rest of the Iberian hinterland, and especially in the very center, there is still a hiatus of 16,000years devoid
of human populations between 42 and 26kacal BP40,110,111.
Figure11. Global and regional Iberian climatic/environmental proxies from 35 to 12.5kacal BP (modied
from [43: g.1] and [40: g.3] in relation to the modeled sequence of human occupation recorded at Peña
Capón. A: δ18O record of the NGRIP ice core, with numbers and grey bars referring to Greenland Stadials67,
and indication of the LGM and HS2 chronology68,69. B: Sea Surface Temperature reconstructions of marine
drilling core MD95-2043 (Alborán Sea)99, and Heinrich Events detected in the same core. C: Percentage
of temperate forest pollen in core MD95-2043100. D: Sea Surface Temperature reconstructions of marine
cores MD95-2042 and SU81-18 (Atlantic)101 and Heinrich Events detected in the same cores. E: Percentage
of temperate forest pollen in cores MD95-2042 and SU81-18. F: Main loess deposition periods and their
sedimentation rates recorded in the Upper Tagus Basin (ochre bars)40 and Maximum extension stages of glaciers
recorded in the Iberian Central System Range (blue bars)102. G: Estimated duration of the human occupation
recorded at Peña Capón, based on the Bayesian Model 2 (Fig.6) as calculated by the ‘date’ command in Oxcal.
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Conclusions
Given the current archaeological record, the argument that climate and environmental variability hampered
the settlement of the Iberian hinterland by modern humans remains a valid hypothesis for the rst phases of
the Initial Upper Paleolithic in most of the inland territories. Yet, the fact that the rst modern human pres-
ence recorded thus far in the deep interior occurred precisely during a cold and arid period of Heinrich Stadial
2, allows us to keep working on the hypothesis that the rst settlement of these regions occurred earlier and
regardless of climatic and environmental variability. As systematic eldwork campaigns are still few in inland
Iberia, ongoing and future research is very much needed to keep testing this and other competing hypotheses.
Anyhow, the results presented in this paper show that the inland Iberian highlands were not an exception to the
wide variety of landscapes to which Paleolithic hunter-gatherers were capable of adapting. Although it is well
documented that large regions of Europe remained depopulated during long periods of the Last Glacial, it has
also been thoroughly demonstrated that, given a certain availability of herbivore and plant resources, humans
were always willing to expand and prosper anywhere, worldwide112117.
Methods
Methods for excavation, spatial recording and sampling are presented in Supplementary Text S2. Methods con-
cerning geomorphology, sedimentology and micromorphology are provided in Supplementary Text S4.
Radiocarbon dating and Bayesian modeling. In CologneAMS, bone samples were processed by col-
lagen extraction and charcoals were AAA (Acid–Alkali–Acid extraction) processed according to sample prepa-
ration described by Rethemeyer etal.118. At ORAU, extraction, purication, and dating of bone collagen were
carried out following ultraltration methods119, while dating of charcoal was undertaken using an ABOx-SC
pretreatment120.
For Bayesian modeling we used the OxCal 4.4 online soware121 and the most recent terrestrial radiocarbon
curve, IntCal20122, to combine the radiocarbon likelihoods with the stratigraphic position of all samples. Since
each sample was three-dimensionally recorded during excavation, relationships between samples and levels
(including depth within a given level) were included within the Bayesian model as prior information. We used
a General t-type Outlier Model123 with a resolution of 20years and assigned 5% chances for each determination
to be an outlier, as it is commonplace in recent research. In OxCal, commands and parameters are written in a
C++ CQL (Command Query Language)121, and Model CQL codes are provided in Supplementary Text S6. e
commands used to constrain the dated events in chronological order, group them within a given stratigraphic
level, and calculate a start and end boundary (Probability Distribution Functions or PDFs) to bracket each
archaeological episode, have been ‘sequence, ‘phase’ and ‘boundary’ respectively121. Furthermore, we used the
date’ command to query further on the accuracy of the time spans of each archaeological layer and the whole
sequence of human occupation at Peña Capón.
In order to best identify outliers in the sequence, we run the model in two stages following the agreement
index method as described by Bronk Ramsey123. us, aer running a Preliminary Model (1) including all
obtained radiocarbon measurements, a Final Model (2) was constructed by removing results with less than 60%
agreement (matching those with posterior probabilities of > 5%), interpreted as potential outliers. Both models
were run 3 times at > 3 million iterations and yielded no signicant variation in their posterior results, thus
showing that they were reproducible and the convergence values were high.
Palynological analysis. During the 2015 season, eight sediment samples of 5 square cm were extracted
for pollen analysis from the southern prole of square 2B (levels 1top, 1base, 2a, 2b, 4, 5top, 5base and 6)
(Supplementary Fig.S23) and one more from the western prole of the same square (level 3) (Supplementary
Fig.S20). Extraction followed standardized techniques for archeological sites124,125. e nine collected samples
were prepared for pollen analysis (10g per sample) at the CSIC labs (Madrid) following standard methods
in archaeopalynology124, using treatment with HCl, 10% KOH, HF and concentration with oulet liquor,
although acetolysis was not carried out to allow the identication of any contamination by modern pollen. e
nal residue was suspended in glycerin and counted until a pollen sum of 250 pollen grains was reached. Count-
ing was undertaken using a Nikon Elipse 50i light microscope at × 400 magnication. Pollen grains were identi-
ed according to Moore etal.126 and Reille127 at the lowest currently possible taxonomical level. Pinus nigra-type
pollen grains were categorized following measurements in Desprat etal.128. Pollen percentages were calculated
using a pollen sum excluding indeterminable pollen grains (i.e., those that were broken, concealed, corroded,
crumpled or degraded), as well as Asterioideae, Carduoideae and Cichorioideae with possible zoophily125, and
presented as bars in a pollen diagram (Fig.7). To establish the zonation of the pollen sequence, we tested several
divisive and agglomerative methods with the program IBM SPSS Statistics 21. Based on the ecological meaning
of the obtained zones, three local pollen assemblage zones (LPAZ-1 to LPAZ-3) were constructed on the basis
of agglomerative constrained cluster analysis of incremental sum of squares (Coniss) with square root trans-
formed percentage data129. e number of statistically signicant zones was determined by using the broken-
stick model130. Tilia and TGView131,132 and CorelDraw soware were used to plot the pollen diagram.
Anthracological analysis. Charcoal remains were sampled by hand during eldwork and by otation in
the laboratories of the University of Alcalá and the Museo Nacional de Ciencias Naturales (Madrid). 131 sam-
ples from levels 1 to 6 have been studied in the Archaeobotanical laboratory of the Autonomous University of
Barcelona and the Environmental Archeology laboratory of the CSIC (Madrid). A total of 197 fragments of
carbonized wood were localized, of which 145 were identied to taxon. Identication of taxa was carried out fol-
lowing standard procedures. e anatomical patterns of each wood species were observed along three sections
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(transversal, longitudinal tangential, and longitudinal radial) using a reected light microscope equipped with
light eld/dark eld and objectives of 50 ×, 100 ×, 200 × and 500 ×. Archeological samples have been compared
with modern woods as well as with wood anatomy atlases133135.
Microvertebrate analysis. In order to collect all small bone and teeth fragments from the fossil assem-
blages, all excavated sediments, bagged by sector, layer and stratigraphic level, were water-screened using
superimposed 1.5mm, 1.0mm, and 0.5mm-mesh screens, both at the site and in the labs of the University of
Alcalá and the Spanish National Natural Sciences Museum (Madrid). A total of 226 bags (3–1kg each) were
wet-screened and 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. e resulted
collections of fossils were then sent to the Department of Earth Sciences of the University of Zaragoza, where
assemblages were examined, photographed and stored. 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, were covered with particles of sediment that impeded the visual analysis. is anatomical
region is needed pristine for the good classication and the morphometric analysis of small mammals. Drawings
were made aer photographs taken with an Olympus SZ61 microscope with a camera attached to it. Images and
measurements were taken with the camera and the LCMicro soware provided for the Olympus equipment.
Classication of small mammals into species was based on the morphology and biometry of the occlusal
surface of the molars, following general criteria of systematic paleontology136138. In each sample, we counted the
number of skeletal elements, mainly dentition, and calculated the minimum number of Identied species (NISP).
Interpretation of the microvertebrate assemblages in palaeoenvironmental terms is based on the taxonomic
association present in each archaeological layer and their ecological preferences, as well as their temperature
and humidity limitations as a whole assemblage.
Zooarcheology. 17,275 faunal remains from levels 1 to 6 of Peña Capón, coming mostly from the 2015
season, were subject to zooarcheological and taphonomic analyses. Studied remains included both identia-
ble and unidentiable fragments and their taxonomic identication were based on reference material held at
the Prehistory Department of the Complutense University of Madrid (Spain). When the identication was not
feasible, epiphyses, axial and sha 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) [see110].
e estimation of NISP (Number of Identied Specimens) and MNI (Minimum Number of Individuals)
were used to quantify the faunal remains and determine the most appropriate features of the faunal taxonomic
distribution. NISP determination follows Lyman139, whereas MNI is based on Brains140 method, which uses bone
laterality and estimated age. Furthermore, skeletal proles and MNI consider sha thickness, section shape and
medullar surface properties141. 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, sensu142), 3) upper appendicular
limbs (humerus, radius, ulna, femur, patella and tibia) and 4) lower appendicular limbs (metapodial, carpals,
tarsals, phalanges and sesamoideal).
Lithic analysis. Aer manual cleaning and removal of adhering concretion (Supplementary Fig.S17), lithic
artifacts were studied at the Prehistory Laboratory of the University of Alcalá. We followed the chaîne opératoire
or ‘operational sequence’ approach [e.g.143,144], combined with the ‘Technological organization’ approach aimed
at examining links between technology and paleoenvironmental change [e.g.145147]. We assigned each lithic arti-
fact to one of the three chaîne opératoire stages commonly recognized in the literature (Supplementary Dataset
5). us, cortical akes, preparation products and tested cores were assigned to the initialization stage or phase
I; raw blanks, core maintenance by-products, thinning akes and productive cores to the exploitation stage or
phase II; and retouched blanks, retouching akes and exhausted cores to the consumption and abandonment
stage or phase III. e study of bifacial reduction sequences aimed at the production of foliate armatures fol-
lowed methods described in Alcaraz-Castaño etal.107 and were ultimately backed in experimental intknapping
works148150.
Data availability
e authors declare that all data supporting this research are available within the paper, its Supplementary
Information and Supplementary Data les. e Peña Capón archaeological assemblages are housed in the His-
tory and Philosophy Department (Prehistory Area) of the University of Alcalá and the Museo de Guadalajara
(Guadalajara, Spain). Both repositories are accessible for all researchers upon request.
Received: 16 April 2021; Accepted: 9 July 2021
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Acknowledgements
is research was carried out in the context of the ERC MULTIPALEOIBERIA project, funded by the European
Research Council (ERC-2018-STG-805478), and the PALEOINTERIOR project, funded by the Spanish Min-
istry of Science and Innovation (HAR2017-82483-C3-3-P). e sedimentological, micromorphological and
part of the radiocarbon analyses were funded by subproject C1 of the CRC 806 “Our way to Europe” (DFG,
German Research Foundation) and the Marie Curie Intra European Fellowship project ‘Hiatus LPleis Iberia
(FP7-2013-IEF-628179). We gratefully acknowledge contributions made by the wide excavation and laboratory
team. Fieldworks at Peña Capón were authorized by the Dirección General de Cultura de la Junta de Comunidades
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de Castilla–La Mancha (Spain) (Exp. 14.0955-P4 and Exp.: 19.248) with permission from the Confederación
Hidrográca del Tajo.
Author contributions
M.A.C. G.C.W and J.J.A.G. designed research. M.A.C. and J.J.A.G. directed eldworks. M.A.C. devised the paper
and coordinated research. M.A.C., J.J.A.G., M.A.H., S.C.J, F.C., G.C.B., M.K., J.A.L.S., L.L., S.P.D., R.P., M.R.A.,
G.C.W and J.Y. performed research and analyzed the data. M.A.C., J.J.A.G., M.A.H., G.C.B., M.K., L.L., J.A.L.S.,
M.R.A. & J.Y. wrote the manuscript and prepared gures. All authors contributed to the manuscript, revised
dras, and approved the nal version.
Competing interests
e authors declare no competing interests.
Additional information
Supplementary Information e online version contains supplementary material available at https:// doi. org/
10. 1038/ s41598- 021- 94408-w.
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The Iberian Peninsula is considered one of the most well-suited regions in Europe to develop studies on the relationship between environmental changes and human adaptations across the Late Pleistocene. Due to its southwesternmost cul-de-sac position and eco-geographical diversity, Paleolithic Iberia was the stage of cyclical cultural/technological changes, linked to fluctuations in climate and environments, human demographics, and the size, extension, and type of social exchange networks. Such dynamics are particularly evident during the Last Glacial Maximum (LGM) timeframe, with a series of innovations emerging in the archaeological record, marking the transition between the traditionally defined Gravettian, Proto-Solutrean, Solutrean, and Magdalenian technocomplexes. Stemming from a workshop organized in Erlangen in 2019 on “The Last Glacial Maximum in Europe - state of knowledge in Geosciences and Archaeology”, this paper presents, in the first part, an updated review on the paleoenvironments and human adaptations across four macro-regions (Northern, Inland, Mediterranean, and Western Atlantic Façade) in Iberia during the LGM; and, in a second part, a discussion on the pronounced inter-regional variability, unresolved research questions, and the most promising research topics for future studies.