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This study presents an exceptional collection of 54 Late Pleistocene human remains that correspond to at least three Neanderthal individuals from Simanya Gran, the main gallery of Cova Simanya, located in the northeastern Iberian Peninsula. The collection comprised 53 unpublished remains that were unearthed during the 1970s and an additional tooth discovered during 2021 excavations. The specimens represent an adult with a small stature, a periadolescent aged approximately 11.5 years, and an immature individual aged approximately 7.7 years, thus offering a more complete demographic perspective. The collection encompasses diverse anatomical parts including upper and lower dentition, mandible, vertebrae, and limb bones from both the upper and lower extremities. Attempts to extract aDNA were unsuccessful. Renewed archaeological investigations at Cova Simanya have facilitated the reevaluation of the original stratigraphic context of these remains, leading to the discovery of the additional tooth, aligning with the periadolescent individual. This assemblage is currently the most extensive Neanderthal collection from the northeastern Mediterranean Iberia, offering invaluable insights into the morphology and evolutionary trajectory of Late Pleistocene hominins. Hence, Simanya Neanderthals will enhance our understanding of Neanderthal demographics and evolution, paving the way for an in-depth examination of the morphological diversity and evolutionary context of Iberian Neanderthals.
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Neanderthals exhibited distinct genetic histories and cultural adaptations along time and space. In the Iberian Peninsula, the Iberian Mediterranean region holds significant importance in understanding Neanderthal evolution and variability, and its archaeological record provides valuable insights into their biological and cultural development as well as their eventual disappearance during marine isotope stage (MIS) 3.
Middle Paleolithic Mousterian sites can be found along the Mediterranean coast from Gibraltar to the Pyrenees, with Neanderthal remains being documented at several locations (
These data suggest that Mediterranean Iberia was continuously inhabited by Neanderthals from MIS-7 onwards, with northeastern Iberia being an especially dense Mousterian cluster (
Herein, we present a novel Neanderthal assemblage from Cova Simanya (Sant Llorenç Savall, Barcelona), first recognized during the analysis of a museum collection and constituting the largest sample documented so far in northeastern Iberia. This discovery has led to a new research project that aimed at reconstructing the genesis, context, and paleobiology of the Neanderthal sample, including renewed excavations at Cova Simanya. These works have provided a robust archeological context for the uncontextualized “Museum Collection” including additional Neanderthal remains.
Simanya’s assemblage may offer insights into the late Neanderthals’ apomorphic traits and their congruence with other known samples, while also raising questions about the genetic diversity and population turnovers within the Iberian Peninsula (
Cova Simanya is a karstic system (UTM ETRS89 417517–4614066, 885 masl) opened in Montcau mountain, running through the Paleogene conglomerates forming the Sant Llorenç del Munt Massif, on the eastern margin of the Ebro Basin, and has over 300 m of graticulated galleries. The Font del Llor stream cuts the system, creating three consecutive entrances known as Simanya Triangle, Simanya Petita, and Simanya Gran (
Simanya Gran is the larger sector of Cova Simanya, comprising the Western Gallery (WG), Main Gallery (MG), Pas de l’Oca Gallery (POG), and the inner part of the Main Gallery (IMG) (
The “Museum Collection” was collected by an amateur, Mr. Miguel Aznar, in 1978–79 and contains a combination of two distinct sources: 1) surface findings and 2) remains from a small excavation. The surface findings include isolated human and animal bones, pottery, and chert fragments. The excavation yielded human and faunal remains, as well as a few samples of charcoal, sediment, and carbonate crusts. The excavation was performed in a breccia deposit of reddish sediments with abundant charcoals located in the IMG sector. No pottery or any other Holocene remains were found during the excavation. The collection containing 143 osteological remains was donated to the Museum of Archaeology of Catalonia in 1986. In 2021, Mr. Aznar delivered two additional bags to us containing small and mostly undiagnostic bone fragments that are not included in the original donation.
Despite being layered in the reddish clay-like sediment, the human bone collection exhibited a notable state of preservation. However, the presence of modern fractures and adhesive-induced alterations from the 1970s necessitated a thorough evaluation of their conservation status. Following the evaluation, a series of measures were taken to inhibit further deterioration. Cleaning procedures involving water and soft brushes were applied, and outdated adhesives were removed and substituted with Paraloid® B-72, a durable and reversible acrylic resin. This resin was also utilized to mend modern fractures and consolidate the overall structure, ensuring the assemblage’s longevity and integrity for future study and preservation.
The 2020–2022 excavation programs targeted Simanya Gran and aimed at identifying the original context of the “Museum Collection” and evaluating the potential of the Pleistocene deposits in the cave. In 2020, five test pits were excavated to reconstruct the longitudinal stratigraphic development along the MG (S1, S2, and S3) and WG (S4–S5). In 2021, a more extensive excavation was carried out in the IMG sector (
Eleven bone samples, both anthropized and non-anthropized remains, and five charcoals (
All 14C ages were calibrated with the IntCal20 calibration curve (
An attempt to extract aDNA from Simanya Neanderthals was made by sampling a tooth and a phalanx from the “Museum Collection” through shotgun sequencing (Illumina NovaSeq 6000) and mitochondrial DNA (mtDNA) capture with Neanderthal-specific baits (Arbor Biosciences).
The litho-stratigraphic sequence documented so far for the MG and IMG sectors of Simanya Gran contains a primary Pleistocene sequence capped by a thin Holocene deposit, 5–30 cm thick, which is documented from the entrance up to 53 meter. From that point of the MG to the end of the IMG, the Pleistocene units are uncapped (
Unit SG4 contains at least 60 cm of pure, dark reddish-brown clays with scarce coarse to very coarse gravels, mainly limestone and slate, documented at S1, S2, S3, and the IMG, and is eroded by Unit SG3. Unit SG3 comprised several coarsening upward sequences (silt/clay to coarse sand) that are cut and filled by at least four poorly sorted (silts to coarse sand) channel deposits. The thickness of Unit SG3 varies from 5 to 45 cm, and it has been documented in S1, S2, S3, and the IMG. In the S1 section of the MG sector, Unit SG3 is conformably overlain by Unit SG2-a, an approximately 50 cm-thick matrix-supported conglomerate of slate and limestone clasts that are more abundant toward the top, while in the S3 section, it is overlain by Units SG2.3-to-1, a succession of brown clays, a poorly sorted yellowish sandy deposit, and a very fine gravel conglomerate, and by the S1 flowstone. The Pleistocene sequence ends with Unit SG1, a reddish-brown, matrix-supported polygenic limestone, sandstone, and slate conglomerate with an erosive base that has been documented exclusively at the IMG.
Three archaeological layers have been documented so far within the litho-stratigraphic sequence at the MG–IMG sector. Layer P1 corresponds to the Holocene phase and yields artifacts from the Late Neolithic to the Late Antiquity, including scattered funerary evidence dated back to the 5th century AD, in a condensed sub-superficial sedimentary palimpsest (
Calibrated dates at 68.3% and 95.4% probability are provided in
Samples selected for 14C dating pretreated at the University of Bologna and AMS results.
Bologna Code | Sample ID | Unit | Sector | Material | Species | Collagen (mg) | Collagen (%) | AMS Code | d13C | F14C | C:N | 14C age | 1s Error | Cal BP age (68.3%) | Cal BP age (95.4%) |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
BRA-6108 | SG21.06.39 | O300-PO400 | 301-70 | Bone | Caprini | 4.8 | 1% | MAMS 56486 | −18.21 | 0.0088 | 2.89 | 38009 | 410 | 42440–42110 | 42620–41930 |
BRA-6109 | SG21.11.08 | O300-PO400 | 301-71 | Bone | Suidae | 0.7 | 0% | — | |||||||
BRA-6110 | SG21.06.78 | O300-PO400 | 301-82 | Bone | Caprini | 0.6 | 0% | — | |||||||
BRA-6111 | SG21.10.58 | O300-PO400 | 301-82 | Bone | Caprini | 0.2 | 0% | — | |||||||
BRA-6112 | SG21.12.10 | O300-PO400 | 301-82 | Tooth | Caprini | — | — | — | |||||||
BRA-6113 | SG21.10.30 | O300-PO400 | 301-83 | Tooth | Caprini | — | — | — | |||||||
BRA-6114 | 121_1978 | Museum Collection | IMG | Tooth | Ursidae | 0.8 | 0% | — | |||||||
BRA-6115 | SG-CA-125 | Museum Collection | IMG | Tooth | Ursidae | 0.5 | 0% | — | |||||||
BRA-6116 | SG-CA-10 | Museum Collection | IMG | Tooth | Ursidae | 0.5 | 0% | — | |||||||
BRA-6117 | SG-CA-62 | Museum Collection | IMG | Bone | Indeterminate | 0 | 0% | — | |||||||
BRA-6118 | SG-CA-137 | Museum Collection | IMG | Bone | Caprini | 1.2 | 0% | — | |||||||
Carbon (mg) | Carbon (%) | ||||||||||||||
BRA-6106 | SG21.11.53 | O300-PO400 | 301-72 | Charcoal |
|
33.3 | 35% | MAMS 56487 | −22.61 | 0.0009 | 137.59 | >49000 | — | — | — |
BRA-6107 | SG21.03.05 | O300-PO400 | 301-86 | Charcoal |
|
50.8 | 49% | MAMS 56488 | −23.43 | 0.0012 | 129.79 | >49000 | — | — | — |
BRA-6264 | SG-CA-200 | Museum Collection | IMG | Charcoal |
|
4.6 | 7% | — | |||||||
BRA-6265 | SG-CA-210 | Museum Collection | IMG | Charcoal |
|
11.7 | 21% | MAMS 62623 | −24.17 | 0.0020 | 87.05 | >49000 | — | — | — |
BRA-6266 | SG-CA-207 | Museum Collection | IMG | Charcoal |
|
6.2 | 16% | MAMS 62624 | −23.79 | 0.0013 | 172.17 | >49000 | — | — | — |
A collection of 53 human remains was recovered by Mr. Aznar in 1978–79, to which an upper third molar (SI-60) found
Foremost Neanderthal remains found at the Simanya Gran gallery from Cova Simanya (Barcelona, Spain). Skeletal remains are distributed according to their attribution to specific individuals.
At least three individuals can be identified in the sample. The most complete one is defined by a complete left humerus SI-1 (
SI-7 is an upper left P3 at the Rc growth stage [according to the work of
Two other immature remains are also preserved in the sample: SI-29 is a hand first phalanx of ray V, whose size is like Juvenile 1 from El Sidrón (
The size of lower dentition I1 (SI-3) and M3 is relatively small, while P3 (SI-5) falls at the upper limit and the upper I1 (SI-45) in the middle of the Neanderthal distribution, while the mesiodistal crown diameter of the I2 (SI-4 and SI-31) falls well above the Neanderthal mean [according to the work of
In order to assess the location of the Simanya Gran specimen SI-1 in the morphospace (
Scatter plot of the PCA (PC1–PC2) shape space of the distal humerus performed on 18 landmarks and a large comparison sample from the work of
Taxonomically, the human remains from Simanya Gran present a large series of Neanderthal apomorphic features distributed across different anatomical systems. Upper central incisors are large with a robust root, a strong dental tubercle, and characteristic flat wear (
Ultimately, no positive results for aDNA were obtained, neither for shotgun sequencing nor for the mtDNA capture.
Excavations at Simanya Gran revealed Pleistocene fossil-bearing deposits, primarily preserved in the layer O300-PO400 from the IMG and likely dismantled from the cave entrance. Its sub-superficial position aligns with Mr. Aznar’s report, and similarities in the taxonomic structure and taphonomic features connect the “Museum Collection” to the documented contexts. The discovery of a Neanderthal upper M3 that is compatible with “Museum Collection” individual 2 supports this interpretation, which is further corroborated by Mr. Aznar identifying the excavation area.
The 14C chronology provided by charcoals from our excavations are compatible with those from the charcoals included in the sediment samples associated to the human remains from the “Museum Collection,” potentially placing Simanya Neanderthals beyond 49,000 years BP. A minimum age of
The Simanya sample is attributed to Neanderthals due to distinct apomorphic traits (see
Simanya Neanderthals’ apomorphic traits align with other late Neanderthals, supporting, together with the chronological data from the excavation, its location in a final evolutionary phase of the evolution of western Neanderthals. However, different genetic backgrounds within Iberian Neanderthal populations cannot be ruled out. The substitution of Neanderthal populations in Western Europe (
The analysis of the Cova Simanya assemblage has allowed the identification of a new Neanderthal sample that will provide essential data on the morphological traits and evolutionary context of late Neanderthals in the Iberian Mediterranean region. Further comparative studies, incorporating morphometric and genetic analyses, with other Iberian samples will elucidate their phylogeography and population dynamics to shed light on a complex evolutionary history comprising different genetic backgrounds and relict genetic groups of late Iberian Neanderthals.
The original contributions presented in the study are included in the article/
Conceptualization: JM and AR. Investigation: JM, AC, MS, AR-H, RH, EM-R, DL, JR, DM-P, AG-T, EA, AG-B, EL, TM-B, ST, CL-F, and AR. Writing—original draft: JM and AR. Writing—review and editing: JM, AC, MS, AR-H, RH, EM-R, DL, JR, DM-P, AG-T, EA, AG-B, EL, TM-B, ST, CL-F, JF, and AR. Project administration: JM, JF, and AR. Funding acquisition: JM, JF, AR, MS, ST, and CL-F. All authors contributed to the article and approved the submitted version.
This research was funded by the Spanish Ministry of Science and Innovation through the projects PID2021-122356NB-I00 (MNCN-CSIC, AR), PID2020-113960GB-I00 (UB, JF), PID2021-124590NB-I00 (MCNB, CL-F), PID2019-103987GB-C31 (IPHES-CERCA), and PID2021-126004NB-100 (IBE-UPF-CSIC, TM-B); the AGAUR through the research groups 2021 SGR 01237 (IPHES-CERCA, EA), 2021 SGR 00337 (UB, JF), and 2021 SGR 00177 (IBE-UPF-CSIC, TM-B); the Culture Department of the Generalitat de Catalunya through the project ARQ001SOL-172-2022; and the Diputació de Barcelona and the Fundación Palarq. MS was funded by the UAM Tomás y Valiente Program, JR by the European Union-Next Generation EU, Ministry of Universities and UA (MARSALAS21-22), and DL by the Xunta de Galicia Grant ED481B-2022-048. ST and TM-B received funding from the European Research Council under the European Union’s Horizon 2020 Research and Innovation Program (grant agreement no. 803147 RESOLUTION,
First and foremost, the authors would like to express their deepest gratitude to the Museu d'Arqueologia de Catalunya for facilitating their access to the essential Aznar collection. The authors would also like to express their gratitude to the Parc Natural de Sant Llorenç del Munt i l'Obac for its supportive role in facilitating the recommencement of excavations at Cova Simanya and for its commitment to the preservation of the site. They also extend their acknowledgement to Mr. Angel Miño and Mr. Josep Canals, whose assistance has been essential to their operations. Furthermore, they would like to acknowledge the consistent support provided by Mr. Joan Lleonart and Mr. Josep Fígols throughout the fieldwork. Their contributions have been beneficial to the progression and outcomes of the research.
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
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