TYPE Brief Research Report PUBLISHED 19 September 2023 DOI 10.3389/feart.2023.1230707 A new assemblage of late OPEN ACCESS Neanderthal remains from Cova EDITED BY Raymond Louis Bernor, Howard University, United States Simanya (NE Iberia) REVIEWED BY Leslea J. Hlusko, Juan I. Morales 1,2,3*, Artur Cebrià 3, María Soto 4,5, National Center for Research on Human Evolution (CENIEH), Spain Antonio Rodríguez-Hidalgo 1,2,6, Raquel Hernando 1,2, John De Vos, Elena Moreno-Ribas 1,2, Diego Lombao 1,7, José R. Rabuñal 8,9, Naturalis Biodiversity Center, Netherlands Marina Melchionna, David M. Martín-Perea 1,6,10, Antonio García-Tabernero 11, University of Naples Federico II, Italy Ethel Allué 1,2, Andrea García-Basanta 5, Esther Lizano 12, *CORRESPONDENCE Juan I. Morales, Tomàs Marquès-Bonet 12,13,14,15, Sahra Talamo 16, Laura Tassoni 16, jmorales@iphes.cat Carles Lalueza-Fox 13,17, Josep M. Fullola 3 and Antonio Rosas 11* Antonio Rosas, 1 arosas@mncn.csic.es IPHES-CERCA, Institut Català de Paleoecologia Humana i Evolució Social, Tarragona, Spain, 2 Departament d’Història i Història de l’Art, Universitat Rovira i Virgili, Tarragona, Spain, 3SERP, RECEIVED 29 May 2023 Departament d’Història i Arqueologia, Secció de Prehistoria, Universitat de Barcelona, Barcelona, Spain, ACCEPTED 04 July 2023 4 Madrid Institute for Advanced Study, MIAS, Madrid, Spain, 5Departamento de Prehistoria y Arqueología, PUBLISHED 19 September 2023 Facultad de Filosofía y Letras, Universidad Autónoma de Madrid, Madrid, Spain, 6Instituto de Evolución en CITATION África (IDEA), Madrid, Spain, 7GEPN-AAT, Departamento de Historia, Facultade de Xeografía e Historia, Morales JI, Cebrià A, Soto M, Universidade Santiago de Compostela, Santiago de Compostela, Spain, 8Instituto Universitario de Rodríguez-Hidalgo A, Hernando R, Investigación en Arqueología y Patrimonio Histórico, Universidad de Alicante, Alicante, Spain, 9 Moreno-Ribas E, Lombao D, Rabuñal JR, Department of Archaeology and Heritage Studies, Aarhus University, Højbjerg, Denmark, 10Department Martín-Perea DM, García-Tabernero A, of Geodynamics, Stratigraphy and Palaeontology, Complutense University of Madrid, Madrid, Spain, 11 Allué E, García-Basanta A, Lizano E, Group of Paleoanthropology, Museo Nacional de Ciencias Naturales (CSIC), Madrid, Spain, 12Institut Marquès-Bonet T, Talamo S, Tassoni L, Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Barcelona, Spain, 13 Lalueza-Fox C, Fullola JM and Rosas A Institute of Evolutionary Biology (UPF-CSIC), University Pompeu Fabra, Barcelona, Spain, 14Catalan (2023), A new assemblage of late Institution of Research and Advanced Studies (ICREA), Barcelona, Spain, 15CNAG-CRG, Centre for Neanderthal remains from Cova Simanya Genomic Regulation (CRG), Barcelona Institute of Science and Technology (BIST), Barcelona, Spain, 16 (NE Iberia). Department of Chemistry Giacomo Ciamician, University of Bologna, Bologna, Italy, 17Museum of Front. Earth Sci. 11:1230707. Natural Sciences of Barcelona, Barcelona, Spain doi: 10.3389/feart.2023.1230707 COPYRIGHT © 2023 Morales, Cebrià, Soto, Rodríguez- Hidalgo, Hernando, Moreno-Ribas, This study presents an exceptional collection of 54 Late Pleistocene human Lombao, Rabuñal, Martín-Perea, García- Tabernero, Allué, García-Basanta, Lizano, remains that correspond to at least three Neanderthal individuals from Simanya Marquès-Bonet, Talamo, Tassoni, Gran, the main gallery of Cova Simanya, located in the northeastern Iberian Lalueza-Fox, Fullola and Rosas. This is an Peninsula. The collection comprised 53 unpublished remains that were open-access article distributed under the terms of the Creative Commons unearthed during the 1970s and an additional tooth discovered during Attribution License (CC BY). The use, 2021 excavations. The specimens represent an adult with a small stature, a distribution or reproduction in other periadolescent aged approximately 11.5 years, and an immature individual aged forums is permitted, provided the original author(s) and the copyright owner(s) are approximately 7.7 years, thus offering a more complete demographic perspective. credited and that the original publication The collection encompasses diverse anatomical parts including upper and lower in this journal is cited, in accordance with dentition, mandible, vertebrae, and limb bones from both the upper and lower accepted academic practice. No use, distribution or reproduction is permitted extremities. Attempts to extract aDNA were unsuccessful. Renewed which does not comply with these terms. 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. Frontiers in Earth Science 01 frontiersin.org Morales et al. 10.3389/feart.2023.1230707 KEYWORDS Late Pleistocene, Middle Paleolithic archeology, human remains, Neanderthal, Cova Simanya, paleoanthropology, Paleolithic archaeology, Iberian Peninsula 1 Introduction Aparicio et al., 2014; Garralda et al., 2014; Quam et al., 2015; Bokelmann et al., 2019; Carrión et al., 2019) (Figure 1A). Neanderthals exhibited distinct genetic histories and cultural These data suggest that Mediterranean Iberia was continuously adaptations along time and space. In the Iberian Peninsula, the inhabited by Neanderthals from MIS-7 onwards, with northeastern Iberian Mediterranean region holds significant importance in Iberia being an especially dense Mousterian cluster (Martínez- understanding Neanderthal evolution and variability, and its Moreno et al., 2010a; Martínez-Moreno et al., 2010b; Rosell et al., archaeological record provides valuable insights into their 2010; Mora et al., 2018; Morales et al., 2022) (Figure 1B), including biological and cultural development as well as their eventual long-term occupation archives such as Abric Romaní (Barcelona) disappearance during marine isotope stage (MIS) 3. (Bischoff et al., 1988; Vallverdú et al., 2010; Carbonell, 2012) or Middle Paleolithic Mousterian sites can be found along the L’Arbreda (Soler et al., 2014), and the southernmost occurrence of Mediterranean coast from Gibraltar to the Pyrenees, with Châtelperronian technologies (Morales et al., 2019; Rodríguez- Neanderthal remains being documented at several locations Hidalgo et al., 2019). However, the paleoanthropological record (Dean et al., 1986; Arsuaga and Bermúdez de Castro, 1987; in this region is restricted to the isolated teeth from Mollet (~215 ka) Daura et al., 2005; Arsuaga et al., 2007; Walker et al., 2008; (Maroto et al., 2012), and the Cova del Gegant assemblage dated Daura et al., 2010; Walker et al., 2010; Rodríguez et al., 2011; back to 55,000–52,000 years ago (Daura et al., 2005; Daura et al., Arsuaga et al., 2012; Maroto et al., 2012; Michel et al., 2013; 2010; Quam et al., 2015; Daura et al., 2022) once the long-time FIGURE 1 (A) Map with the distribution of Neanderthal remains in the Iberian Peninsula; (B) main Middle Paleolithic sites from NE Iberia and the situation of Cova Simanya; (C) Cova Simanya karstic system, in red the areas excavated between 2020–2022, and (D) main stratigraphic sections for the Pleistocene sequence. Frontiers in Earth Science 02 frontiersin.org Morales et al. 10.3389/feart.2023.1230707 disputed Banyoles mandible does not show any derived Neanderthal their conservation status. Following the evaluation, a series of features (Keeling et al., 2023). measures were taken to inhibit further deterioration. Cleaning Herein, we present a novel Neanderthal assemblage from Cova procedures involving water and soft brushes were applied, and Simanya (Sant Llorenç Savall, Barcelona), first recognized during the outdated adhesives were removed and substituted with Paraloid® analysis of a museum collection and constituting the largest sample B-72, a durable and reversible acrylic resin. This resin was also documented so far in northeastern Iberia. This discovery has led to a utilized to mend modern fractures and consolidate the overall new research project that aimed at reconstructing the genesis, structure, ensuring the assemblage’s longevity and integrity for context, and paleobiology of the Neanderthal sample, including future study and preservation. renewed excavations at Cova Simanya. These works have provided a robust archeological context for the uncontextualized “Museum Collection” including additional Neanderthal remains. 2.3 2020–2022 excavations Simanya’s assemblage may offer insights into the late Neanderthals’ apomorphic traits and their congruence with other The 2020–2022 excavation programs targeted Simanya Gran and known samples, while also raising questions about the genetic aimed at identifying the original context of the “Museum Collection” diversity and population turnovers within the Iberian Peninsula and evaluating the potential of the Pleistocene deposits in the cave. In (Vernot et al., 2021). 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 2 Materials and methods IMG sector (Figure 1D). The latter provided a similar fossil assemblage to the “Museum Collection,” including a new Neanderthal tooth (SI-60) 2.1 Cova Simanya found in the area reported by Mr. Aznar as the origin of the collection (Supplementary Table S1). In 2022, we continued with the excavation of Cova Simanya is a karstic system (UTM ETRS89 the IMG and expanded the S3 test pit of the MG to a surface of 10 m2. A 417517–4614066, 885 masl) opened in Montcau mountain, visual recognition of Mr. Aznar excavations (trenches, sections, or running through the Paleogene conglomerates forming the Sant backfills) was not possible due to the homogenization of the Llorenç del Munt Massif, on the eastern margin of the Ebro Basin, sedimentary surface by the unrestricted public access. 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 (Figure 1C 2.4 Radiocarbon dating and Supplementary Figure S1). Simanya Gran is the larger sector of Cova Simanya, comprising Eleven bone samples, both anthropized and non-anthropized the Western Gallery (WG), Main Gallery (MG), Pas de l’Oca Gallery remains, and five charcoals (Pinus sylvestris type) from the (POG), and the inner part of the Main Gallery (IMG) (Figure 1C). “Museum Collection” and our own excavations at the IMG were The cave has been known to scholars at least since the 18th century, selected for radiocarbon pretreatment at the University of Bologna’s and the presence of archeological remains is known since the first BRAVHO facilities. Talamo et al. (2021) followed the collagen excavation in the 1930s. extraction method, and only a non-anthropized Capra pyrenaica humerus (SG21.06.39) from the excavation yielded sufficient collagen for 14C dating. Charcoal pretreatment used the ABOx 2.2 The “Museum Collection” method proposed by Tassoni et al. (2022), adapting Bird et al.’s (1999) method to small, poorly preserved samples by adjusting the The “Museum Collection” was collected by an amateur, Mr. HCl solution’s molarity and reducing the oxidation step’s reaction Miguel Aznar, in 1978–79 and contains a combination of two time. After the pretreatment, the collagen and charcoal samples were distinct sources: 1) surface findings and 2) remains from a small graphitized at the BRAVHO lab (Tassoni et al., 2022). During the excavation. The surface findings include isolated human and animal charcoal sample SG-CA-200 combustion, the C integral peak bones, pottery, and chert fragments. The excavation yielded human obtained was very low for analysis and was excluded. The and faunal remains, as well as a few samples of charcoal, sediment, resulting graphite was pressed into targets and sent to the and carbonate crusts. The excavation was performed in a breccia Mannheim AMS Laboratory at the Curt-Engelhorn-Center for deposit of reddish sediments with abundant charcoals located in the Archeometry for measurement. IMG sector. No pottery or any other Holocene remains were found All 14C ages were calibrated with the IntCal20 calibration curve during the excavation. The collection containing 143 osteological (Reimer et al., 2020) using OxCal v4.4 software (Bronk Ramsey, remains was donated to the Museum of Archaeology of Catalonia in 2009). 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. 2.5 aDNA Despite being layered in the reddish clay-like sediment, the human bone collection exhibited a notable state of preservation. An attempt to extract aDNA from Simanya Neanderthals was However, the presence of modern fractures and adhesive-induced made by sampling a tooth and a phalanx from the “Museum alterations from the 1970s necessitated a thorough evaluation of Collection” through shotgun sequencing (Illumina NovaSeq Frontiers in Earth Science 03 frontiersin.org Morales et al. 10.3389/feart.2023.1230707 6000) and mitochondrial DNA (mtDNA) capture with Within this horizon, a small and flat combustion feature was also Neanderthal-specific baits (Arbor Biosciences). documented. The presence of core-trimming flakes, a sidescraper, multifaceted butts, dorso-ventral right angles, and centripetal dorsal scar patterns in the lithic assemblage clearly indicates a Mousterian 3 Results affinity (Morales et al., 2022). Additionally, a third upper human molar (SI-60) was found in Layer O300-PO400 (301–80 meter). 3.1 Stratigraphic context and archeo- Finally, Layer P300, documented exclusively at S3, has provided a paleontological record faunal assemblage composed of highly cemented skeletal remains of Capra pyrenaica and isolated remains of Cervus elaphus, without The litho-stratigraphic sequence documented so far for the MG any evidence of human activity so far. 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 3.2 Dating the MG to the end of the IMG, the Pleistocene units are uncapped (Supplementary Figure S2.1) and four main litho-stratigraphic units Calibrated dates at 68.3% and 95.4% probability are provided in have been described (Figure 1D), namely, from the bottom to the Table 1. The bone sample is the only one that results in a final top, Unit SG4 (cave clays), Unit SG3 (channel deposits), Unit SG2 calibrated range of 42,440–42,110 cal BP (68.3%). All the other four (complex unit), and Unit SG1 (diamicton). charcoal samples are older than the radiocarbon method (>49,000). 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 3.3 Paleoanthropology SG3. Unit SG3 comprised several coarsening upward sequences (silt/ clay to coarse sand) that are cut and filled by at least four poorly A collection of 53 human remains was recovered by Mr. Aznar sorted (silts to coarse sand) channel deposits. The thickness of Unit in 1978–79, to which an upper third molar (SI-60) found in situ SG3 varies from 5 to 45 cm, and it has been documented in S1, S2, during the excavations (Supplementary Table S2) must be added. S3, and the IMG. In the S1 section of the MG sector, Unit SG3 is The assemblage includes upper and lower dentition, a fragment of conformably overlain by Unit SG2-a, an approximately 50 cm-thick the mandible, vertebrae, the left and right upper limb (humeri and matrix-supported conglomerate of slate and limestone clasts that are hand bones), and foot bones. Some of the specimens are preserved in more abundant toward the top, while in the S3 section, it is overlain complete and good condition, while others are fragmentary with old by Units SG2.3-to-1, a succession of brown clays, a poorly sorted dry (diagenetic) and modern fractures (Figure 2) (probably broken yellowish sandy deposit, and a very fine gravel conglomerate, and by during the excavation). No evidence of pathological processes has the S1 flowstone. The Pleistocene sequence ends with Unit SG1, a been detected in the sample, while the preserved dentition does not reddish-brown, matrix-supported polygenic limestone, sandstone, display strong evidence of the use of the mouth in heavy-duty tasks. and slate conglomerate with an erosive base that has been At least three individuals can be identified in the sample. The documented exclusively at the IMG. most complete one is defined by a complete left humerus SI-1 Three archaeological layers have been documented so far within (Figure 2A). Several other postcranial remains (right proximal the litho-stratigraphic sequence at the MG–IMG sector. Layer humerus, hand 1 and hand 2, vertebra; Figures 2B–D) can be P1 corresponds to the Holocene phase and yields artifacts from securely associated based on the size and taphonomic features. It the Late Neolithic to the Late Antiquity, including scattered funerary corresponds to an adult, with a stature estimation for the humerus of evidence dated back to the 5th century AD, in a condensed sub- 154 ± 4.89 cm based on the Sjøvold (1990) equation (max. length: superficial sedimentary palimpsest (Supplementary Figure S2.1). 293 mm), which is consistent with a Neanderthal female individual. The Pleistocene layer O300-PO400 (Unit SG1) was documented SI-7 is an upper left P3 at the Rc growth stage [according to the in the IMG and is characterized by a paleontological background work of Moorrees et al. (1963)], with the root length completed with composed mostly of Ursus cf. arctos, U. cf. spelaeus, and Capra parallel ends (Figure 2E). This provides an estimated age of pyrenaica. The preliminary analysis of the assemblage suggests a approximately 11.5 years by modern human standards [according combination of taphonomic agents and processes in the formation to the work of AlQahtani et al. (2010)]. SI-60 is a complete upper left of the record, including bear deaths during hibernation, M3 crown (Figure 2E) with a defined pulp roof [CRc, according to accumulation of Iberian ibex by a large carnivore (likely a felid), the work of Moorrees et al. (1963)], which would provide an age of and transportation and/or reworking by physical processes such as 12.5–13.5 years [according to the work of AlQahtani et al. (2010)]. water flow and gravity. This taxonomic profile is consistent with the However, M3 development is thought to be accelerated in “Museum Collection,” which mostly consists of U. cf. arctos and Neanderthals as compared to modern humans (Guatelli- Capra pyrenaica. Evidence of occasional Middle Paleolithic Steinberg, 2009), which makes it probable that the M3 (SI-60) Mousterian occupation has also been found to be interspersed in and P3 (SI-7) may come from a single individual of the paleontological background of O300-PO400, over 70 m inside approximately 11.5 years old. the cave (Supplementary Figures S2.2, 2.3, S3). The evidence Two other immature remains are also preserved in the sample: associated to human occupations contains a small lithic SI-29 is a hand first phalanx of ray V, whose size is like Juvenile assemblage and some faunal remains (Sus scrofa and Cervus 1 from El Sidrón (Rosas et al., 2017) (Figure 2F). This provides an elaphus) that exhibit cut marks or green breakage patterns. approximate age of 7.61–7.78 years for this individual. SI-37 is a Frontiers in Earth Science 04 frontiersin.org Frontiers in Earth Science Morales et al. 14 TABLE 1 Samples selected for C dating pretreated at the University of Bologna and AMS results. Bologna Sample Unit Sector Material Species Collagen Collagen AMS d13C F14C C:N 14C 1s Cal BP Cal BP Code ID (mg) (%) Code age Error age (68.3%) age (95.4%) BRA-6108 SG21.06.39 O300-PO400 301-70 Bone Caprini 4.8 1% MAMS −18.21 0.0088 2.89 38009 410 42440–42110 42620–41930 56486 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 IMG Tooth Ursidae 0.8 0% — Collection BRA-6115 SG-CA-125 Museum IMG Tooth Ursidae 0.5 0% — Collection BRA-6116 SG-CA-10 Museum IMG Tooth Ursidae 0.5 0% — 05 Collection BRA-6117 SG-CA-62 Museum IMG Bone Indeterminate 0 0% — Collection BRA-6118 SG-CA-137 Museum IMG Bone Caprini 1.2 0% — Collection Carbon (mg) Carbon (%) BRA-6106 SG21.11.53 O300-PO400 301-72 Charcoal Pinus sylvestris 33.3 35% MAMS −22.61 0.0009 137.59 >49000 — — — type 56487 BRA-6107 SG21.03.05 O300-PO400 301-86 Charcoal Pinus sylvestris 50.8 49% MAMS −23.43 0.0012 129.79 >49000 — — — type 56488 BRA-6264 SG-CA-200 Museum IMG Charcoal Pinus sylvestris 4.6 7% — Collection type 10.3389/feart.2023.1230707 BRA-6265 SG-CA-210 Museum IMG Charcoal Pinus sylvestris 11.7 21% MAMS −24.17 0.0020 87.05 >49000 — — — Collection type 62623 BRA-6266 SG-CA-207 Museum IMG Charcoal Pinus sylvestris 6.2 16% MAMS −23.79 0.0013 172.17 >49000 — — — Collection type 62624 frontiersin.org Morales et al. 10.3389/feart.2023.1230707 FIGURE 2 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. (A) Left humerus; (B) shaft, caudal view; (C) bones of the foot; (D) bones of the right hand; (E) upper P3 and M3; (F) fragment of the atlas and first phalanx of the hand; and (G) fragment of the ascending branch. fragment of an atlas from the right side, preserving upper and lower morphometric techniques based on landmark configurations, articular facets. The surface of the neurocentral junction can be according to Rosas et al.’s (2015) procedures and comparative appreciated, meaning that the anterior arch of the atlas was still samples (see Supplementary Material). unfused even though the size of the specimen is relatively large. This Taxonomically, the human remains from Simanya Gran present is fully compatible with the growth pattern established for a large series of Neanderthal apomorphic features distributed across Neanderthals by Rosas et al. (2017) and congruent with the age different anatomical systems. Upper central incisors are large with a of 7.7 years established by the SI-37 phalanx. robust root, a strong dental tubercle, and characteristic flat wear The size of lower dentition I1 (SI-3) and M3 is relatively small, (Supplementary Figure S4). The upper lateral incisors present a while P3 (SI-5) falls at the upper limit and the upper I1 (SI-45) in conspicuous and characteristic shovel shape, where the marginal the middle of the Neanderthal distribution, while the mesiodistal ridges are extremely thickened and invade the lingual fossa, crown diameter of the I 2 (SI-4 and SI-31) falls well above the circumscribing a narrow longitudinal groove. The mandible Neanderthal mean [according to the work of Pinilla and Trinkaus shows a large, moderately truncated gonial area, with a large (2017)] (Supplementary Table S3, Supplementary Figure S4). medial pterygoid tubercle, and a lateral position of the condyle in Historical and taphonomic conditions of the assemblage make relation to the mandibular notch (Figure 2G). Humerus SI-1 it feasible to assign some of these teeth to individual 1. However, (Figures 2A, 3; Supplementary Material) displays a massive and because of the noted size differences, the presence of other anteriorly projecting lesser tubercle. A deep olecranon fossa, which individuals cannot be ruled out. For the time being, no is elliptical in shape, wider than high, and a marked medial individual has been assigned to adult dentition. The upper and epicondyle are moderately prominent. The long lateral lower permanent teeth exhibit moderate occlusal wear [stage 5 of supracondylar crest and coracobrachial insertion are located the work of Skinner (1997)], indicating a mature adult. more distally than the distal limit of the deltoid tuberosity, while Considering that Neanderthal anterior permanent teeth wear the deltoid tuberosity displays two parallel and proximal ridges. The faster or commence wear earlier (Skinner, 1997), an age of humeral torsion is moderate. The first metacarpal is short and 30–40 years may be considered. displays a flange of bone along the distodorsoradial margin, In order to assess the location of the Simanya Gran specimen SI- marking the insertion of the opponens pollicis muscle. The first 1 in the morphospace (Figure 3), we used 3D geometric pollex phalanx is relatively short. The first phalanx of the second ray Frontiers in Earth Science 06 frontiersin.org Morales et al. 10.3389/feart.2023.1230707 FIGURE 3 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 Rosas et al. (2015). Landmark 8 of the Simanya humerus has been estimated by the thin plate spline method. Ellipses for the prediction interval of a single new observation (0.95% probability) are shown only for the Neanderthal and modern human samples. Note the position of the Simanya’s humerus clearly within the Neanderthal distribution. presents a robust and prominent radial tubercle, and the distal consistent with the open character of the layer that could have been phalanges show an extremely wide distal end (Figure 2D). The big deposited during a relatively long period including different time- toe metatarsal and distal phalanx are gracile, but the distal head of segregated events. the metatarsal is proportionally bulky (Figure 2C). The Simanya sample is attributed to Neanderthals due to Ultimately, no positive results for aDNA were obtained, neither distinct apomorphic traits (see Figure 3). It includes at least three for shotgun sequencing nor for the mtDNA capture. individuals—an adult, a periadolescent (~11.5 years), and an immature (~7.5 years)—with preserved dentition, mandible, vertebrae, upper limbs, and foot bones. Some specimens are 4 Discussion and conclusion complete and well preserved, while others are fragmentary with diagenetic and modern fractures. Taphonomic conditions allow Excavations at Simanya Gran revealed Pleistocene fossil-bearing assigning some teeth to individual 1, but based on size deposits, primarily preserved in the layer O300-PO400 from the differences, a fourth individual is possible. IMG and likely dismantled from the cave entrance. Its sub- Simanya Neanderthals’ apomorphic traits align with other late superficial position aligns with Mr. Aznar’s report, and Neanderthals, supporting, together with the chronological data from similarities in the taxonomic structure and taphonomic features the excavation, its location in a final evolutionary phase of the connect the “Museum Collection” to the documented contexts. The evolution of western Neanderthals. However, different genetic discovery of a Neanderthal upper M3 that is compatible with backgrounds within Iberian Neanderthal populations cannot be “Museum Collection” individual 2 supports this interpretation, ruled out. The substitution of Neanderthal populations in which is further corroborated by Mr. Aznar identifying the Western Europe (Dalén et al., 2012) and a population turnover excavation area. with mtDNA diversity loss (~100 ka ago) have been identified The 14C chronology provided by charcoals from our excavations (Vernot et al., 2021). The presence of primitive features in other are compatible with those from the charcoals included in the samples, such as the 49,000 old from El Sidrón (Rosas et al., 2020), sediment samples associated to the human remains from the allows for hypothesizing the potential persistence of relict groups. “Museum Collection,” potentially placing Simanya Neanderthals Comparative studies with other Iberian samples may clarify late beyond 49,000 years BP. A minimum age of circa 42,000 cal BP has Iberian Neanderthals’ phylogeography. been obtained for the paleontological assemblage that is also present The analysis of the Cova Simanya assemblage has allowed the in the layer O300-PO400. The discrepancy between both ages is identification of a new Neanderthal sample that will provide Frontiers in Earth Science 07 frontiersin.org Morales et al. 10.3389/feart.2023.1230707 essential data on the morphological traits and evolutionary context agreement no. 803147 RESOLUTION, https://site.unibo.it/ of late Neanderthals in the Iberian Mediterranean region. Further resolution-erc/en (ST), and no. 864203 (TM-B)]. IPHES-CERCA comparative studies, incorporating morphometric and genetic received financial support from the Spanish Ministry of Science and analyses, with other Iberian samples will elucidate their Innovation through the “María de Maeztu” program for Units of phylogeography and population dynamics to shed light on a Excellence (CEX 2019-000945-M). complex evolutionary history comprising different genetic backgrounds and relict genetic groups of late Iberian Neanderthals. Acknowledgments Data availability statement First and foremost, the authors would like to express their deepest gratitude to the Museu d’Arqueologia de Catalunya for The original contributions presented in the study are included in facilitating their access to the essential Aznar collection. The authors the article/Supplementary Material; further inquiries can be directed would also like to express their gratitude to the Parc Natural de Sant to the corresponding authors. 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 Author contributions acknowledgement to Mr. Angel Miño and Mr. Josep Canals, whose assistance has been essential to their operations. Furthermore, they Conceptualization: JM and AR. Investigation: JM, AC, MS, AR-H, would like to acknowledge the consistent support provided by Mr. RH, EM-R, DL, JR, DM-P, AG-T, EA, AG-B, EL, TM-B, ST, CL-F, Joan Lleonart and Mr. Josep Fígols throughout the fieldwork. Their and AR. Writing—original draft: JM and AR. Writing—review and contributions have been beneficial to the progression and outcomes editing: JM, AC, MS, AR-H, RH, EM-R, DL, JR, DM-P, AG-T, EA, of the research. 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. Conflict of interest The authors declare that the research was conducted in the Funding absence of any commercial or financial relationships that could be construed as a potential conflict of interest. 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- Publisher’s note 124590NB-I00 (MCNB, CL-F), PID2019-103987GB-C31 (IPHES- CERCA), and PID2021-126004NB-100 (IBE-UPF-CSIC, TM-B); All claims expressed in this article are solely those of the authors the AGAUR through the research groups 2021 SGR 01237 and do not necessarily represent those of their affiliated organizations, (IPHES-CERCA, EA), 2021 SGR 00337 (UB, JF), and 2021 SGR or those of the publisher, the editors, and the reviewers. Any product 00177 (IBE-UPF-CSIC, TM-B); the Culture Department of the that may be evaluated in this article, or claim that may be made by its Generalitat de Catalunya through the project ARQ001SOL-172- manufacturer, is not guaranteed or endorsed by the publisher. 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 Supplementary material UA (MARSALAS21-22), and DL by the Xunta de Galicia Grant ED481B-2022-048. ST and TM-B received funding from the The Supplementary Material for this article can be found online European Research Council under the European Union’s at: https://www.frontiersin.org/articles/10.3389/feart.2023.1230707/ Horizon 2020 Research and Innovation Program (grant full#supplementary-material References AlQahtani, S. J., Hector, M. P., and Liversidge, H. M. (2010). Brief communication: Arsuaga, J. 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