Reconstructing Contact and Potential Interbreeding Geographical Zone for Neanderthals and Anatomically Modern Humans

Reconstructing Contact and Potential Interbreeding Geographical Zone for Neanderthals and Anatomically Modern Humans | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Reconstructing Contact and Potential Interbreeding Geographical Zone for Neanderthals and Anatomically Modern Humans Saman Heydari-Guran, Masood Yosefi, Anoshe Kafash, Elham Ghasidian This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4298125/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 03 Sep, 2024 Read the published version in Scientific Reports → Version 1 posted 10 You are reading this latest preprint version Abstract Whilst the admixture of the Neanderthal and anatomically modern human is precisely proven, due to the shortage of fossils and absence of appropriate DNA, the timing and geography of it, are the subjects of questions. In this study, we applied ecological niche modelling (maximum entropy approach) and GIS to reconstruct Neanderthal and modern human palaeodistribution and identify their contact and potential interbreeding zone during the marine isotope stage 5. We used climatic variables characterising environmental conditions of marine isotope stage 5 ca. 120 to 80 thousand years ago (Kyr) along with topography and coordinates of Neanderthal and modern human archaeological sites to build each species palaeodistribution. Overlapping the models showed that Zagros Mountains was a contact and potential interbreeding zone for the two sub-human species. We believe that Zagros Mountains acted as a corridor connected the Palearctic/Afro-Arabian realms, facilitated northward dispersal of anatomically modern human and southward dispersal of Neanderthal during marine isotope stage 5. Our analyses are comparable with archaeological and genetic evidence collected during the last decades. Zagros Mountains Ecological niche palaeoenvironment Neanderthals anatomically modern humans Figures Figure 1 Figure 2 Introduction Following the ground-breaking discovery of biocultural admixture of Late Pleistocene different early human groups of Neanderthals, archaic/modern human, and Denisovans, a large and growing body of research concerning the nature and evolutionary history of these events are presented. Besides of significant consequences the biological exchanges that have had to subspecies (e.g. Quach et al 2016) and related issues, the time (Green et al 2010; Prüfer et al 2014 and 2017) and geography of contact and interbreeding are subject of intense debate (Stewart and Stringer 2012). Neanderthals are an extinct lineage of hominins that emerged around 400 Kyr and died off around 40 Kyr (Higham et al 2014). Fossil localities and morphological evidence of Neanderthals indicate that they were companionable with the Palearctic biogeographical realm, which includes from western Europe to Altai Mountains in Siberia at 55° latitude and down until around 31° 1atitude in the Western Asia (Kolobova et al. 2020, Heydari-Guran et al. 2021, Been et al. 2017). The chronological settlement patterns of the Neanderthals' sites indicate of their expansion to the east and south west Asia at least from 150 Kyr (Grün and Stringer 2000). At the other side, anatomically modern humans (AMH) have evolved in Africa more than 300 Kyr (Hublin et al 2017; Richter et al 2017). The evidence including physical remains and morphological analyses suggest that they have exited Africa over and over during a period of at least 200kyr (Groucutt et al 2018; Hershkovitz et al 2018; Harvati et al 2019). AMH, also reached Eastern Asia around 120kyr (Liu et al 2010) and later on into Europe around 60 Kyr (Slimak et al 2022; Hublin et al 2020). Recent accurate archaeological and Paleoenvironmental data suggest that AMH rapidly adapted to the new and extreme environments beyond Africa, such as high-plateaus, mountain systems and palearctic ecosystem (Roberts and Stewart 2018). Meanwhile, the archaeological and fossil evidence indicate that AMH have entered mainly to south western Asia during MIS 5 (Groucutt et al 2012; Scerri et al 2015; Groucutt et al 2018, Heydari-Guran and Ghasidian 2021). There is strong evidence of multiple interbreeding between two groups of Neanderthal and archaic/modern human in the western Eurasia (e.g. Prüfer et al 2014; Slon et al 2018). Meanwhile, many attempts have been made to estimate the time for this admixture and significant successes have been achieved (e.g. Krause et al 2010; Yuan et al 2021). The Paleogenetic studies support interbreeding between AMH and Neanderthals go back to at least 130 Kyr (Prüfer et al 2017). In some studies, researchers suggest that the lower latitude regions of southwestern Asia have a high potential for the first overlap between Neanderthal and AMH. Sanchez Goñi (2020) uses the patterns of expansion of Neanderthal and AMH and share same ecological niches under the certain climatic conditions during the late Pleistocene. Recently Churchill et al. (2022) found facial morphological similarities between Neanderthals and AMH in the Near East, indicating key region for the interbreeding between the two lineages. But still it is unclear where the two species met and probably interbreed. Species Distribution Models (SDMs) are very practical tools to investigate geography of the two species palaeodistribution and potential interbreeding areas (Ruan et al 2023). SDMs which have found important application in paleobiogeography, archaeological and paleo-anthropological (Varela et al., 2011; Svenning et al., 2011 26 ; Franklin et al., 2015; Giampoudakis et al., 2017; Benito et al., 2017). They use occurrence data of a target species including ancient humans and paleoenvironmental variables to calculate probability of a species or hominin species’ presence in a defined geographic region (Guisan et al. 2017) . These models have successfully been used to reconstruct different hominin species (Giampoudakis et al 2017; Benito et al 2017; Jones et al 2019), identify refugia during the ice ages and reconstructing dispersal corridors (Kafash et al 2023), their niche overlap with each other (Ruan et al 2023) and niche overlap with their prey species (Yousefi et al 2020 33 ). For instance, Ruan et al. (2023) have used SDMs to identify contact zone of Neanderthal and Denisova. In another study, Benito et al. (2017) applied the ecological niche and distribution of the Neanderthal during the Last Interglacial in Europe and the Irano-Turonian region. Thus, SDMs are used to model Neanderthal and AMHs palaeodistribution and locate geography of their niche overlap (Benito et al. 2017; Ruan et al. 2023). The aims of present study are to first reconstruct palaeodistribution of Neanderthal and AMHs during the MIS 5, identify the most important predictor of the two species, investigate the two species response to environmental variables and identify geographic overlap for the two species at the meeting zone of three continents, Asia, Africa and Europe. Considering previous studies on the importance of Southwest Asia the African continent and southern part of the Middle East are located in the Afro-Arabian biogeographical realms () which matched with distribution of AMH and the lands above Levant. The northern part of the Persian and Anatolian plateaus, Caucasus to European continent and Siberia are setting in the Palearctic realm were considered to be the distribution range of the Neanderthals. Here, we hypothesized that these two species first met and interbreed at the border of the two biogeographic realms where environmental condition facilitated niche overlap and resource partitioning by providing highly diverse habitat rich in resources. In this regard climate is a major determinant of species distributions (Sexton et al 2009) particular at large spatial scales, we are expecting climate to be more effective than topography in shaping the two sub species of Neanderthal and AMH. Results Reconstructing contact and interbreeding zone. The models developed in this study for Neanderthal (AUC=941) and AMH (AUC= 895) performed well according to the AUC model performance metric. Our model of paleodistribution of Neanderthal shows that north and west of Mediterranean Sea toward Levant, vast patches in Turkey, around the Black Sea, south of Caspian Sea, Taurus, Caucasus and Zagros Mountains have high suitability for the species during the MIS 5 (Figure 1). AMH paleo-distribution model identified large and continues suitable patches and in Africa, Arabia and Iranian Plateau. Our model identified Zagros Mountains as a contact and potential interbreeding zone Variable importance and response curve. We estimated relative contributions of the environmental variables to the Maxent model of Neanderthal and AMH. We found that maximum temperature of the warmest month (with 58.5% contribution), minimum temperature of coldest month (with 19.7% contribution), and annual precipitation (16.5% contribution) were the most important predictor of the Neanderthal palaeodistribution (Benito et al 2017). Maximum temperature of the warmest month has a negative association with presence of the Neanderthals. Slope (with 35.6% contribution), topographic diversity (with 26% contribution) and precipitation of the warmest quarter (with 14% contribution) were the most important variables in shaping AMH palaeodistribution. Both species showed similar response to slope and habitat suitability decrease in areas with high slope. Figure 2 shows how each environmental variable affects the Maxent prediction for Neanderthal (a) and AMH (b). The curves show how the predicted probability of presence changes as each environmental variable is varied, keeping all other environmental variables at their average sample value. Discussion Hominin interbreeding is an important topic in paleo-anthropological studies but when and where it happened is remained largely unknown. Among the different hominin species interbreeding of Neanderthal and AMH is particularly important as it contributed to genetic of our own species. Here we applied SDMs and GIS and showed that Zagros Mountains of Iran as a potentially highly suitable geographic unit for niche overlap and potential interbreeding zone of the two species. Our niche models predicted niche overlap for the two species in Zagros Mountains. In support of this finding, it should be noted various research of genetic data (e.g. Lazaridis et al. 2018, Vallini et al. 2024), ecological modelling (e.g. Roberts and Stewart 2018) archaeological and genetic records (e.g. Heydari-Guran and Ghasidian 2021, Vallini et al. 2024) and fossils (Churchill et al 2022) are in concordance with our niche overlap model. The expansion of the Neanderthals to Zagros must have happened in accordance with the Palearctic environment and karstic terrains from both sides of the Black Sea, i.e. the Caucasus and Anatolia crossing towards the southern regions. The latest evidence has shown, the southernmost expanse Neanderthal, until the latitudes of around 31° in an arm shape stretched to the south in two different directions alongside of the Anti-Lebanon and Zagros Mountains (Heydari-Guran et al. 2021). Neanderthals' territories further towards east, such as those found in present-day Uzbekistan (Glantz et al. 2008), Tajikistan (Trinkaus et al. 2000) and Asian Russia (Mednikova 2011, Krause et al. 2004) are known as Central and North Asian Neanderthals. So far, the evidence of the presence of Neanderthal is consistent with Southwestern humid mountainous zones, including Anti-Lebanon in the Levant (Tillier and Arensburg 2016), Anatolia (Harvati and Roksandic 2017), Caucasus (Hajdinjak et al. 2018, Gasparyan and Glauberman 2022) and Zagros (Zanolli et al. 2019, Heydari-Guran et al. 2021). The data on the MP period in the Zagros Mountains region is rich and more up to date due to the discoveries of stratified sites associated with absolute dates, hominin physical remains, and lithic artefacts. Among the large number of MP sites, four have yielded Neanderthal fossils. The best-known of these is Shanidar Cave, where the remains of ten Neanderthals were discovered (Pomeroy et al 2020). Approximately 350 km to the southeast (around latitude 34˚), Wezmeh and Bisetun caves in the Kermanshah region have also yielded Neanderthal remains (Trinkaus and Biglari 2006, Zanolli et al. 2019). However, the recent discovery of the Neanderthal remains from Bawa Yawan Rockshelter is significant since it yielded an in situ Neanderthal tooth in association with the Zagros Mousterian lithic artefacts (Heydari-Guran et al 2021). The tooth has been dated to between 65 to 60 Kyr, while the age of the Mousterian layer, so far, goes back until 83 Kyr (Heydari et al 2024). Due to the evidence of various physical remains, it has been determined that the region of Southwest Asia was inhabited by AMHs modern humans in the late Pleistocene. AMHs have inhabited the Levant during at least two periods between 177-194 Kyr, as evidenced at the site of Misliya (Hershkovitz 2018) and between ~120 Kyr and 90 Kyr, as shown at the sites of Skhul and Qafzeh (Mercier et al 1993), before the area was permanently occupied by H. sapiens around 55,000 years (Hershkovitz et al 2015). There is a vast data of hominin (including AMH) occupations from 400 kyr to 50 Kyr in Arabia associated with Eastern African lithic technology (see: Groucutt et al 2021 and references therein) and moreover physical remains including finger bone from Al-Wusta dated to ca. 85 Kyr (Groucutt et al. 2018), all indicate Arabia as a gateway to Eurasia during Middle to Late Pleistocene. There is evidence of the presence of nun-Mousterian MP artifacts dated back to 80 Kyr in the southern regions of the Persian Plateau, both in the Zagros (Heydari et al 2021) and in the southern to central parts of the Persian Plateau (Heydari-Guran and Ghasidian 2021). There are several reasons that why Zagros Mountains is a suitable place for two species niche overlap and potential interbreeding zone. Firstly, Zagros Mountains is characterised with environmental conditions of the Palearctic realm which is known as a birth place of Neanderthal (references). At the same time surrounding areas of the Zagros are characterised with environmental conditions of Afro-Arabian; the birth place of AMH. Thus, Zagros Mountains could have been visited repeatedly by people who were living in the border areas of the Palaearctic and Afro-Arabian realms during Pleistocene climatic shifts. Therefore, the possibility of interaction between different hominins, including AMHs and Neanderthals, was higher in these areas. Secondly, Zagros is a vast region (overs 1500 km from Lake Van at the Turkish Kurdistan to south-eastern Iran) to support stable human populations. Thirdly, Zagros is exceptionally diverse in terms of topography and biodiversity (Heydari-Guran 2014; Kafash et al 2020, 2021; Yousefi et al., 2022, 2023) making it capable of supporting presence of two sub-species at the same time. Our initial hypothesis was that climatic factors would be the predominant force in predicting the distribution of both Neanderthal and AMHs. However, our findings revealed a nuanced picture: while climate indeed emerged as the key determinant of Neanderthal habitats, AMHs distribution was significantly influenced by topographical variations. Climate is homogenous but the topography is heterogenous across the AMHs distribution areas. These likely grants topography a more pronounced role in sculpting the distribution patterns of AMH. Our study contributes to the growing body of evidence that underscores the complex interplay between environmental factors in determining species distribution. Our results are in line with prey overlap (Yousefi et al. 2020) showed that annual precipitation and maximum temperature of the warmest month were the most important predictor of Neanderthal distribution on the Persian Plateau. Climate was the most important determinant of the Neanderthal distribution in Europe and the Irano-Turanian region during the Last Interglacial, yet, the influence of topography was confined to local scales (Benito et al. 2017). Conclusion One particular application of species distribution models (SDMs) is to identify suitable areas for the presence of target species where no observations have been made (Fois et al. 2015, Becker et al. 2022). Field surveys guided by SDMs have led to the discovery of new populations and rare species (Fois et al., 2015; Becker et al., 2022), thereby proving the utility of SDMs in this context. Our model, which predicts the interbreeding areas of Neanderthals and AMH, is assigned a very high priority for future field investigations and excavations. Although field testing of SDMs in archaeological studies is limited (Yousefi et al. 2020), we encourage Iranian archaeologists to conduct field excavations in this potential interbreeding area to evaluate the practicality of the models in archaeological research. Moreover, the use of SDMs can guide the allocation of resources for archaeological excavations, ensuring that efforts are concentrated in areas with the highest potential for significant findings. By prioritizing these high-probability locations, researchers can maximize the efficiency of their fieldwork, leading to more targeted and fruitful excavations. Our research shows the high potential of the Persian Plateau, in particular, its western parts: the Zagros Mountains. Northern to central Zagros region is one of the nexus points for Neanderthal migration into in southwest Asia as ecozones shifted MIS 5 and the entry point for AMHs as they spread across southern plateau and first geographical contact at the southern end of the Neanderthal occupation. The possibility of attracting different hominin groups in the Zagros is justified by the geographical conditions of this region, since it is located in two different biogeographical zones of Palearctic and Afro-Arabian realms. The border areas of two realms are important in biology since they operate as refugium for species from glacial environmental conditions. Consequently, some parts of the Zagros Mountains could have been visited repeatedly by people who were living in the border areas of the Palaearctic and Afro-Arabian realms during Pleistocene climatic shifts. Therefore, the possibility of interaction between different hominins, including AMHs and Neanderthals, was possibly higher in these areas. In concordance with our initial expectation the interaction and potential interbreeding zone of Neanderthal and AMHs located in the contact zone of Afro-Arabian and Palearctic, the Zagros Mountains. These mountains have high topographic and species diversity providing rich food resources to humans (Heydari-Guran and Ghasidian 2020). The Zagros Mountains is vast, making it an ideal place for two species to overlap. These mountains facilitated niche overlap of other taxa with similar niche in the same habitat (Grant 1975, Yousefi et al. 2020). The mountains are known to play a very significant role in species distribution by acting as a dispersal barrier or as a dispersal corridor (Yousefi et al. 2023). All above mentioned facts support the results of our study. Our findings can be further supported by new fossil discovery in the Zagros mountains and new genetic data (Vallini et al., 2024). According to our hypothesis, a migration route into the Central Plateau from other directions, including the south via Arabia, the Persian Gulf, and the Oman Sea is plausible. This route might have followed the coastal lines towards the north, and eventually into the inner parts of the Persian Plateau. Recent evidence of hominin occupations scattered on the surface in the areas located on the southernmost part of the Persian Plateau supports our hypothesis (Heydari-Guran and Ghasidian 2021, Heydari et al 2021, Shoaee et al 2021). Before this study, our understanding of the interbreeding of AMH and Neanderthal was based on genetic data alone (Tobler et al 2023). Here for the first time, we applied SDMs as additional and independent line of information to locate possible geographic location of the two species interbreeding. Our study identified Persian Plateau, particularly Zagros Mountains as potential interbreeding area for AMHs and Neanderthals. Adding to our results, the idea of the Persian Plateau as a hub for Homo sapiens after the main out of Africa dispersal (Vallini et al., 2024), we conclude that this Plateau contributed significantly to the hominin distribution (Heydari-Guran 2014, Yousefi et al. 2020), dispersal (Heydari-Guran and Ghasidian 2021, Ghasidian et al. 2023, Shoaee et al. 2023) and evolution (Ghasidian et al. 2019, Vallini et al. 2024) and awaiting many exciting discoveries to shed light on human evolution and dispersal. Methods Archaeological sites. We obtained 38 occurrence points for Neanderthal and 45 for AMHs, extracted from multiple sources including “Role of Culture in Early Expansions of Humans Out of Africa (ROCEEH: http://www.roceeh.net) Database (ROAD30,31) and Appendix S1 in Benito et al (2017). Association of each arachnological site to one/two species was made by fossil records and lithic artefacts. Since our research fucuses on the time frame MIS 5 (c.g.120-80 Kyr) we have just used the archaeological sites associated with presence of Neanderthals during this period for western Asia and southeastern Europe. We carefully examined each coordinate and removed duplicates. Since, environmental data were at 1 km spatial resolution we thinned the occurrence data to km to avoid pseudo-replication (Yousefi et al 2020). Environmental predictors. We considered environmental variables related to past climate and topography to reconstruct AMH and Neanderthals niche during the MIS 5. As paleoclimatic variables we added maximum temperature of warmest month, minimum temperature of coldest month, annual precipitation and precipitation of warmest quarter to the niche models for time span of MIS 5. Paleoclimatic data were obtained from Oscillayers, which is a dataset of climatic oscillations over Plio-Pleistocene time-scales at high spatial-temporal resolution (Gamisch 2019). We estimated average values for each of above-mentioned variables during the MIS 5 using the raster package v. 3.4–13 (Hijmans 2021) implemented in R environment (R Core Team 2023). To take into account topography we included slope and topographic heterogeneity (Benito et al. 2017, Yousefi et al. 2020) which were downloaded from EarthEnv (Amatulli et al. 2018). In order to avoid multicollinearity among the predictors we calculated a variance inflation factor (VIF; [Quinn and Keough 2002]) using the “vifstep” function in the “usdm” package (Naimi, 2015) and ensured that collinearity among predictors is low (VIF < 10). Ecological niche modelling. In this study, we used Maximum Entropy Modelling approach (Phillips et al. 2006) to reconstruct ecological niche models of Homo sapiens and Homo neanderthalensis during the MIS 5. Maxent version 3.4.4 was used to build the niche models (Phillips et al. 2006). We used Maxent model because it has been shown to perform better than other niche modelling methods (Elith et al. 2006, Guisan et al. 2017). Then we overlapped the two palaeodistribution models to identify potential areas for their contact zones in QGIS (www.qgis.org). Performance of the niche models was assessed using the Area Under the Curve (AUC) metric of the Receiving Operator Characteristic (ROC) curve (Phillips et al. 2006). An AUC value of 0.5 indicates that the performance of the model is not better than random, while values closer to 1.0 indicate better model performance (Swets 1988). The ROC plots were created by selecting 80% of the data for training and 20% for testing. Declarations Data availability All data needed to evaluate the conclusions in the paper are present in the paper or the references cited here within. We obtained archaeological sites data from the ROCEEH Out of Africa Database (ROAD) (http://www.roceeh.org) and references cited in the manuscript. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4298125","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":294203686,"identity":"63416c31-2838-40ad-80ec-01e0b47d3e68","order_by":0,"name":"Saman Heydari-Guran","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2klEQVRIiWNgGAWjYFAD9gYgYWBBihaeAyAtEqRokUgAk4QVyrefTvxc8OuwvLnk86sbfhRIMPC3dyfg1WJwJnez9My+w4Y7Z+eU3ewBOkzizNkN+LUw5G6Q5u05zLjhdk7aDR6gFgOJXPxa5Pvfbv4N1GK/4eaZtJt/iNHCcCN3mzTPj8OJG26wH7tNlC0GN95us+ZtSE/ecCaH7baMgQQPQb/I9+duvs3zx9p2w/Hjz26++WMjx9/eS8BhIMDYBiJ5DMAkYeVg8AdEsD8gUvUoGAWjYBSMNAAAjQNMt1BuQH4AAAAASUVORK5CYII=","orcid":"","institution":"University of Cologne","correspondingAuthor":true,"prefix":"","firstName":"Saman","middleName":"","lastName":"Heydari-Guran","suffix":""},{"id":294203687,"identity":"8f9ba48f-ec6f-4cad-b5fc-4a68421c493d","order_by":1,"name":"Masood Yosefi","email":"","orcid":"","institution":"Neanderthal Museum","correspondingAuthor":false,"prefix":"","firstName":"Masood","middleName":"","lastName":"Yosefi","suffix":""},{"id":294203688,"identity":"ef0f776d-38e1-4c48-b158-4ff8c75cc363","order_by":2,"name":"Anoshe Kafash","email":"","orcid":"","institution":"School of Culture and Society, Department of Archeology and Heritage Studies , Aarhus","correspondingAuthor":false,"prefix":"","firstName":"Anoshe","middleName":"","lastName":"Kafash","suffix":""},{"id":294203689,"identity":"9af96220-02f9-40ff-b865-c6bee8555d2f","order_by":3,"name":"Elham Ghasidian","email":"","orcid":"","institution":"Neanderthal Museum","correspondingAuthor":false,"prefix":"","firstName":"Elham","middleName":"","lastName":"Ghasidian","suffix":""}],"badges":[],"createdAt":"2024-04-20 15:40:36","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4298125/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4298125/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-024-70206-y","type":"published","date":"2024-09-03T16:05:42+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":55335612,"identity":"22a46d86-dbed-4564-b33f-35775fb4ff96","added_by":"auto","created_at":"2024-04-25 22:21:28","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":590511,"visible":true,"origin":"","legend":"\u003cp\u003eHabitat suitability models of the two \u003cem\u003eHomo\u003c/em\u003e species and their potential contact and interbreeding zone. This figure is generated in QGIS 3.14.1 (www.qgis.org).\u003c/p\u003e","description":"","filename":"Figure1.png","url":"https://assets-eu.researchsquare.com/files/rs-4298125/v1/ff98a24c79fb26f146b82815.png"},{"id":55335761,"identity":"7e291eda-67b9-4261-8f26-1bcaff877cb8","added_by":"auto","created_at":"2024-04-25 22:29:28","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":92876,"visible":true,"origin":"","legend":"\u003cp\u003eResponse curves showing how the presence of Neanderthal (a) and AMH (b) is related to the environmental variables (Bio5: maximum temperature of warmest month, Bio6: minimum temperature of coldest month, Bio12: annual precipitation and Bio18: precipitation of warmest quarter). (https://biodiversityinformatics.amnh.org/open_source/maxent/).\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-4298125/v1/c6df7de4bbeecbdd4dff9ab8.png"},{"id":64185964,"identity":"78d7f3c3-4213-42f7-aa82-f1ccdd94a687","added_by":"auto","created_at":"2024-09-09 16:23:11","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":946737,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4298125/v1/da37b49c-fa7b-4dbc-a4a8-ec8e40b02e07.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Reconstructing Contact and Potential Interbreeding Geographical Zone for Neanderthals and Anatomically Modern Humans","fulltext":[{"header":"Introduction","content":"\u003cp\u003eFollowing the ground-breaking discovery of biocultural admixture of Late Pleistocene different early human groups of Neanderthals, archaic/modern human, and Denisovans, a large and growing body of research concerning the nature and evolutionary history of these events are presented. Besides of significant consequences the biological exchanges that have had to subspecies (e.g. Quach et al 2016) and related issues, the time (Green et al 2010; Pr\u0026uuml;fer et al 2014 and 2017)\u003csup\u003e\u0026nbsp;\u003c/sup\u003eand geography of contact and interbreeding are subject of intense debate (Stewart and Stringer 2012). Neanderthals are an extinct lineage of hominins that emerged around 400 Kyr and died off around 40 Kyr (Higham et al 2014). Fossil localities and morphological evidence of Neanderthals indicate that they were companionable with the Palearctic biogeographical realm, which includes from western Europe to Altai Mountains in Siberia at 55\u0026deg; latitude and down until around 31\u0026deg; 1atitude in the Western Asia (Kolobova et al. 2020, Heydari-Guran et al. 2021, Been et al. 2017). The chronological settlement patterns of the Neanderthals\u0026apos; sites indicate of their expansion to the east and south west Asia at least from 150 Kyr (Gr\u0026uuml;n and Stringer 2000). \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;At the other side, anatomically modern humans (AMH) have evolved in Africa more than 300 Kyr (Hublin et al 2017; Richter et al 2017). The evidence including physical remains and morphological analyses suggest that they have exited Africa over and over during a period of at least 200kyr (Groucutt et al 2018; Hershkovitz et al 2018; Harvati et al 2019). AMH, also reached Eastern Asia around 120kyr (Liu et al 2010) and later on into Europe around 60 Kyr (Slimak et al 2022; Hublin et al 2020). Recent accurate archaeological and Paleoenvironmental data suggest that AMH rapidly adapted to the new and extreme environments beyond Africa, such as high-plateaus, mountain systems and palearctic ecosystem (Roberts and Stewart 2018). Meanwhile, the archaeological and fossil\u003cspan dir=\"RTL\"\u003e\u0026nbsp;\u003c/span\u003eevidence indicate that AMH have entered mainly to south western Asia during MIS 5 (Groucutt et al 2012; Scerri et al 2015; Groucutt et al 2018, Heydari-Guran and Ghasidian 2021).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;There is strong evidence of multiple interbreeding between two groups of Neanderthal and archaic/modern human in the western Eurasia (e.g. Pr\u0026uuml;fer et al 2014; Slon et al 2018). Meanwhile, many attempts have been made to estimate the time for this admixture and significant successes have been achieved (e.g. Krause et al 2010; Yuan et al 2021). \u0026nbsp;The Paleogenetic studies support interbreeding between AMH and Neanderthals go back to at least 130 Kyr (Pr\u0026uuml;fer et al 2017). In some studies, researchers suggest that the lower latitude regions of southwestern Asia have a high potential for the first overlap between Neanderthal and AMH. Sanchez Go\u0026ntilde;i (2020) uses the patterns of expansion of Neanderthal and AMH and share same ecological niches under the certain climatic conditions during the late Pleistocene. Recently Churchill et al. (2022) found facial morphological similarities between Neanderthals and AMH in the Near East, indicating key region for the interbreeding between the two lineages. But still it is unclear where the two species met and probably interbreed. Species Distribution Models (SDMs) are very practical tools to investigate geography of the two species palaeodistribution and potential interbreeding areas (Ruan et al 2023). SDMs which have found important application in paleobiogeography, archaeological and paleo-anthropological (Varela et al., 2011; Svenning et al., 2011\u003csup\u003e26\u003c/sup\u003e; Franklin et al., 2015; Giampoudakis et al., 2017; Benito et al., 2017). They use occurrence data of a target species including ancient humans and paleoenvironmental variables to calculate probability of a species or hominin species\u0026rsquo; presence in a defined geographic region\u0026nbsp;(Guisan et al. 2017)\u003cspan dir=\"RTL\"\u003e.\u003c/span\u003e These models have successfully been used to reconstruct different hominin species (Giampoudakis et al 2017; Benito et al 2017; Jones et al 2019), identify refugia during the ice ages and reconstructing dispersal corridors (Kafash et al 2023), their niche overlap with each other (Ruan et al 2023) and niche overlap with their prey species (Yousefi et al 2020\u003csup\u003e33\u003c/sup\u003e). For instance, Ruan et al. (2023) have used SDMs to identify contact zone of Neanderthal and Denisova. In another study, Benito et al. (2017) applied the ecological niche and distribution of the Neanderthal during the Last Interglacial in Europe and the Irano-Turonian region. Thus, SDMs are used to model Neanderthal and AMHs palaeodistribution and locate geography of their niche overlap (Benito et al. 2017; Ruan et al. 2023).\u003c/p\u003e\n\u003cp\u003eThe aims of present study are to first reconstruct palaeodistribution of Neanderthal and AMHs during the MIS 5, identify the most important predictor of the two species, investigate the two species response to environmental variables and identify geographic overlap for the two species at the meeting zone of three continents, Asia, Africa and Europe. Considering previous studies on the importance of Southwest Asia the African continent and southern part of the Middle East are located in the Afro-Arabian biogeographical realms () which matched with distribution of AMH and the lands above Levant. The northern part of the Persian and Anatolian plateaus, Caucasus to European continent and Siberia are setting in the Palearctic realm were considered to be the distribution range of the Neanderthals. Here, we hypothesized that these two species first met and interbreed at the border of the two biogeographic realms where environmental condition facilitated niche overlap and resource partitioning by providing highly diverse habitat rich in resources. In this regard climate is a major determinant of species distributions (Sexton et al 2009) particular at large spatial scales, we are expecting climate to be more effective than topography in shaping the two sub species of Neanderthal and AMH.\u0026nbsp;\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003eReconstructing contact and interbreeding zone.\u003c/strong\u003e The models developed in this study for Neanderthal (AUC=941) and AMH (AUC= 895) performed well according to the AUC model performance metric. Our model of paleodistribution of Neanderthal shows that north and west of Mediterranean Sea toward Levant, vast patches in Turkey, around the Black Sea, south of Caspian Sea, Taurus, Caucasus and Zagros Mountains have high suitability for the species during the MIS 5 (Figure 1). AMH paleo-distribution model identified large and continues suitable patches and in Africa, Arabia and Iranian Plateau. Our model identified Zagros Mountains as a contact and potential interbreeding zone\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eVariable importance and response curve.\u0026nbsp;\u003c/strong\u003e We estimated relative contributions of the environmental variables to the Maxent model of Neanderthal and AMH. We found that maximum temperature of the warmest month (with 58.5% contribution), minimum temperature of coldest month (with 19.7% contribution), and annual precipitation (16.5% contribution) were the most important predictor of the Neanderthal palaeodistribution (Benito et al 2017). Maximum temperature of the warmest month has a negative association with presence of the Neanderthals. Slope (with 35.6% contribution), topographic diversity (with 26% contribution) and precipitation of the warmest quarter (with 14% contribution) were the most important variables in shaping AMH palaeodistribution. Both species showed similar response to slope and habitat suitability decrease in areas with high slope. Figure 2 shows how each environmental variable affects the Maxent prediction for Neanderthal (a) and AMH (b). The curves show how the predicted probability of presence changes as each environmental variable is varied, keeping all other environmental variables at their average sample value.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eHominin interbreeding is an important topic in\u0026nbsp;paleo-anthropological studies but when and where it happened is remained largely unknown. Among the different hominin species interbreeding of Neanderthal and AMH is particularly important as it contributed to genetic of our own species. Here we applied SDMs and GIS and showed that Zagros Mountains of Iran as a potentially highly suitable geographic unit for niche overlap and potential interbreeding zone of the two species.\u003c/p\u003e\n\u003cp\u003eOur niche models predicted niche overlap for the two species in Zagros Mountains. In support of this finding, it should be noted various research of genetic data (e.g. Lazaridis et al. 2018, Vallini et al. 2024), ecological modelling (e.g. Roberts and Stewart 2018) archaeological and genetic records (e.g. Heydari-Guran and Ghasidian 2021, Vallini et al. 2024) and fossils (Churchill et al 2022) are in concordance with our niche overlap model. The expansion of the Neanderthals to Zagros must have happened in accordance with the Palearctic environment and karstic terrains from both sides of the Black Sea, i.e. the Caucasus and Anatolia crossing towards the southern regions. The latest evidence has shown, the southernmost expanse Neanderthal, until the latitudes of around 31\u0026deg; in an arm shape stretched to the south in two different directions alongside of the Anti-Lebanon and Zagros Mountains (Heydari-Guran et al. 2021). Neanderthals\u0026apos; territories further towards east, such as those found in present-day Uzbekistan (Glantz et al. 2008), Tajikistan (Trinkaus et al. 2000) and Asian Russia (Mednikova 2011, Krause et al. 2004) are known as Central and North Asian Neanderthals. So far, the evidence of the presence of Neanderthal is consistent with Southwestern humid mountainous zones, including Anti-Lebanon in the Levant (Tillier and Arensburg 2016), Anatolia (Harvati and Roksandic 2017), Caucasus (Hajdinjak et al. 2018, Gasparyan and Glauberman 2022) and Zagros (Zanolli et al. 2019, Heydari-Guran et al. 2021).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;The data on the MP period in the Zagros Mountains region is rich and more up to date due to the discoveries of stratified sites associated with absolute dates, hominin physical remains, and lithic artefacts. Among the large number of MP sites, four have yielded Neanderthal fossils. The best-known of these is Shanidar Cave, where the remains of ten Neanderthals were discovered (Pomeroy et al 2020). Approximately 350 km to the southeast (around latitude 34˚), Wezmeh and Bisetun caves in the Kermanshah region have also yielded Neanderthal remains (Trinkaus and Biglari 2006, Zanolli et al. 2019). However, the recent discovery of the Neanderthal remains from Bawa Yawan Rockshelter is significant since it yielded an in situ Neanderthal tooth in association with the Zagros Mousterian lithic artefacts (Heydari-Guran et al 2021). The tooth has been dated to between 65 to 60 Kyr, while the age of the Mousterian layer, so far, goes back until 83 Kyr (Heydari et al 2024).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;Due to the evidence of various physical remains, it has been determined that the region of Southwest Asia was inhabited by AMHs modern humans in the late Pleistocene. AMHs have inhabited the Levant during at least two periods between 177-194 Kyr, as evidenced at the site of Misliya (Hershkovitz 2018) and between ~120 Kyr and 90 Kyr, as shown at the sites of Skhul and Qafzeh (Mercier et al 1993), before the area was permanently occupied by H. sapiens around 55,000 years (Hershkovitz et al 2015). There is a vast data of hominin (including AMH) occupations from 400 kyr to 50 Kyr in Arabia associated with Eastern African lithic technology (see: Groucutt et al 2021 and references therein) and moreover physical remains including finger bone from Al-Wusta dated to ca. 85 Kyr (Groucutt et al. 2018), all indicate Arabia as a gateway to Eurasia during Middle to Late Pleistocene. There is evidence of the presence of nun-Mousterian MP artifacts dated back to 80 Kyr in the southern regions of the Persian Plateau, both in the Zagros (Heydari et al 2021) and in the southern to central parts of the Persian Plateau (Heydari-Guran and Ghasidian 2021).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;There are several reasons that why Zagros Mountains is a suitable place for two species niche overlap and potential interbreeding zone. Firstly, Zagros Mountains is characterised with environmental conditions of the Palearctic realm which is known as a birth place of Neanderthal (references). At the same time surrounding areas of the Zagros are characterised with environmental conditions of Afro-Arabian; the birth place of AMH. Thus, Zagros Mountains could have been visited repeatedly by people who were living in the border areas of the Palaearctic and Afro-Arabian realms during Pleistocene climatic shifts. Therefore, the possibility of interaction between different hominins, including AMHs and Neanderthals, was higher in these areas. Secondly, Zagros is a vast region (overs 1500 km from Lake Van at the Turkish Kurdistan to south-eastern Iran) to support stable human populations. Thirdly, Zagros is exceptionally diverse in terms of topography and biodiversity (Heydari-Guran 2014; Kafash et al 2020, 2021; Yousefi et al., 2022, 2023) making it capable of supporting presence of two sub-species at the same time.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eOur initial hypothesis was that climatic factors would be the predominant force in predicting the distribution of both Neanderthal and AMHs. However, our findings revealed a nuanced picture: while climate indeed emerged as the key determinant of Neanderthal habitats, AMHs distribution was significantly influenced by topographical variations. Climate is homogenous but the topography is heterogenous across the AMHs distribution areas. These likely grants topography a more pronounced role in sculpting the distribution patterns of AMH. Our study contributes to the growing body of evidence that underscores the complex interplay between environmental factors in determining species distribution. Our results are in line with prey overlap (Yousefi et al. 2020) showed that annual precipitation and maximum temperature of the warmest month were the most important predictor of Neanderthal distribution on the Persian Plateau. Climate was the most important determinant of the Neanderthal distribution in Europe and the Irano-Turanian region during the Last Interglacial, yet, the influence of topography was confined to local scales (Benito et al. 2017).\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eOne particular application of species distribution models (SDMs) is to identify suitable areas for the presence of target species where no observations have been made (Fois et al. 2015, Becker et al. 2022). Field surveys guided by SDMs have led to the discovery of new populations and rare species (Fois et al., 2015; Becker et al., 2022), thereby proving the utility of SDMs in this context. Our model, which predicts the interbreeding areas of Neanderthals and AMH, is assigned a very high priority for future field investigations and excavations. Although field testing of SDMs in archaeological studies is limited (Yousefi et al. 2020), we encourage Iranian archaeologists to conduct field excavations in this potential interbreeding area to evaluate the practicality of the models in archaeological research.\u0026nbsp;Moreover, the use of SDMs can guide the allocation of resources for archaeological excavations, ensuring that efforts are concentrated in areas with the highest potential for significant findings. By prioritizing these high-probability locations, researchers can maximize the efficiency of their fieldwork, leading to more targeted and fruitful excavations.\u003c/p\u003e\n\u003cp\u003eOur research shows the high potential of the Persian Plateau, in particular, its western parts: the Zagros Mountains. Northern to central Zagros region is one of the nexus points for Neanderthal migration into in southwest Asia as ecozones shifted MIS 5 and the entry point for AMHs as they spread across southern plateau and first geographical contact at the southern end of the Neanderthal occupation.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe possibility of attracting different hominin groups in the Zagros is justified by the geographical conditions of this region, since it is located in two different biogeographical zones of Palearctic and Afro-Arabian realms. The border areas of two realms are important in biology since they operate as refugium for species from glacial environmental conditions. Consequently, some parts of the Zagros Mountains could have been visited repeatedly by people who were living in the border areas of the Palaearctic and Afro-Arabian realms during Pleistocene climatic shifts. Therefore, the possibility of interaction between different hominins, including AMHs and Neanderthals, was possibly higher in these areas.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn concordance with our initial expectation the interaction and potential interbreeding zone of Neanderthal and AMHs located in the contact zone of Afro-Arabian and Palearctic, the Zagros Mountains. These mountains have high topographic and species diversity providing rich food resources to humans (Heydari-Guran and Ghasidian 2020). The Zagros Mountains is vast, making it an ideal place for two species to overlap. \u0026nbsp;These mountains facilitated niche overlap of other taxa with similar niche in the same habitat (Grant 1975, Yousefi et al. 2020). The mountains are known to play a very significant role in species distribution by acting as a dispersal barrier or as a dispersal corridor (Yousefi et al. 2023). All above mentioned facts support the results of our study. Our findings can be further supported by new fossil discovery in the Zagros mountains and new genetic data (Vallini et al., 2024). According to our hypothesis, a migration route into the Central Plateau from other directions, including the south via Arabia, the Persian Gulf, and the Oman Sea is plausible. This route might have followed the coastal lines towards the north, and eventually into the inner parts of the Persian Plateau. Recent evidence of hominin occupations scattered on the surface in the areas located on the southernmost part of the Persian Plateau supports our hypothesis\u003cspan dir=\"RTL\"\u003e\u0026nbsp;\u003c/span\u003e(Heydari-Guran and Ghasidian 2021, Heydari et al 2021, Shoaee et al 2021).\u003c/p\u003e\n\u003cp\u003eBefore this study, our understanding of the interbreeding of AMH and Neanderthal was based on genetic data alone (Tobler et al 2023). Here for the first time, we applied SDMs as additional and independent line of information to locate possible geographic location of the two species interbreeding. Our study identified Persian Plateau, particularly Zagros Mountains as potential interbreeding area for AMHs and Neanderthals. Adding to our results, the idea of the Persian Plateau as a hub for \u003cem\u003eHomo sapiens\u003c/em\u003e after the main out of Africa dispersal (Vallini et al., 2024), we conclude that this Plateau contributed significantly to the hominin distribution (Heydari-Guran 2014, Yousefi et al. 2020), dispersal (Heydari-Guran and Ghasidian 2021, Ghasidian et al. 2023, Shoaee et al. 2023) and evolution (Ghasidian et al. 2019, Vallini et al. 2024) and awaiting many exciting discoveries to shed light on human evolution and dispersal.\u0026nbsp;\u003c/p\u003e"},{"header":"Methods","content":"\u003cp\u003e\u003cstrong\u003eArchaeological sites.\u0026nbsp;\u003c/strong\u003e We obtained 38 occurrence points for Neanderthal and 45 for AMHs, extracted from multiple sources including \u0026ldquo;Role of Culture in Early Expansions of Humans Out of Africa (ROCEEH: http://www.roceeh.net) Database (ROAD30,31) and Appendix S1 in Benito et al (2017). Association of each arachnological site to one/two species was made by fossil records and lithic artefacts. \u0026nbsp;Since our research fucuses on the time frame MIS 5 (c.g.120-80 Kyr) we have just used the archaeological sites associated with presence of Neanderthals during this period for western Asia and southeastern Europe. \u0026nbsp;We carefully examined each coordinate and removed duplicates. Since, environmental data were at 1 km spatial resolution we thinned the occurrence data to km to avoid pseudo-replication (Yousefi et al 2020).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEnvironmental predictors.\u0026nbsp;\u003c/strong\u003eWe considered environmental variables related to past climate and topography to reconstruct AMH and Neanderthals niche during the MIS 5. As paleoclimatic variables we added maximum temperature of warmest month, minimum temperature of coldest month, annual precipitation and precipitation of warmest quarter to the niche models for time span of MIS 5. Paleoclimatic data were obtained from Oscillayers, which is a dataset of climatic oscillations over Plio-Pleistocene time-scales at high spatial-temporal resolution (Gamisch 2019). We estimated average values for each of above-mentioned variables during the MIS 5 using the raster package v. 3.4\u0026ndash;13 (Hijmans 2021) implemented in R environment (R Core Team 2023). To take into account topography we included slope and topographic heterogeneity (Benito et al. 2017, Yousefi et al. 2020) which were downloaded from EarthEnv (Amatulli et al. 2018). In order to avoid multicollinearity among the predictors we calculated a variance inflation factor (VIF; [Quinn and Keough 2002]) using the \u0026ldquo;vifstep\u0026rdquo; function in the \u0026ldquo;usdm\u0026rdquo; package (Naimi, 2015) and ensured that collinearity among predictors is low (VIF \u0026lt; 10).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eEcological niche modelling.\u003c/strong\u003e In this study, we used Maximum Entropy Modelling approach (Phillips et al. 2006) to reconstruct ecological niche models of \u003cem\u003eHomo sapiens\u003c/em\u003e and \u003cem\u003eHomo neanderthalensis\u003c/em\u003e during the MIS 5. Maxent version 3.4.4 was used to build the niche models (Phillips et al. 2006). We used Maxent model because it has been shown to perform better than other niche modelling methods (Elith et al. 2006, Guisan et al. 2017). Then we overlapped the two palaeodistribution models to identify potential areas for their contact zones in QGIS (www.qgis.org). Performance of the niche models was assessed using the Area Under the Curve (AUC) metric of the Receiving Operator Characteristic (ROC) curve (Phillips et al. 2006). An AUC value of 0.5 indicates that the performance of the model is not better than random, while values closer to 1.0 indicate better model performance (Swets 1988). The ROC plots were created by selecting 80% of the data for training and 20% for testing.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eData availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll data needed to evaluate the conclusions in the paper are present in the paper or the references cited here within. We obtained archaeological sites data from the ROCEEH Out of Africa Database (ROAD) (http://www.roceeh.org) and references cited in the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe are grateful to the ROCEEH Out of Africa Database (ROAD) (http://www.roceeh.org) for making the data available to us.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eAuthor contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eSHG, M.Y, and contributed equally to the design and conceptualization and writing and A.K in performance and analysis of the data. \u0026nbsp;E. G. in review \u0026amp; editing. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe research is funded by the funded by Deutsche Forschungsgemeinschaft (DFG), Priority program 2176:\u003c/p\u003e\n\u003cp\u003eThe Iranian highlands: Resiliencies and integration in pre-modern societies, \u0026ldquo;Last Neanderthal and early Homo sapiens occupations in the Bawa Yawan Rockshelter, Kermanshah, West-Central Zagros Mountains (Iran). Project number 402379177. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;The authors declare no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAmatulli, G., Domisch, S., Tuanmu, M.-N., Parmentier, B., Ranipeta, A., Malczyk, J., and Jetz, W. (2018). 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Journal of human evolution, 135, 102643.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Zagros Mountains, Ecological niche, palaeoenvironment, Neanderthals, anatomically modern humans","lastPublishedDoi":"10.21203/rs.3.rs-4298125/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4298125/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Whilst the admixture of the Neanderthal and anatomically modern human is precisely proven, due to the shortage of fossils and absence of appropriate DNA, the timing and geography of it, are the subjects of questions. In this study, we applied ecological niche modelling (maximum entropy approach) and GIS to reconstruct Neanderthal and modern human palaeodistribution and identify their contact and potential interbreeding zone during the marine isotope stage 5. We used climatic variables characterising environmental conditions of marine isotope stage 5 ca. 120 to 80 thousand years ago (Kyr) along with topography and coordinates of Neanderthal and modern human archaeological sites to build each species palaeodistribution. Overlapping the models showed that Zagros Mountains was a contact and potential interbreeding zone for the two sub-human species. We believe that Zagros Mountains acted as a corridor connected the Palearctic/Afro-Arabian realms, facilitated northward dispersal of anatomically modern human and southward dispersal of Neanderthal during marine isotope stage 5. Our analyses are comparable with archaeological and genetic evidence collected during the last decades.","manuscriptTitle":"Reconstructing Contact and Potential Interbreeding Geographical Zone for Neanderthals and Anatomically Modern Humans","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-04-25 22:21:24","doi":"10.21203/rs.3.rs-4298125/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2024-06-23T11:13:33+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-06-11T15:11:45+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-06-03T08:44:03+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"38603928872506425918029724366388503094","date":"2024-05-21T14:03:21+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"30283430298047002543909106573375043305","date":"2024-05-07T07:37:14+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-05-02T08:38:04+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-04-25T18:37:49+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2024-04-22T16:55:07+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-04-22T16:37:52+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2024-04-20T15:34:24+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"8b8ef12c-0c05-43bf-bed9-838e3727cb75","owner":[],"postedDate":"April 25th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-09-09T16:14:50+00:00","versionOfRecord":{"articleIdentity":"rs-4298125","link":"https://doi.org/10.1038/s41598-024-70206-y","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2024-09-03 16:05:42","publishedOnDateReadable":"September 3rd, 2024"},"versionCreatedAt":"2024-04-25 22:21:24","video":"","vorDoi":"10.1038/s41598-024-70206-y","vorDoiUrl":"https://doi.org/10.1038/s41598-024-70206-y","workflowStages":[]},"version":"v1","identity":"rs-4298125","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4298125","identity":"rs-4298125","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}