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Initial Upper Paleolithic

The Initial Upper Paleolithic (IUP) is a transitional techno-cultural of the early period, characterized by the production of blades using methods that blend Levallois techniques—such as faceted platforms, hard hammer percussion, and flat-faced cores—with emerging Upper Paleolithic traits like unidirectional volumetric reduction and laminar . This phase originated in the around 50,000–45,000 years ago, as evidenced by assemblages from sites like Boker Tachtit (layer 4) and Ksar Akil (layers XXV–XXI), where it marks a shift from bidirectional Levallois-like flaking to more specialized technologies. Geographically, the IUP extends across , from the Levant and (e.g., Grotte Mandrin in , dated to ~54,000 years ago) to Central and East (e.g., in and Tolbor Valley in , dated to ~47,000–40,000 calibrated years ), spanning over 230,000 km² in the latter region alone. Key features of IUP assemblages include asymmetrical core reduction for elongated blades, intentional blade fragmentation for tool production, and burin-core reduction yielding small blades or bladelets, often involving 13–21 procedural units in a complex knapping system comparable to Middle Stone Age technologies. These technologies are associated with the earliest evidence of Homo sapiens dispersals out of Africa, replacing Neanderthals in Europe around 45,000 years ago, as seen at Bacho Kiro Cave in Bulgaria, where IUP layers contain stone tools, bone implements, and personal ornaments linked to modern human arrivals. The IUP is interpreted as potentially representing multiple waves of migration—such as an initial pulse ~54,000 years ago into the Rhône Valley and a second ~45,000 years ago into Atlantic Europe—rather than a single event, though debates persist on whether it reflects cultural transmission or independent technological convergence across regions.

Definition and Chronology

Definition

The Initial (IUP) represents the earliest phase of the Upper Paleolithic period, dating to approximately 50,000–40,000 years (), and is characterized by the initial expansion of anatomically modern humans (Homo sapiens) into from their origins. This transitional cultural phase marks a distinct shift in lithic technologies and human adaptations, coinciding with the dispersal of modern human populations across southwest and beyond, often overlapping with lingering industries. The term "Initial Upper Paleolithic" was first proposed in the early 1980s by archaeologists Lawrence E. Marks and Hilary W. Volkman to describe the uppermost assemblages (level 4) at the Boker Tachtit site in Israel's , distinguishing them from preceding traditions and subsequent, more fully developed industries. This designation highlighted a novel techno-complex that bridged earlier and later phases, later formalized in publications by Marks and C. Richard Ferring in 1988, and expanded by Steven L. Kuhn and others in the 1990s to encompass similar assemblages across the and . The IUP thus serves as a for understanding the gradual emergence of modern human behaviors during this expansive migratory episode. Key diagnostic features of the IUP include a that blends elements, such as the Levallois reduction technique for producing predetermined flakes and points, with incipient innovations like systematic unidirectional or bidirectional production and the appearance of backed tools. These traits reflect an experimental phase in manufacture, adapting to new environmental and social demands during human colonization of diverse Eurasian landscapes. Scholars debate whether the IUP's widespread distribution signals —direct population migration carrying technological packages—or cultural transmission, where ideas and practices spread through contact without substantial demographic replacement. Evidence from genetic, archaeological, and chronological data supports complex scenarios involving both mechanisms, potentially tied to multiple waves of H. sapiens movement. Within the broader , the IUP lays foundational groundwork for , evidenced by early symbolic elements like personal ornaments and pigment use, signaling enhanced cognitive and among dispersing modern humans.

Chronological Framework

The Initial Upper Paleolithic (IUP) encompasses a timeframe of approximately 50,000–40,000 calibrated years before present (cal BP), marking the initial stage of the Upper Paleolithic and associated with early dispersals of anatomically modern humans into Eurasia. This period is established through a combination of accelerator mass spectrometry (AMS) radiocarbon dating on organic materials like bone and charcoal, and optically stimulated luminescence (OSL) dating on sedimentary deposits, which provide complementary insights into human occupation layers. These methods address the scarcity of datable material in open-air and cave contexts, though challenges persist due to potential contamination in radiocarbon samples and the need for single-grain OSL to account for incomplete bleaching in sediments. The IUP is typically phased into an Early IUP (ca. 50,000–45,000 cal ), linked to the onset of modern human migrations , and a Late IUP (ca. 45,000–40,000 cal ), reflecting technological consolidation and regional adaptations before transitioning to more established industries. For instance, at the Kara-Bom site in , IUP occupations in layers 5 and 6 yield AMS radiocarbon dates of 47,000–43,000 cal , illustrating the phasing through Bayesian modeling of stratigraphic sequences. Regional chronologies exhibit slight variations, with the recording IUP from ca. 48,000–42,000 cal , as seen in the Boker Tachtit sequence dated to 50,000–44,000 cal via integrated radiocarbon and OSL analyses. In , the IUP appears between ca. 54,000–41,000 cal , evidenced by human remains and artifacts at Grotte Mandrin in (~54,000 cal ) and in dated to 45,930–42,580 cal using ultrafiltration pretreatment on for AMS radiocarbon. East Asian records span ca. 45,000–40,000 cal , with the Shuidonggou Locality 1 site in northern providing evidence around 41,000 cal through , though OSL dates suggest up to ~46,000 cal (debated). The IUP temporally overlaps with the terminal , including the decline of industries associated with Neanderthals around 40,000 cal , and serves as a precursor to the Early , such as the proto-Aurignacian, which emerges around 43,000 cal in parts of . This transitional framework underscores the dynamic interplay of cultural innovations during a period of hominin coexistence and .

Geographic Distribution and Key Sites

Sites in the Levant and Near East

The and represent the core region for the Initial Upper Paleolithic (IUP), serving as the primary corridor for dispersal into during the late Middle to early Upper Paleolithic transition. Archaeological evidence from this area documents temporal overlap between incoming Homo sapiens populations and late Neanderthal groups, suggesting potential interactions including coexistence and possible interbreeding in shared landscapes around 50,000–40,000 years ago. Key sites in this zone reveal transitional lithic technologies and early signs of , underscoring the 's role in facilitating the spread of IUP innovations northward. Boker Tachtit, an open-air site in the Desert of southern , is the type-site for the IUP, originally defined based on its stratified assemblages exhibiting a shift from techniques to blade production. The site's four main levels, separated by sterile sediments, date to approximately 48,000–42,000 calibrated years (cal ), marking the onset and persistence of IUP occupations in the . These layers contain Levallois points alongside early bladelets, reflecting a transitional technology that combined prepared-core reduction methods with nascent prismatic blade knapping, indicative of short-term camp-like settlements by mobile hunter-gatherers. Further north, Üçağızlı Cave in the of south-central preserves IUP assemblages dated to around 45,000 cal BP, providing evidence of blade production techniques that evolved from Levallois-like methods toward more standardized forms. The site's early layers, spanning Marine Isotope Stage 3, include over 20 radiocarbon dates confirming continuous occupation and the presence of shell beads made from marine species like Columbella rustica, among the earliest known ornaments in the and signaling emerging symbolic behaviors. Ksar Akil, a deeply stratified rockshelter near in , offers one of the longest IUP sequences in the , with transitional layers beginning around 50,000 cal BP and extending into the early . These levels feature Emireh points—distinctive foliate tools retouched on both edges—as hallmarks of the IUP, alongside dentalium shell beads used for personal adornment, which appear in association with lithic tools and hearths, pointing to structured camp activities. Excavations at these Levantine sites, including renewed work by Deborah Olszewski and collaborators from the 1980s through the 2000s, have illuminated camp-like occupations characterized by domestic features such as hearths and faunal remains, emphasizing repeated short-term use by dispersing modern human groups. These investigations, building on earlier digs, highlight the region's environmental diversity—from desert wadis to coastal caves—as a conduit for IUP expansion, with artifacts underscoring adaptive strategies during a period of climatic instability.

Sites in Europe and Central Asia

The Initial Upper Paleolithic (IUP) in and reflects the expansion of modern human populations into higher latitudes and diverse continental environments following dispersals from origins around 50,000 years ago. Key sites in these regions provide evidence of technological continuity with southern assemblages, including Levallois-derived blade production and bifacial tools, while demonstrating adaptations to colder climates. These manifestations highlight the rapid colonization of north of the , with assemblages dated primarily between 47,000 and 40,000 cal BP. In , Grotte Mandrin in the Valley of represents one of the earliest IUP sites in , with layer GA dated to approximately 54,000 years ago through optically stimulated luminescence and associated fauna. The assemblages include small blades and points produced via Levallois-like methods, alongside a modern human child's tooth, indicating an early incursion of Homo sapiens into Neanderthal territories before their replacement around 45,000 years ago. This site underscores episodic modern human presence in during the IUP phase. In , the Kara-Bom site in the of southern stands as a pivotal locus for understanding IUP dispersal into northern territories. Excavations reveal multilayered deposits with IUP layers dated to approximately 47,000–43,000 cal BP through new radiocarbon analyses of bone and charcoal samples. These layers contain characteristic bifacial foliate points, prismatic blade cores, and Levallois blanks, indicating a technological package akin to early IUP but adapted for local raw materials like and . The site's faunal remains, including and , underscore its role in tracking human movement across Siberian steppes, positioning Kara-Bom as one of the earliest confirmed IUP occupations beyond the . Further west in , the Peștera Muierii cave in yields evidence of presence associated with transitional lithic industries. Human remains, including a female cranium and postcranial elements, are directly dated to approximately 35,000 cal via radiocarbon on bone collagen, placing them within the broader early timeframe that encompasses IUP influences. Accompanying tools feature bladelets, endscrapers, and burins on local flint, suggesting an assemblage blending and elements consistent with initial modern human incursions into southeastern . This site illustrates the penetration of IUP-like technologies into the Carpathian region, with isotopic analysis of the remains indicating a diet reliant on large herbivores suited to forested uplands. In , Kent's Cavern in , , offers debated but significant signals of early IUP activity. A fragmentary (Kent's Cavern 4) from the cave's deposits has been dated to 44,000–41,000 cal using radiocarbon pretreatment on adhering bone, though stratigraphic mixing with earlier layers has sparked controversy over its contextual integrity. Associated lithics include blades and flakes potentially attributable to proto-Aurignacian or IUP traditions, recovered alongside fauna like and , pointing to intermittent occupation during the onset of modern human expansion into . The site's position on the northwestern fringe of the continent underscores the challenges and timelines of IUP signals reaching isolated refugia amid fluctuating glacial climates. Extending back into Central Asia, the Obi-Rakhmat Grotto in Uzbekistan preserves multilayered sequences bridging Middle and Upper Paleolithic phases. Stratum 5, interpreted as transitional to IUP, is dated to circa 45,000 cal BP based on radiocarbon dates from hearth charcoal and stratified sediments, revealing assemblages that integrate Levallois point production with elongated blades and backed tools. These artifacts, crafted from chert and quartz, show technological affinities to both Levantine IUP and local Middle Paleolithic variants, including possible projectile points indicative of hunting strategies. The site's location along trade and migration corridors in the Tian Shan foothills highlights Central Asia's role as a conduit for IUP dissemination eastward and northward. Across these European and Central Asian sites, IUP populations demonstrated remarkable adaptability to cold steppe-tundra and forest ecosystems, as evidenced by faunal profiles dominated by such as , , and . Stable isotope data from human and animal remains at sites like Peștera Muierii confirm reliance on grasses-fueled herbivores, reflecting specialized hunting of large game in open landscapes during Marine Isotope Stage 3. This environmental niche exploitation facilitated sustained occupation in conditions, with toolkits emphasizing efficient blade production for composite weapons and processing hides for cold-weather survival.

Sites in East Asia and Oceania

The in , , represents a key locus for Initial Upper (IUP) occupation in , with Locality 1 (SDG1) yielding blade-based assemblages dated to approximately 46–41 ka cal BP through optically stimulated (OSL) and radiocarbon methods. These small tool kits feature recurrent Levallois-like reduction methods for producing blades and elongated flakes, alongside early microblade components and volumetric cores, indicating technological continuity with western IUP traditions but adapted to local siliceous resources. Such findings challenge Eurocentric narratives of innovation by demonstrating that IUP technologies dispersed rapidly across , reaching via southern or eastern routes rather than solely northern continental pathways. Further east, Tianyuan Cave near provides direct evidence of presence associated with IUP contexts, where a partial skeleton (Tianyuan 1) dated to 42–39 ka cal BP via radiocarbon was recovered from Layer III. This burial, lacking associated stone artifacts but embedded in a faunal-rich deposit, exhibits modern human with some archaic traits, linking it to foundational East Asian populations that diverged during the . The site's temporal placement aligns with the IUP transition, underscoring northern China's role in early dispersals of anatomically modern humans into eastern . On the , the Nwya Devu site at 4,600 meters above stands as the highest-altitude IUP locality worldwide, occupied intermittently from 40–30 ka BP based on OSL dating of quartz grains. Artifacts include a blade-dominated lithic assemblage with prismatic cores and limited retouched tools, alongside implements, reflecting adaptations to high-elevation environments amid cold-adapted such as blue sheep and . This discovery extends IUP evidence into extreme altitudes, suggesting human technological and physiological resilience facilitated southern highland expansions during Marine Isotope Stage 3. In Oceanic extensions, exhibits potential early IUP signals through archaeological traces of modern human activity dated to around 47–43 cal BP at sites like on , including behavioral shifts toward symbolic practices and adaptation to island ecosystems. These indicate maritime crossings into the region by IUP populations, though the core IUP phase (50–40 BP) precedes the full colonization of around 50 BP, highlighting as a transitional corridor for southern dispersals. Recent ancient genomic analyses reinforce these archaeological patterns, with Bennett et al. (2024) modeling East Asian ancestry divergence around 46 ka BP via a southern route, supported by data from individuals like Tianyuan 1 showing basal affinities to both northern and coastal lineages. This framework posits sequential population splits along southern Eurasian pathways, integrating genetic evidence with site distributions to emphasize alternative migration dynamics beyond northern routes.

Technological Characteristics

Lithic Industries

The lithic industries of the Initial (IUP) are defined by a transitional technological repertoire that bridges Middle and Upper Paleolithic traditions, emphasizing efficient flake and blade production for versatile tool forms. Core reduction strategies retain elements of the Levallois method, particularly for producing pointed flakes and blanks suitable for points, while incorporating prismatic blade knapping to yield elongated, parallel-sided blanks. This combination is evident in Levallois-like cores with preferential flake removal for pointed tools alongside unidirectional prismatic cores for blades, reflecting a deliberate integration of established and novel reduction sequences. Key tool types include regionally distinct forms such as Emireh points in the , characterized by butt-thinning and marginal retouch on Levallois points; backed bladelets in , which are finely retouched, elongated microliths often under 3 cm long; and bifacial leaf points in , biconvex tools with invasive retouch on both faces. Microliths, including backed bladelets and geometric segments, typically measure 2–5 cm in length, enabling precise applications in hunting and processing. Raw materials were predominantly high-quality flint and chert, sourced from local nodules or outcrops within radii of 10–20 km, with evidence of in some assemblages to improve flaking predictability, as seen at Kara-Bom where silicites show thermal alteration traces. Technological variability manifests regionally, with Near Eastern assemblages dominated by from prismatic and Levallois cores, contrasting with more flake-oriented strategies in where core-and-flake reduction prevails alongside sporadic technologies at key sites. In and , hybrid approaches combine Levallois flakes with burin-core bladelet , highlighting adaptive responses to local availability and mobility patterns. Burin-core reduction in these regions involves complex sequences of 13–21 procedural units, yielding small blades or bladelets through spalling, comparable to systems. A major innovation was the shift toward standardized bladelets produced via intentional blade fragmentation and burin spalling, facilitating the creation of composite tools such as hafted projectiles through backing and segmentation for secure attachment to shafts or handles. This standardization enhanced tool efficiency and modularity, supporting increased specialization during IUP dispersals.

Tool Assemblages and Innovations

Tool assemblages in the Initial Upper Paleolithic (IUP) typically consist primarily of lithic artifacts, supplemented by and implements such as awls and points, along with rare wooden artifacts. These non-lithic elements represent a minor but significant portion of the toolkit, enabling diverse functions beyond stone-based production. At sites like Kamenka in , dated to around 45,000 years ago, tools and engraved bones appear alongside lithics, highlighting the integration of organic materials in early IUP adaptations. Functional categories within IUP assemblages emphasize practical subsistence needs, including hunting tools like projectile points (e.g., Emireh points with bifacial thinning for secure attachment), processing implements such as scrapers and burins for hide and wood working, and domestic perforators like awls for leatherworking. Evidence of technologies underscores the development of composite tools that enhanced efficiency in these activities. This is evident in sites like Emireh Cave, where points were designed for mounting on shafts, marking a key innovation in weapon and tool design. Innovations in osseous materials further distinguish IUP toolkits, with early examples of bone and antler worked into needles and awls, indicating adaptations for sewing clothing and possibly fishing with harpoon-like implements. At in , formal bone tools including eyed needles and awls date to the initial Upper Paleolithic around 50,000 BP, suggesting tailored responses to colder climates through improved garment production. These developments reflect a broadening technological repertoire that supported mobile lifestyles. Subsistence strategies are illuminated by faunal remains across IUP sites, revealing a focus on of large herbivores such as (Rangifer tarandus), (Equus sp.), (Bison/Bos sp.), and caprines, complemented by gathering of plant resources but without any evidence of . At in (ca. 45,000 years ago), zooarchaeological analysis of over 7,000 remains shows selective exploitation of medium-to-large ungulates, with cut marks indicating human butchery and transport of high-utility body parts to the site. This pattern of intensive large-mammal hunting persisted as a core economic strategy, with limited reliance on small game. Regional variations include adaptations to harsh environments, such as hybrid core reduction in Siberia for small, sharp blades suitable for insertion into handles, facilitating precise cutting and cold-weather tool maintenance.

Cultural and Symbolic Aspects

Artistic Expressions

The Initial Upper Paleolithic (IUP) marks the appearance of some of the earliest documented artistic expressions associated with anatomically modern humans in Eurasia, including the processing and use of pigments, personal adornments, and simple engravings. These artifacts suggest an emerging capacity for symbolic communication and aesthetic production, though they remain relatively simple compared to later Upper Paleolithic developments. Evidence from key sites indicates that such expressions were integral to IUP cultural practices, potentially linked to social identity and ritual activities. One of the hallmark features of IUP artistic behavior is the processing of red , an , dated to approximately 45,000 years (BP) in the . At sites like Ksâr 'Akil in , red ochre fragments occur in Initial Upper Paleolithic layers, showing signs of grinding and mixing, likely for application as body decoration or in ritual contexts. This practice represents a continuity from traditions but increases in frequency and complexity during the IUP, possibly facilitating social signaling among dispersing human groups. Similar ochre processing is evident at other IUP sites, underscoring its role in early symbolic . Personal ornamentation also emerges prominently in IUP assemblages, with pierced marine shells and animal teeth serving as beads for body adornment around 42,000–41,000 BP. At Üçağızlı Cave in southern , excavations have uncovered over 100 such items, including gibbosulus shells with drilled perforations and fox canines with notches, indicating deliberate manufacture for necklaces or other jewelry. These ornaments, often sourced from coastal environments up to 200 km away, imply trade networks and a concern for personal or group identity, distinguishing IUP populations from preceding groups where such items are rare. The use of red ochre in with these beads further suggests multifunctional roles. Portable art in the form of abstract engravings on appears in IUP contexts, providing evidence of non-figurative artistic experimentation. At in , dated to about 46,000–44,000 , bone tools and ornaments include incised lines and geometric patterns on and remains, interpreted as decorative or functional modifications. Cave art, while rarer, is represented by IUP-adjacent examples such as the incised animal figures and abstract lines at Peștera Coliboaia in , dated to around 35,000–30,000 , though these align more closely with the subsequent phase. These engravings, executed with stone tools, reflect early abstract thinking without the narrative complexity seen in later periods. The chronological emergence of these artistic expressions postdates 45,000 , coinciding with the expansion of IUP technologies across and a marked increase in symbolic artifacts compared to the sparse and utilitarian record. This shift is attributed to behavioral innovations among modern humans, enabling more diverse forms of expression. Interpretations view IUP art as foundational evidence of abstract , fostering social cohesion, though it lacks the elaborate figurative styles of later Venus figurines.

Evidence of Symbolic Behavior

The partial skeleton of an , dated to approximately 40,000 years (), was recovered from Layer III of Tianyuan Cave in northern . The remains show no signs of carnivore damage but are disarticulated and dispersed over a small area, consistent with post-depositional disturbance by natural processes rather than intentional burial. Although no definitive were identified, the site's association with processing and marine shell fragments in contemporary East Asian contexts points to emerging ritualistic elements in human mortuary treatment during this period. Recent findings from Manot Cave in the reveal evidence of deep cave use for collective s around 46,000–42,000 calibrated years , including structured deposits of artifacts and possible activities, suggesting organized and behaviors among IUP populations. in IUP sites further demonstrates organization through structured living spaces that reflect planning and investment. At Kara-Bom in the Siberian , dated to around 43,000–41,000 , excavations revealed multiple hearths arranged in linear patterns and storage pits for provisions, indicating deliberate campsite design for prolonged occupation and resource management beyond immediate survival needs. These features suggest investment in communal spaces, where fire and storage served not only practical functions but also roles in group . Long-distance networks, a hallmark of symbolic and , are attested by the presence of marine shells at inland IUP sites, implying or systems that facilitated cultural . In the , sites like Üçağızlı Cave and Ksar 'Akil, dated to approximately 45,000 BP, yielded perforated shells sourced from Mediterranean coasts, transported over 200 km inland, which were used as beads and likely held symbolic value in personal adornment or rituals. This pattern underscores the of inter-group connections, essential for sharing ideas and materials across landscapes. The cognitive underpinnings of IUP symbolic behavior are debated, particularly regarding the onset of and potential for . Tool assemblages, characterized by Levallois-like blade production and composite technologies, exhibit complexity that implies abstract planning and possibly linguistic transmission of knowledge, though direct evidence for remains elusive. Scholars argue that these innovations mark the gradual emergence of modern cognition around 50,000–40,000 BP, challenging earlier models of a sudden " revolution" and emphasizing continuity from traditions. Social organization in the IUP is inferred from site densities and faunal remains, supporting bands of 20–50 individuals engaged in communal hunting. Analysis of base camps like those in the and shows clustered activity areas and large game processing (e.g., and ), consistent with group sizes optimal for coordinated and risk-sharing in societies, reflecting symbolic norms of reciprocity and alliance formation.

Human Populations and Archaeogenetics

Fossil Evidence

The Initial Upper Paleolithic (IUP) is characterized by sparse but significant fossil evidence of early anatomically modern humans (Homo sapiens), primarily consisting of fragmentary skeletal remains associated with transitional lithic industries dated between approximately 50,000 and 40,000 years (BP). These fossils, often recovered from cave contexts in , provide the earliest direct indications of modern human presence outside during this period, with key specimens including isolated bones and partial skeletons that exhibit a mix of derived modern traits and occasional features. Prominent examples include the Ust'-Ishim from , a proximal left femur dated to around 45,000 through direct , representing one of the oldest known modern human remains in northern . This specimen, belonging to an individual, displays anatomical features consistent with early modern humans, such as a relatively gracile build, though it lacks associated archaeological material, complicating precise cultural attribution to the IUP. In , the Oase Cave in yielded two key fossils: Oase 1, a robust from an dated to approximately 40,500 , and Oase 2, a nearly complete cranium from an individual estimated at 17-18 years old, also around 40,000 . These remains, found in a karstic system with hydraulic transport evidence, show modern human including a prominent chin and reduced brow ridges, but with some robust traits possibly influenced by regional variation. Further south in the , in provided four H. sapiens fragments—a tooth and three bones—directly dated to 45,930-42,580 , stratigraphically linked to IUP layers containing Levallois-like tools and personal ornaments. In , the Tianyuan Cave near , , preserved a partial (Tianyuan 1) of an , radiocarbon-dated to about 40,000 calendar years , including cranial and postcranial elements associated with early blade technologies suggestive of IUP affinities. Anatomically, IUP fossils generally feature high, vertical foreheads, rounded cranial vaults, and gracile that distinguish them from contemporaneous Neanderthals, whose skulls were more elongated with prominent supraorbital tori and robust postcrania. For instance, the Oase 2 cranium exhibits a globular braincase and orthognathic face, while the Tianyuan shows slender long bones and dental indicating a varied through moderate wear patterns on molars and premolars. These traits reflect an overall reduction in robusticity compared to , adapted potentially to diverse ecological niches during rapid dispersals. Demographic insights from these remains suggest representation of both sexes, as evidenced by the male mandibles at Oase 1 and Tianyuan, alongside the likely juvenile or female-associated Oase 2 cranium, with no consistent patterns of perimortem indicating interpersonal in the preserved sample. Preservation of IUP fossils poses significant challenges due to taphonomic processes, including fragmentation from post-depositional movement in caves and poor organic survival in humid or acidic sediments, resulting in mostly isolated elements rather than complete skeletons. Recent advances in () extraction from petrous bones and teeth, however, have enabled successful recovery from specimens like those at Bacho Kiro and Oase, despite these degradative factors, offering new windows into without relying on genomic interpretations.

Genetic Studies and Ancestry

Genetic studies of Initial Upper Paleolithic (IUP) populations rely on () extraction and sequencing using next-generation technologies, such as Illumina platforms, which allow for the recovery of low-coverage genomes from skeletal remains despite degradation. These methods involve or targeted capture of nuclear and , followed by authentication to detect postmortem damage signatures like cytosine deamination. However, challenges persist, including potential contamination from modern human sources—estimated and mitigated using tools like AuthentiCT—and limited sample sizes, with only a handful of IUP individuals yielding sufficient endogenous DNA for population-level inferences. Ancestry analyses indicate that IUP populations in included a component, a lineage that diverged from other non-African groups prior to 50,000 years ago, likely in the or region, representing an early branch with reduced admixture. In , the ~40,000-year-old Tianyuan individual from exhibits close genetic relatedness to the ancestors of present-day East Asians and , forming a basal branch to modern East Asian diversity via a southern coastal dispersal route. This supports models of multiple out-of-Africa migrations, with East Asian IUP groups showing affinities to later regional populations through shared derived alleles. Neanderthal admixture in IUP individuals ranges from 2% to 9%, exceeding the ~2% average in modern non-Africans due to more recent interbreeding events dated to 50,000–60,000 years ago, shortly after out-of-Africa dispersals. For instance, the Oase 1 individual from (~40,000 years old) carries 6–9% Neanderthal ancestry in long, uninterrupted segments, indicating a Neanderthal ancestor within the previous four to six generations, though this signal did not persist substantially into later populations. Other IUP genomes, such as those from , show elevated Neanderthal (~3-3.8%) with evidence of recent , highlighting variable admixture pulses with limited long-term to contemporary humans. Migration patterns are illuminated by uniparental markers, including Y-chromosome —the root of non-African paternal lineages—and haplogroups M and N, which trace back to out-of-Africa waves around 60,000–70,000 years ago and diversified during IUP expansions. Bennett et al. (2024) provide evidence for a southern route into , with ancient genomes from the region linking early IUP arrivals to the formation of East Eurasian ancestry clusters. Population dynamics during IUP reveal bottlenecks associated with rapid expansions into , reducing effective population sizes and leading to high . In , IUP groups like those at Bacho Kiro contributed less than 1% to subsequent ancestries, largely replaced by later waves, whereas in , lineages akin to Tianyuan persist as a foundational component in modern populations, comprising a substantial portion of regional .

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