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Paleolithic

The Paleolithic period, also known as the Old Stone Age, represents the earliest and longest phase of human prehistory, extending from approximately 3.3 million years ago, when the earliest stone tools were crafted at Lomekwi 3 in Africa,^[1] to around 10,000 BCE, marking the transition to more settled lifestyles with the advent of agriculture.^[2]^[3] During this era, early hominins such as Australopithecus, Paranthropus, and various Homo species, including Homo habilis, Homo erectus, Neanderthals (Homo neanderthalensis), and eventually anatomically modern Homo sapiens, lived as nomadic hunter-gatherers, relying on foraging wild plants and hunting or scavenging animals for sustenance without domesticating crops or animals.^[2]^[4] The Paleolithic is conventionally divided into three main stages based on technological and cultural advancements: the Lower Paleolithic (roughly 2.6 million to 300,000 years ago), characterized by simple stone tools like choppers and hand axes made through basic flaking techniques; the Middle Paleolithic (about 300,000 to 50,000 years ago), featuring more refined flake tools, prepared cores, and evidence of complex behaviors such as controlled use of fire and possible symbolic practices among Neanderthals and early Homo sapiens; and the Upper Paleolithic (around 50,000 to 10,000 BCE), which saw sophisticated innovations including blade tools, bone and antler implements, needles for clothing, and the emergence of symbolic expression through cave art, figurines, and personal ornaments, primarily associated with modern humans.^[3]^[2] These stages reflect gradual migrations out of Africa starting around 2 million years ago, with human populations adapting to diverse environments across Eurasia, including cold glacial periods that influenced tool use and social organization.^[3] Key notable aspects include the period's role in human evolution, where brain size increased dramatically alongside tool complexity, enabling survival in varied climates and the control of fire by at least 1 million years ago for cooking and warmth.^[4] Socially, Paleolithic groups lived in small, mobile bands of 20–50 individuals, with evidence of cooperation in hunting megafauna like mammoths and the development of language-like communication in the later stages.^[3] Artifacts from this era, numbering in the thousands and spanning over 50 countries, provide insights into cognition, environmental adaptation, and cultural diffusion, underscoring the Paleolithic as a foundational epoch that shaped the trajectory of human innovation before the Neolithic Revolution.^[2]

Introduction and Terminology

Definition and Chronology

The Paleolithic, also known as the Old Stone Age, represents the earliest and longest phase of human prehistory, characterized by the initial development and use of stone tools by hominins. It spans from the appearance of the oldest known stone tools, dated to approximately 3.3 million years ago at Lomekwi 3 in Kenya, to the end of the Pleistocene epoch around 11,650 years before present (BP), marking the transition to the Mesolithic or Neolithic periods in most regions.^[5]^[4] This period encompasses the gradual evolution of tool-making technologies and adaptive behaviors in response to environmental changes, without evidence of agriculture or permanent settlements.^[6] The Paleolithic is traditionally subdivided into three main chronological phases based on technological and cultural advancements. The Lower Paleolithic, from about 3.3 million to 300,000 years ago, includes the earliest tool industries such as the Lomekwian and Oldowan, followed by the Acheulean, reflecting basic flaking and bifacial handaxe production.^[5]^[4] The Middle Paleolithic, spanning roughly 300,000 to 50,000 years ago, is associated with more refined prepared-core techniques like the Levallois method and the Mousterian industry, primarily linked to Neanderthals and early modern humans in Eurasia and Africa.^[7] The Upper Paleolithic, from approximately 50,000 to 11,650 years ago, features advanced blade technologies, symbolic artifacts, and the emergence of behavioral modernity, coinciding with the global dispersal of anatomically modern humans.^[8] These divisions, while Eurocentric in origin, are applied globally with adjustments for regional archaeological sequences. The end of the Paleolithic is defined by the onset of the Holocene epoch and the advent of sedentism, plant domestication, and animal husbandry, though exact dates vary regionally due to differing environmental and cultural trajectories. In Eurasia and Africa, it generally concludes around 10,000 BCE with the warming climate facilitating Neolithic innovations. In the Americas, where human presence is more recent, the equivalent Paleoindian period extends later, with cultures like Clovis dating to 13,050–12,750 calibrated years BP, and some foraging traditions persisting until about 8,000 BCE before transitioning to Archaic periods.^[9] Recent archaeological findings have refined the early chronology; for instance, 2023 excavations at Nyayanga, Kenya, uncovered Oldowan tools dated to 3.03–2.58 million years ago, expanding the geographic range of early lithic technologies beyond East Africa.^[10] Additionally, 2025 analyses at the same site revealed evidence of raw material transport over 10 kilometers, indicating planned foraging behaviors by 2.6 million years ago, while reaffirming the Lomekwi origins at 3.3 million years.^[11]

Etymology and Periodization

The term "Paleolithic" was coined by British archaeologist and naturalist John Lubbock in 1865, in his seminal work Pre-historic Times, as Illustrated by Ancient Remains and the Manners and Customs of Modern Savages, deriving from the Greek words palaios ("old") and lithos ("stone") to designate the "Old Stone Age."^[12]^[13] This nomenclature distinguished it from the "New Stone Age" or Neolithic, which Lubbock contrasted with polished stone tools and early agriculture.^[14] Lubbock's framework built upon the three-age system earlier proposed by Danish antiquarian Christian Jürgensen Thomsen in 1836, which categorized prehistory into Stone, Bronze, and Iron Ages based on predominant materials in artifacts at the National Museum of Denmark.^[15]^[16] The periodization of the Paleolithic evolved significantly from its 19th-century origins, initially relying on stratigraphic sequences and typological classifications of stone tools from European sites. Lubbock's broad division was refined by French archaeologist Gabriel de Mortillet in the 1880s, who subdivided the Paleolithic into Lower, Middle, and Upper phases based on tool morphologies observed in French caves, such as the Chellean (early handaxes) and Aurignacian (blades).^[17] By the mid-20th century, absolute dating revolutionized this system; potassium-argon (K-Ar) dating, introduced in the 1960s, provided chronologies for volcanic layers enclosing early tools, while uranium-series (U-series) dating applied to carbonate deposits in caves enabled precise timelines for later assemblages.^[18]^[19] These methods revealed chronological overlaps and regional disparities, sparking debates on synchronizing African and Eurasian timelines—for example, whether the African Middle Stone Age aligns temporally with the Eurasian Middle Paleolithic or reflects independent trajectories influenced by local ecologies.^[20]^[21] Paleolithic subdivisions are fundamentally rationalized by shifts in lithic technology and associated behaviors, rather than uniform temporal markers, allowing for flexible application across continents. The Lower Paleolithic emphasizes rudimentary flaking, as in the simple pebble tools of the Oldowan industry (~2.6 million to 1.7 million years ago), progressing to the more refined bifacial handaxes of the Acheulean.^[1] The Middle Paleolithic marks a leap in premeditation with prepared-core methods, exemplified by the Levallois technique in the Mousterian toolkit, enabling efficient flake production for diverse implements.^[22] The Upper Paleolithic (~50,000 to 12,000 years ago) showcases heightened sophistication, including thin blades, burins for engraving, and composite tools, often linked to symbolic expressions like art.^[23] In Africa, the contemporaneous Middle Stone Age parallels these Eurasian developments through innovations like heat-treated silcrete blades but incorporates regionally specific adaptations, such as backed tools, underscoring that periodization prioritizes techno-cultural transitions over rigid chronology.^[24]^[25] Criticisms of early 20th-century models highlight their Eurocentric bias, which imposed French sequences globally and undervalued African evidence; post-2020 refinements, incorporating genomic data from ancient DNA, have integrated behavioral modernity—evidenced by complex planning and symbolism—back to around 300,000 years ago in Africa, associated with early Homo sapiens at sites like Jebel Irhoud, rather than confining it to the Eurasian Upper Paleolithic.^[26]^[27] This genetic-archaeological synthesis challenges outdated linear progressions, emphasizing mosaic evolution across regions.^[28]

Geological and Environmental Setting

Paleogeography

During the Pleistocene epoch, which encompasses the Paleolithic period from approximately 2.58 million to 11,700 years ago, Earth's continental configurations remained largely stable compared to earlier geological eras, with the supercontinent Pangaea long fragmented into the modern continents. Tectonic activity was minimal, featuring gradual shifts such as the continued separation of Africa from Eurasia and minor subduction along plate boundaries, but these did not drastically alter landmasses relevant to human dispersal. Africa served as the primary cradle for early hominins, characterized by diverse habitats including expansive savannas and woodlands that supported evolutionary adaptations.^[29] Eurasia exhibited a mosaic of environments, from steppes and tundras in the north to forests and grasslands in the south, while later expansions reached Australia around 65,000 years ago and the Americas, with habitats ranging from arid interiors to coastal zones.^[30] Significant paleogeographic features emerged due to glacial-interglacial cycles, notably land bridges exposed by lowered sea levels. The Beringia land bridge, connecting Siberia to Alaska, was intermittently exposed during glacial maxima, facilitating human migration into the Americas between approximately 20,000 and 13,000 years ago as a corridor of tundra and steppe environments.^[31] In Southeast Asia, Sundaland formed a vast low-lying landmass linking modern Indonesia, Malaysia, and surrounding islands, encompassing rainforests and riverine habitats that enabled early human movements.^[32] Similarly, Doggerland in northern Europe connected Britain to the mainland, featuring wetlands, rivers, and coastal plains that supported Mesolithic populations before inundation.^[33] Sea level fluctuations, driven by the accumulation and melting of continental ice sheets, profoundly shaped these configurations. At the Last Glacial Maximum (approximately 26,500–19,000 years ago), global sea levels dropped by about 120 meters, exposing extensive coastal shelves and creating migration routes now submerged.^[34]^[35] Recent 2025 archaeological evidence from lithic tool analyses, including microblade technologies, supports a Pacific coastal migration route to North America, with artifacts linking East Asian traditions to submerged Pacific Rim pathways used over 20,000 years ago.^[36]^[37]

Climate Variations and Impacts

The Pleistocene epoch, encompassing the Paleolithic period from about 2.6 million to 11,700 years ago, featured pronounced glacial-interglacial cycles driven by Milankovitch orbital variations and amplified by feedback mechanisms such as ice-albedo effects. In the European Alps, these cycles are traditionally divided into four major glaciations—Günz (approximately 2.6–0.65 million years ago), Mindel (0.65–0.3 million years ago), Riss (0.3–0.13 million years ago), and Würm (0.13 million to 11,700 years ago)—interspersed with warmer interglacial intervals that facilitated biotic recoveries and expansions.^[38] Correlative stages in North America include the Nebraskan, Kansan, Illinoian, and Wisconsinan glaciations, with the latter peaking around 20,000 years ago during the Last Glacial Maximum (LGM).^[39] Interglacials, such as the Eemian (about 130,000–115,000 years ago), brought global warming of several degrees, enabling forest regrowth and faunal dispersals across previously glaciated landscapes.^[40] During glacial maxima, global temperatures declined by up to 10°C in mid-latitudes relative to interglacials, with sea-surface temperatures dropping similarly in the North Atlantic and precipitation shifting toward drier conditions in many continental interiors.^[41] These shifts intensified aridity in tropical regions and expanded polar fronts southward, culminating in widespread megafauna extinctions; for instance, woolly mammoths vanished around 10,000 years ago amid the terminal Pleistocene warming transition from the LGM.^[42] Superimposed on these long-term cycles were abrupt millennial-scale fluctuations, including Dansgaard-Oeschger (D-O) events—rapid Greenland warmings of 8–15°C lasting centuries, occurring roughly 25 times between 115,000 and 11,700 years ago—and Heinrich events, involving iceberg armadas that cooled the North Atlantic by 2–5°C for millennia through freshwater influx disrupting ocean circulation.^[43]^[44] Ecologically, glacial advances transformed Africa's equatorial belt from humid tropical forests to expansive arid savannas and grasslands during episodes like 135,000–90,000 years ago, reducing vegetation cover and altering herbivore distributions.^[45] In Eurasia, cooling promoted tundra and steppe biomes, with tundra expanding southward to cover much of northern Europe and Asia between 21,000 and 14,000 years ago, supporting cold-adapted megafauna while constraining woodland habitats.^[46] Volcanic perturbations, such as the Toba supereruption in Indonesia around 74,000 years ago—a VEI 8 event ejecting over 2,800 km³ of material—induced a multi-year volcanic winter with global temperature drops of 3–5°C and reduced photosynthesis, yet archaeological and genetic evidence reveals human populations exhibited resilience without a near-total bottleneck.^[47] Recent 2023 analyses of bipolar ice cores from Greenland and Antarctica refine Toba's timing to approximately 73,700 years ago and indicate only modest, short-lived radiative forcing, underscoring early modern humans' adaptive capacity amid such stressors rather than extinction-level collapse.^[48]

Human Evolution and Populations

Early Hominins and Tool Makers

The earliest hominins associated with the onset of the Paleolithic include species predating the genus Homo, such as Australopithecus afarensis, which is exemplified by the partial skeleton known as "Lucy," discovered in 1974 at Hadar, Ethiopia, and dated to approximately 3.2 million years ago.^[49] This species exhibits key adaptations like bipedalism, evidenced by skeletal features such as a curved phalanges indicating partial arboreality alongside upright walking, which likely facilitated the carrying of tools or food over distances.^[50] A. afarensis had a small brain size, estimated at 400–500 cubic centimeters, comparable to that of modern chimpanzees, suggesting limited cognitive capacity for complex behaviors despite evidence of potential tool use inferred from associated cut marks on animal bones.^[51] Other pre-Homo species potentially linked to early tool making include Australopithecus garhi, known from fossils dated to about 2.5 million years ago in the Afar region of Ethiopia, where cut-marked bones and primitive stone tools were found in proximity, indicating possible involvement in scavenging and meat processing. Similarly, Kenyanthropus platyops, discovered in 1998 at Lomekwi, Kenya, and dated to around 3.5 million years ago, features a flat face, small molars, and a brain size of approximately 400 cubic centimeters, with its location near early tool sites suggesting it as a candidate for rudimentary tool use among diverse hominin lineages. These species highlight a mosaic of traits, including bipedal locomotion that freed the hands for manipulation, though their small cranial capacities imply that tool behaviors were opportunistic rather than systematically innovative.^[52] The origins of Oldowan technology, the simplest known stone tool industry, are traced to approximately 2.6 million years ago at sites in Gona, Ethiopia, where flakes were produced by striking cores to create choppers and scrapers primarily for processing animal carcasses through opportunistic scavenging rather than active hunting.^[53] Anatomical evidence from these early tool makers includes bipedalism, which enabled efficient transport of unmodified stones or simple implements across savanna landscapes, as supported by locomotor reconstructions from fossil pelvises and femora.^[54] Additionally, analysis of cut marks on bones from Oldowan contexts reveals a bias toward oblique incisions consistent with right-handed tool wielding, providing the earliest indications of population-level manual laterality among hominins.^[55] Recent discoveries have challenged the traditional attribution of tool use exclusively to Homo, with 2023 findings at Nyayanga, Kenya, uncovering 2.9-million-year-old stone tools used for butchering hippopotamus remains and pounding plant materials, expanding the geographic and temporal scope of pre-Homo lithic activities.^[10] Similarly, at Dikika, Ethiopia, bones dated to 3.3 million years ago bear cut marks interpreted as evidence of stone-tool-assisted defleshing, potentially linked to A. afarensis or related taxa, though some debate persists over whether these marks result from tools or trampling. A 2025 hypothesis posits that early lithic technologies may have been influenced by behavioral patterns observed in non-human primates, such as percussive nut-cracking in chimpanzees and capuchins, which produce sharp flakes as by-products and suggest convergent evolutionary pathways in tool modification without cumulative cultural transmission.^[56]

Key Homo Species

Homo habilis Homo habilis, often translated as "handy man," represents one of the earliest members of the genus Homo, emerging around 2.4 million years ago and persisting until approximately 1.4 million years ago in East Africa.^[57] This species is characterized by a brain size averaging about 600 cubic centimeters, slightly larger than that of preceding australopiths, along with reduced facial prognathism and smaller teeth adapted for a more varied diet.^[57] Fossils, including notable specimens from Olduvai Gorge in Tanzania such as OH 7 and OH 24, reveal a body size similar to modern humans but with more primitive limb proportions.^[58] H. habilis is closely associated with the Oldowan tool industry, consisting of simple stone flakes and choppers used for processing food, marking an early advancement in cultural technology during the Paleolithic.^[57] Homo erectus Homo erectus, appearing around 1.9 million years ago and enduring until about 110,000 years ago, was the first hominin to exhibit modern body proportions, including longer legs suited for endurance walking and running.^[59] Brain size in this species ranged from approximately 900 to 1,200 cubic centimeters, showing a significant increase over earlier Homo and enabling enhanced cognitive capabilities.^[59] As the first global disperser, H. erectus fossils are found across Africa, Asia, and Europe, including the Dmanisi site in Georgia dated to 1.8 million years ago and the Trinil site in Java, Indonesia, where Eugene Dubois discovered "Java Man" in 1891.^[59] This species is linked to the Acheulean tool tradition, featuring symmetrical bifacial hand axes that demonstrate improved planning and skill in lithic technology.^[60] Homo heidelbergensis Homo heidelbergensis, dating from about 700,000 to 200,000 years ago, served as a transitional or "bridge" species between earlier Homo erectus and later forms like Neanderthals and anatomically modern humans.^[61] With a brain size averaging around 1,200 cubic centimeters, this species displayed a more rounded cranium, reduced brow ridges compared to H. erectus, and robust skeletal features indicative of high physical activity.^[61] Key evidence includes wooden spears from the Schöningen site in Germany, dated to approximately 300,000 years ago, which represent the earliest known complex hunting weapons and suggest advanced predatory strategies.^[62] Fossils from sites like Broken Hill in Zambia and the type specimen from Mauer, Germany, highlight its wide distribution across Africa and Eurasia, underscoring its role as a precursor to regional human variants.^[63] Homo neanderthalensis Homo neanderthalensis, commonly known as Neanderthals, thrived from roughly 400,000 to 40,000 years ago, primarily in Europe and western Asia, with adaptations for cold climates including a stocky build, broad noses for warming air, and large nasal cavities.^[64] Their brain size averaged about 1,500 cubic centimeters, larger than that of modern humans, potentially supporting sophisticated problem-solving and social structures.^[64] Neanderthals are renowned for the Mousterian tool industry, which involved prepared-core techniques for producing Levallois flakes, indicating foresight in tool manufacture.^[65] Evidence of intentional burials, such as the child interment at La Ferrassie in France dated to around 70,000 years ago, points to symbolic cognition and possible ritualistic behaviors.^[66] Recent genetic analyses from 2024 have refined the timeline of interbreeding between Neanderthals and early Homo sapiens, confirming gene flow occurred around 50,000 years ago as modern humans dispersed into Eurasia.^[67] In Asia, Denisovans—a sister group to Neanderthals known from limited fossils in Siberia and Tibet—also interbred with Homo sapiens, contributing genetic variants related to high-altitude adaptation in modern populations.^[68]

Dispersal and Anatomical Modern Humans

Anatomically modern humans, or Homo sapiens, first emerged in Africa approximately 300,000 years ago, as evidenced by fossils from Jebel Irhoud in Morocco, which display a mix of modern facial features and archaic braincase morphology. These early H. sapiens possessed a cranial capacity averaging around 1,350 cubic centimeters and a gracile skeletal build, characterized by lighter bones and more rounded skulls compared to earlier hominins.^[69] This anatomical modernity reflects adaptations for enhanced cognitive and social capabilities, marking the onset of our species' global trajectory during the Paleolithic.^[70] The major dispersal of H. sapiens out of Africa occurred between 70,000 and 50,000 years ago, primarily via a northern route through the Levant, allowing populations to enter Eurasia.^[71] From there, migrants reached Australia by around 65,000 years ago, as indicated by archaeological layers at Madjedbebe rock shelter containing stone tools and ochre.^[72] In Europe, H. sapiens arrived approximately 45,000 years ago, represented by Cro-Magnon individuals associated with the Aurignacian culture, overlapping with Neanderthal populations.^[73] The peopling of the Americas involved multiple routes, with pre-Clovis evidence from human footprints at White Sands National Park in New Mexico dating to about 23,000 years ago, predating the traditional Clovis horizon by over 10,000 years.^[74] A 2025 analysis of stone tools from Pacific Rim sites further supports a coastal migration pathway alongside the Beringian land bridge, linking East Asian seafarers to North American entry points during the Last Glacial Maximum.^[37] These migrations were facilitated briefly by geographic features like the exposed Beringia land bridge, enabled by lowered sea levels.^[75] Population dynamics during these dispersals included severe bottlenecks, such as the one around 74,000 years ago potentially linked to the Toba supervolcano eruption, which reduced global human genetic diversity through a near-extinction event affecting early migrants.^[76] Interactions with archaic humans involved both replacement and assimilation; non-African H. sapiens carry 1-2% Neanderthal DNA from interbreeding in Eurasia, while some Asian and Oceanian populations show up to 5% Denisovan admixture from hybridizations.^[77] Recent 2023-2025 research reinforces earlier African modernity, with genetic analyses suggesting structured dispersals within Africa predating 300,000 years ago and ongoing Denisovan contributions in hybrid Asian populations.^[78] These findings highlight a complex history of admixture shaping modern human variation.^[79]

Technological Developments

Lithic Technology and Tools

Lithic technology represents the cornerstone of Paleolithic innovation, encompassing the systematic production of stone tools through knapping techniques that evolved over millions of years. The earliest evidence of intentional stone tool manufacture dates to the Lower Paleolithic, with Oldowan assemblages characterized by simple flakes and choppers produced by striking pebbles or cores with hammerstones, primarily using quartzite and other locally available materials. These tools, found in eastern Africa, exhibit basic flaking patterns that suggest opportunistic reduction of nodules to create sharp edges for cutting and scraping. Recent analyses confirm Oldowan artifacts from localities dating between approximately 2.9 and 2.6 million years ago, pushing back the onset of lithic technology and highlighting early hominin adaptation to resource availability.^[80] The Acheulean industry, emerging around 1.7 million years ago and persisting until about 200,000 years ago, marked a significant advance with the introduction of bifacial handaxes and cleavers shaped through symmetric flaking on both sides, indicating greater planning and standardization in tool production. These large cutting tools, often made from quartzite or basalt sourced from river gravels, featured teardrop or pointed forms with refined edges, facilitating diverse functions such as butchery and woodworking. Assemblages from sites like Konso in Ethiopia demonstrate the gradual refinement of these bifaces, with early examples showing thick profiles and sinusoidal edges that evolved toward more symmetrical designs over time.^[81] In the Middle Paleolithic, the Levallois technique revolutionized lithic production by employing prepared cores to detach predetermined flakes of consistent shape and size, a method that enhanced efficiency and versatility around 300,000 years ago. Associated predominantly with Neanderthals in Europe and the Near East, this core reduction strategy involved hierarchical flaking sequences to create a Levallois flake from a tortoise-like core, often using high-quality flint or chert transported from specific outcrops up to tens of kilometers away. This period also saw the emergence of bone tools alongside lithics, with Neanderthals shaping ribs into specialized lissoirs for hide processing, reflecting integrated organic and stone technologies.^[82]^[83] The Upper Paleolithic witnessed further sophistication in blade technology, where long, thin blades were struck from prismatic cores starting around 45,000 years ago, enabling the production of composite tools with interchangeable parts. These blades, typically crafted from fine-grained flint sourced through long-distance procurement networks, supported the creation of microliths—small, geometrically shaped inserts for hafted projectiles—and burins, chisel-like tools for engraving bone or wood. Microliths, often backed or tanged for secure attachment, appeared widely in late Upper Paleolithic assemblages, optimizing cutting efficiency for hunting armatures. Burins, with their acute-angled working edges formed by deliberate notching, facilitated precise incisions essential for artistic and functional modifications.^[84]^[85] Knapping techniques across the Paleolithic relied on percussion and pressure methods, with raw material selection emphasizing fracture predictability; flint's conchoidal breakage produced sharp, uniform edges, while quartzite's durability suited heavy-duty tasks despite coarser flaking. Sourcing strategies involved targeted exploitation of secondary deposits, as evidenced by geochemical analyses of flint artifacts showing transport distances exceeding 100 kilometers in some regions, underscoring territorial knowledge and exchange. A 2025 hypothesis posits that early lithic innovation may have been inspired by observations of natural rock breakage, such as from trampling or geological processes, bridging the gap between opportunistic use of sharp stones and deliberate manufacture prior to 2.6 million years ago.^[86]^[87]

Control of Fire and Cooking

The earliest evidence for the use of fire by hominins dates to approximately 1.0 million years ago, as indicated by microscopic traces of wood ash, burned bone fragments heated to 400–500°C, and ashed plant remains discovered in Stratum 10 of Wonderwerk Cave in South Africa.^[88] These findings, located 30 meters inside the cave, rule out natural causes like wildfires or spontaneous combustion, suggesting intentional burning events associated with early Acheulean tool makers, likely Homo erectus.^[88] Controlled use of fire, implying repeated and purposeful management, is evidenced around 790,000 years ago at the Gesher Benot Ya'aqov site in Israel, where clusters of burned flint, wood, seeds, and fruits occur across multiple sedimentary layers, consistent with hearth-like features and hominin activity.^[89] This site demonstrates fire's integration into daily behaviors, including possible cooking, as supported by the spatial distribution of heat-altered materials away from natural fire sources.^[89] Fire-starting methods during the Paleolithic are inferred primarily from indirect evidence and experimental archaeology, with percussion techniques—striking flint against pyrite or manganese dioxide to produce sparks—appearing in Middle Paleolithic contexts around 50,000 years ago among Neanderthals, and likely earlier for Homo erectus based on the presence of such minerals at sites.^[90] Friction methods, such as hand drills or bow drills using wood, are hypothesized for the Lower Paleolithic but lack direct artifacts, though ethnographic analogies and residue analysis on tools suggest their feasibility for maintaining hearths.^[91] Fire transport was probably achieved via slow-burning materials like moss or bark in portable hearths, enabling mobility as seen in dispersed burnt sediments at occupation sites.^[91] Archaeological evidence for fire and cooking includes charred animal bones indicating temperatures above 300°C, burnt sediments with ash layers, and heat-fractured lithics, all diagnostic of anthropogenic heating rather than opportunistic scavenging.^[88] More recent analyses, such as those from Gesher Benot Ya'aqov, reveal cooked fish remains dated to 780,000 years ago, with scales and bones showing exposure to low-heat baking around 500–600°C, providing direct proof of plant and animal processing over fire.^[92] Charred plant fragments from African Paleolithic sites (such as Klasies River Caves in South Africa) further confirm cooking of starchy foods, altering microstructures consistent with boiling or roasting by 120,000 years ago.^[93] The control of fire profoundly impacted Paleolithic survival and evolution, offering protection from predators, warmth during glacial periods, and light for extended activities, which expanded habitable ranges into colder environments.^[89] Cooking, by denaturing proteins and breaking down starches, reduced digestion time and energy costs by up to 50%, freeing metabolic resources for larger brains as outlined in the expensive tissue hypothesis, where gut size decreased while encephalization increased in Homo erectus and later species. This dietary shift, enabled by fire, supported cognitive advancements and population growth, with brain sizes doubling over 2 million years partly attributable to more efficient nutrient extraction from cooked foods.

Other Innovations

Paleolithic peoples constructed various forms of shelter to adapt to diverse environments, ranging from natural rock shelters to built structures. Rock shelters, such as those in the Dordogne region of France and the Levant, provided natural protection and often contain archaeological evidence of repeated occupation, including hearths and tool scatters.^[94] In open landscapes, early evidence of constructed shelters appears in the form of post holes—depressions in the ground indicating wooden supports—dating back to around 400,000 years ago at sites like Terra Amata in France, suggesting the use of lightweight frames covered with hides or branches.^[95] More elaborate semi-permanent dwellings emerged during the Upper Paleolithic, exemplified by the mammoth bone huts at Mezhirich in Ukraine, dated to approximately 15,000 years ago, where large bones formed circular walls up to 2 meters high, enclosing oval living spaces of about 20 square meters, possibly roofed with hides and thatch.^[96] These structures, clustered in small settlements, demonstrate organized resource use and seasonal habitation during cold glacial periods.^[97] Clothing technologies developed to enable survival in varied climates, with evidence pointing to the use of animal hides processed into garments. Genetic analysis of human lice indicates that body lice, adapted to clothing, diverged from head lice lineages between 83,000 and 170,000 years ago, suggesting regular use of body coverings by anatomically modern humans in Africa during this period to combat environmental exposure.^[98] The earliest direct artifacts for sewing appear in Eurasia, including eyed bone needles from Denisova Cave in Siberia, dated to around 40,000 calibrated years before present, crafted from bird bone and likely used to stitch hides into fitted clothing for cold steppe conditions.^[99] These needles, measuring about 7 centimeters long with a perforation for thread, represent a key innovation in composite manufacturing, allowing for tailored apparel that enhanced mobility and thermoregulation.^[100] Transport innovations facilitated dispersal across barriers, inferred from colonization patterns requiring watercraft. The presence of Homo floresiensis on Flores Island, Indonesia, with remains dated to about 50,000 years ago, implies seafaring capabilities, as the island lies beyond swimming distance from mainland Asia, separated by deep straits even during glacial lowstands.^[101]^[102] Recent 2025 analysis of stone tools from Pacific Northwest sites in North America reveals technological links to Upper Paleolithic tools from Hokkaido, Japan, supporting a coastal migration route around 20,000 years ago that likely involved watercraft to navigate the Pacific shoreline and reach the Americas.^[37] Such voyages would have required rafts or simple boats constructed from logs and fibers, enabling the transport of people and goods over open water.^[103] Beyond basic shelters and apparel, Paleolithic innovations included composite tools and binding materials that amplified utility. Hafted tools, where stone points were attached to wooden or bone handles, appear as early as 300,000 years ago in Africa and Europe, with residues indicating secure fastening for spears and knives.^[104] Neanderthals produced birch bark tar adhesives as early as around 200,000 years ago, for example at Campitello Quarry in Italy, and later at sites like Königsaue in Germany (~40,000–80,000 years ago), heating bark in oxygen-poor conditions to create a sticky resin for hafting stone tools, demonstrating advanced pyrotechnic knowledge.^[105]^[106] Evidence of cordage from plant fibers, such as bast, dates to at least 41,000 years ago in Europe; a 3-ply twisted cord fragment from Grotte du Renne in France, adhering to a stone tool, was made from pine inner bark and used for binding or netting.^[107] Similar rope-making techniques, involving twisting and plying fibers, are documented in Aurignacian contexts over 41,000 years old, underscoring the role of fiber technology in crafting nets, bags, and structural elements.^[108]

Subsistence Strategies

Diet and Nutrition

The Paleolithic diet was primarily foraging-based, consisting of a diverse array of animal and plant resources that varied by region and season. Animal proteins formed a significant component, particularly from large herbivores such as mammoths, bison, and reindeer, which provided high-quality protein and fats essential for energy demands in cold climates.^[109] Coastal populations often incorporated marine resources like fish and shellfish, contributing omega-3 fatty acids and other nutrients, while inland groups relied more on terrestrial game and gathered plants.^[110] Seasonal fluctuations influenced intake, with greater emphasis on preserved or stored foods during scarcity, and broader dietary breadth emerging in the Upper Paleolithic as evidenced by isotopic signatures indicating mixed carnivory and herbivory.^[111] Nutritionally, this diet supported brain development and overall health through balanced macronutrients and micronutrients. Seafood consumption supplied long-chain omega-3 fatty acids like DHA, which are linked to enhanced cognitive function and neural growth in early humans.^[112] The high diversity of plants, including tubers, nuts, seeds, and wild fruits, ensured adequate fiber, vitamins, and minerals, making deficiencies in essentials like vitamin C or B vitamins rare compared to more monotonous modern diets.^[113] Estimated daily caloric intake ranged from 2,500 to 3,000 kcal, sufficient for active lifestyles, with proteins comprising 20-35% of energy, fats 30-40%, and carbohydrates from plants filling the remainder.^[114] Archaeological and biochemical evidence elucidates this dietary profile. Stable isotope analysis of bone collagen, particularly elevated δ¹⁵N levels, indicates substantial carnivorous intake from megafauna, reflecting trophic positions high in food chains.^[115] Dental microwear textures reveal abrasive patterns from plant processing and meat consumption, supporting a mixed diet rather than meat dominance.^[116] Recent studies, including a 2022 PNAS review of paleobiological data, challenge meat-centric views by providing biogeochemical and dental evidence of significant plant consumption in early hominins, such as C4 grasses and tubers. Analyses of hunter-gatherer subsistence ratios indicate plant foods could contribute 35–55% of energy in ecologically variable settings.^[113]^[117] A 2024 isotopic study from Moroccan sites further confirms high plant reliance (over 50% of diet) via low δ¹³C and δ¹⁵N in remains, highlighting underappreciated botanical contributions.^[118] Early cooking, facilitated by fire control, enhanced nutrient bioavailability from plants and tubers, as shown in dental calculus analyses revealing processed starches.^[119] Health outcomes from this nutrition were generally positive, with Paleolithic skeletons displaying robust bone density and minimal signs of nutritional stress.^[120] Low incidences of chronic conditions like atherosclerosis or diabetes are inferred from the absence of pathological markers in preserved remains, contrasting sharply with post-agricultural increases in such diseases due to dietary shifts.^[121] This nutritional resilience likely stemmed from the diet's anti-inflammatory profile and physical activity synergy, promoting longevity and physical vigor absent in sedentary modern contexts.^[120]

Hunting, Gathering, and Resource Use

During the Lower Paleolithic, early hominins primarily relied on scavenging for meat, supplemented by opportunistic hunting of small animals, as evidenced by cut-mark patterns on bones from sites like Olduvai Gorge that suggest access to carcasses rather than primary kills.^[122] By the Middle Paleolithic, subsistence strategies became more flexible, incorporating both hunting and scavenging of larger game, with Neanderthals demonstrating planned exploitation of herd animals through ambush tactics at sites such as Kebara Cave.^[123] The transition to the Upper Paleolithic marked a shift toward organized cooperative hunting, enabled by advanced projectile technologies and group coordination, allowing modern humans to target a broader spectrum of prey more efficiently. Similar practices emerged in Eurasia during this period, while in the terminal Pleistocene Americas (Paleoindian period), groups used Clovis points for big-game drives, herding large herbivores like mammoths into traps or over cliffs, as reconstructed from kill sites featuring these fluted points embedded in mammoth bones, indicating deliberate projectile use around 13,000 years ago.^[124]^[125] These fluted stone points, hafted to spears, facilitated communal hunts that maximized returns from megafauna, with evidence from sites like Murray Springs showing multiple animals processed in single events.^[126] The timing of Pleistocene megafauna extinctions varied regionally—staggered over 50,000–10,000 years ago in Eurasia, around 13,000–10,000 years ago in the Americas, and 50,000–40,000 years ago in Australia—coinciding with human hunting pressures and climate changes, prompting shifts to smaller, more resilient prey worldwide.^[127] Smaller game was pursued using snares and traps, as inferred from bone accumulations and tool microwear at Middle Stone Age sites like Sibudu Cave in South Africa, dated to over 70,000 years ago, where passive capture methods supplemented direct encounters.^[128] Such techniques, including pitfalls and nets implied by ethnographic analogies, allowed for sustained exploitation of fast-moving species like rabbits and birds during the Late Paleolithic.^[129] This evolution in strategies contributed to dietary diversification, though detailed nutritional impacts are addressed elsewhere.^[130] Gathering of wild plants, including nuts, tubers, and seeds, formed a critical component of Paleolithic economies, providing reliable carbohydrates and often comprising the majority of caloric intake in non-coastal environments.^[131] Ethnographic studies of recent hunter-gatherer societies, such as the Hadza and !Kung, infer that women typically led these efforts, collecting seasonally available resources during daily forays that required knowledge of plant phenology and processing techniques.^[132] Tools like wooden digging sticks, preserved at waterlogged sites, facilitated extraction of underground storage organs; for instance, over 30 such implements from the Gantangqing site in southwest China, dated to 300,000 years ago, show deliberate shaping for probing and uprooting tubers.^[133] These perishable artifacts highlight the underrepresentation of plant-based tools in the archaeological record but underscore gathering's role in risk buffering against hunting variability.^[134] Resource exploitation involved strategic mobility, with groups establishing seasonal camps near resource patches to optimize access to food and materials, as seen in the patterned distribution of artifacts at open-air sites like those in the Basque region, where lithic scatters indicate repeated occupations tied to herd migrations.^[135] Raw material quests drove long-distance travel for high-quality flint and obsidian, with Upper Paleolithic foragers transporting nodules up to 200 kilometers, as documented by sourcing analyses from sites in southwest Europe that reveal planned provisioning for tool maintenance during hunts.^[136] Recent excavations at Late Paleolithic sites in the southeastern margin of China's Badain Jaran Desert, reported in 2025, uncover diverse activities including hunting, plant processing, and lithic production, illustrating adaptive resource use in arid inland settings previously underrepresented in global narratives.^[137] Lithic tools for hunting, such as spear points, were central to these expeditions but are detailed in technological contexts.^[138]

Group Organization and Behavior

Paleolithic societies are inferred to have been organized into small, egalitarian bands typically ranging from 20 to 50 individuals, with fluid membership allowing for fission-fusion dynamics based on resource availability and seasonal needs.^[139] Archaeological evidence from camp scatters, such as clustered hearths and tool distributions at sites like Ohalo II in Israel, supports these modest group sizes, indicating temporary aggregations rather than permanent settlements.^[140] This structure, drawn from comparisons with modern hunter-gatherers and primate social groups, promoted mobility and low hierarchy, as larger units would strain foraging efficiency in varied Paleolithic environments.^[141] Kinship ties likely formed the core of these bands, with roles showing a flexible division of labor where hunting large game was often male-dominated, while gathering plant resources and small game collection involved both sexes, though recent analyses of ethnographic and later prehistoric evidence challenge rigid gender binaries, suggesting the possibility of female participation in big-game hunts.^[142] Evidence of elder care and compassion emerges from Neanderthal remains, such as the Shanidar 1 individual, who survived severe injuries with group support, and a child with Down syndrome from Cueva del Sidrón, Spain, who lived to at least age six despite debilitating conditions, suggesting empathetic caregiving within family units.^[143]^[144] These practices highlight relational bonds beyond immediate survival, integrating vulnerable members through shared responsibilities. Cooperative behaviors underpinned group stability, with widespread food sharing evident from faunal remains at sites like Kebara Cave, where diverse animal bones indicate communal processing and distribution to mitigate individual foraging risks.^[145] Conflict resolution likely involved rituals and alliances, inferred from low violence markers in skeletal records and ethnographic analogies.^[146] Footprint assemblages at White Sands National Park, dated to 21,000–23,000 years ago, preserve tracks of multiple individuals—including adults, adolescents, and children—suggesting coordinated group travel and possibly collective hunting of megafauna like giant sloths.^[147] Recent findings from Tinshemet Cave in Israel (as of 2025) indicate prolonged coexistence and possible interactions between Neanderthals and early Homo sapiens around 70,000–60,000 years ago, suggesting complex social dynamics across groups.^[148] Inferences from ethnographic analogies suggest relatively egalitarian gender dynamics, with women often central to gathering, which provided the majority of caloric intake, fostering mutual dependence without pronounced hierarchies.^[141]

Language, Symbolism, and Cognition

The cognitive evolution of Homo sapiens during the Paleolithic involved significant genetic and neurological changes that facilitated advanced communication and abstract thinking. Key genetic adaptations included derived mutations in the FOXP2 gene, associated with speech and language production, shared with Neanderthals and arising in their common ancestor with modern humans approximately 400,000–600,000 years ago.^[149] Concurrently, the human brain underwent substantial reorganization, including a threefold increase in size and structural shifts toward a more globular shape, enhancing cognitive capacities for complex problem-solving and social interaction.^[150]^[151] These developments marked a departure from earlier hominins, enabling sapiens to process symbolic information more efficiently. Evidence for early language capabilities in Paleolithic populations includes anatomical and environmental indicators suggesting proto-language use. The discovery of a well-preserved hyoid bone from a Neanderthal individual at Kebara Cave, dated to around 60,000 years ago, indicates that Neanderthals possessed the vocal tract anatomy necessary for producing a range of sounds similar to modern humans, supporting the potential for complex vocal communication.^[152] Acoustic properties of Paleolithic caves, such as enhanced reverberation in chambers with visual motifs, suggest that early humans may have selected sites for vocalization, where sound resonance could amplify group interactions and possibly aid in the development of spoken language.^[153] Additionally, the gestural origins hypothesis posits that language evolved from manual and facial gestures used by early hominins for coordination during activities like toolmaking, with archaeological evidence from tool assemblages indicating gestural signaling as a precursor to vocal speech around 2 million years ago, persisting into the Paleolithic.^[154] Symbolic behavior emerged as a hallmark of Paleolithic cognition, with material evidence pointing to intentional use of pigments and ornaments for non-utilitarian purposes. At Blombos Cave in South Africa, engraved ochre pieces dated to approximately 100,000 years ago demonstrate early abstract marking, likely serving symbolic roles in social or ritual contexts beyond practical applications like hide processing.^[155] Similarly, perforated Nassarius shell beads from Grotte des Pigeons in Morocco, dated to about 82,000 years ago, represent deliberate adornment, implying personal or group identity signaling through body modification.^[156] Advanced cognition is further evidenced by behaviors requiring foresight and empathy, such as intentional burials and resource planning. Paleolithic burials, beginning around 120,000 years ago at sites like Qafzeh Cave, suggest an awareness of others' mental states—termed theory of mind—through deliberate positioning of bodies with grave goods, indicating beliefs in post-mortem existence or social mourning.^[157] Tool caches, like those of prepared cores and unused implements found in Middle Paleolithic sites, reflect forward planning and mental representation of future needs, as early humans stockpiled materials anticipating scarcity or group mobility.^[158] Recent discoveries, including oversized handaxes from Kent, England, dated to 300,000 years ago, push the timeline for symbolic thought even earlier, potentially indicating ceremonial or status-related production among pre-sapiens hominins.^[159]

Cultural Expressions

Rock Art and Portable Art

Paleolithic rock art encompasses a range of visual expressions created on cave walls and rock shelters, primarily through the application of mineral pigments and incising techniques. These artworks, dating from the Upper Paleolithic period, often depict animals, human hands, and abstract signs, providing insights into early human symbolic behavior. Notable examples include the cave of Lascaux in France, where paintings of large herbivores such as horses, aurochs, and deer were created around 17,000 years ago using red ochre and black charcoal pigments mixed with binders like animal fat or water, applied via finger daubing, blowing, or rudimentary brushes made from moss or feathers.^[160]^[161] Engravings were also common, achieved by scraping the rock surface with flint tools to create outlines and textures that interacted with the cave's natural relief.^[162] In northern Spain's Altamira Cave, hand stencils—produced by placing a hand on the wall and blowing pigment over it—date to the Upper Paleolithic Gravettian to Middle Magdalenian periods, approximately 22,000–13,000 years ago, highlighting the use of red iron oxide for vivid contrast against limestone surfaces.^[163]^[164] Portable art, in contrast, consists of small-scale sculptures and carvings that could be carried by mobile hunter-gatherer groups, often fashioned from stone, bone, or ivory. These objects, emerging around 30,000 years ago, include anthropomorphic and zoomorphic figures that suggest personal or communal significance. The Venus of Willendorf, a limestone statuette from Austria standing about 11 cm tall, dates to 28,000–25,000 years ago and features exaggerated female forms—prominent breasts, hips, and abdomen—possibly symbolizing fertility or abundance, carved using stone tools to shape and polish the oolitic limestone surface coated with red ochre.^[165] In eastern Siberia, at sites like Mal'ta associated with the Mal'ta-Buret' culture, mammoth ivory was extensively used for portable art around 24,000–15,000 years ago; examples include delicately carved bird and bear figurines, crafted with fine burins and abrasives to achieve smooth contours and detailed features, reflecting advanced micro-lithic technology.^[166]^[167] Thematic content in Paleolithic art frequently revolves around fauna and human-animal interactions, with hunting scenes portraying dynamic pursuits of prey, as seen in later European caves, and a possible undercurrent of animism where animals were viewed as possessing spiritual essences integral to human survival.^[168]^[169] These motifs emphasize realism in European rock art, with animals rendered in profile using shading for depth, while abstract geometric patterns and hand motifs appear alongside, potentially serving mnemonic or ritual functions. Regional variations are evident: European parietal art prioritizes large-scale, naturalistic animal depictions in deep caves, whereas African examples, such as the engraved ochre plaques from Blombos Cave in South Africa dated to about 73,000 years ago, feature abstract cross-hatched designs incised with pointed stones, indicating an earlier tradition of non-figurative symbolism predating European cave paintings by tens of thousands of years.^[170]^[171] Recent archaeological findings have expanded the global scope of Paleolithic art, challenging Eurocentric narratives of its origins. In 2024, a cave painting in Leang Karampuang Cave, Sulawesi, Indonesia, depicting therianthropic figures hunting a warty pig, was dated to at least 51,200 years ago using uranium-series analysis of overlying calcite, surpassing many European examples in age and illustrating narrative complexity with human-animal hybrids.^[172] Similarly, African discoveries like the Blombos abstract drawing underscore that symbolic art likely emerged first in Africa, with portable and engraved forms appearing up to 73,000 years ago, well before the florescence of European cave art around 40,000 years ago. These advancements highlight the diverse, worldwide development of artistic expression during the Paleolithic, driven by cognitive capacities for symbolism briefly linked to broader social practices.^[173]

Music, Instruments, and Rituals

Evidence for musical practices during the Paleolithic period primarily derives from archaeological finds of instruments and contextual analyses of soundscapes, suggesting that sound played a significant role in social and performative activities among early modern humans and possibly Neanderthals.^[174] The earliest undisputed aerophones, dating to the Aurignacian culture around 43,000 years ago, were discovered in the Swabian Jura region of southwestern Germany, including the Hohle Fels cave where a flute crafted from the wing bone of a griffon vulture was found, featuring five finger holes and a V-shaped notch for blowing.^[174] These instruments indicate a developed musical tradition capable of producing melodic tones, with experimental reconstructions demonstrating diatonic scales similar to modern systems.^[174] Additional flutes from the same period, such as those made from mammoth ivory at Geißenklösterle cave, further attest to the use of diverse materials for sound production, with radiocarbon dating placing them between 42,000 and 43,000 calibrated years before present.^[174] Percussion instruments are inferred from modified animal bones, including mammoth scapulae and long bones showing wear patterns consistent with rhythmic striking, as evidenced by experimental archaeology replicating sounds from Ukrainian Upper Paleolithic sites.^[175] A controversial earlier example is the Divje Babe I artifact from Slovenia, a cave bear femur with two holes dated to approximately 60,000 years ago and associated with Neanderthals; while some analyses suggest intentional modification for musical use, others attribute the perforations to carnivore bites, though recent reviews (up to 2023) maintain the possibility of it being a simple flute.^[176]^[177] Sonic archaeology, or archaeoacoustics, has revealed how Paleolithic caves amplified and altered sounds, with acoustic modeling of sites like La Garma and Las Chimeneas in Spain showing that visual motifs such as dots and lines often align with areas of high reverberation or low-frequency resonance, implying deliberate selection of spaces for auditory effects in performances.^[153] Ethnographic analogies from modern hunter-gatherer societies suggest that pre-instrumental music likely involved vocalizations and body percussion, including rhythmic clapping and foot-stamping, which could have facilitated group synchronization without durable artifacts.^[178] Ritualistic uses of music are inferred from associations between instruments and symbolic artifacts, such as the dynamic "dancing" figurine from the Vogelherd cave in Germany, dated to around 40,000 years ago and found near multiple flutes, indicating performative dances during seasonal gatherings.^[179] Shamanistic elements appear in contexts where art and sound intersect, with cave acoustics potentially inducing altered states through echoes and resonances, as modeled in Upper Paleolithic sites, supporting rituals that blended auditory and visual symbolism for communal bonding or spiritual experiences.^[180] These practices likely reinforced social cohesion, though direct evidence remains sparse due to the perishable nature of non-osseous elements.^[178]

Spiritual Beliefs and Practices

Paleolithic spiritual beliefs are primarily inferred from archaeological evidence of intentional burials, symbolic artifacts, and artistic representations that suggest concepts of animism, an afterlife, and ritual mediation between the physical and spirit worlds. The earliest indications of such practices appear in the Middle Paleolithic, around 100,000 years ago, with intentional human burials accompanied by red ochre, a pigment often associated with symbolic or ritual significance. At Qafzeh Cave in Israel, excavations uncovered the remains of at least 15 early Homo sapiens individuals in shallow pits, some flexed in fetal positions and covered or surrounded by red ochre chunks and tools stained with the pigment, dating to approximately 92,000–115,000 years ago.^[181]^[182] This use of ochre, sourced from distant locations and processed for application, implies deliberate ritual actions possibly linked to beliefs in transformation or the afterlife, marking a shift from incidental body disposal to structured mortuary behavior.^[183] Neanderthals also exhibit evidence of burial practices during the Middle Paleolithic, suggesting shared or parallel spiritual concerns across hominin species. At Shanidar Cave in Iraq, multiple Neanderthal skeletons from around 60,000–70,000 years ago were found in clustered pits, with one (Shanidar IV) initially interpreted as a "flower burial" due to concentrated pollen from bee-pollinated plants like yarrow and hellebore around the body, hinting at intentional floral offerings as part of a ritual to honor the dead or invoke spiritual continuity.^[184] However, recent analyses, including 2023 sediment studies, attribute the pollen to ancient bee burrows rather than deliberate placement, challenging the flower offering hypothesis while affirming the intentionality of the interments themselves as evidence of emerging ritual complexity.^[185] These Middle Paleolithic burials, often simple pits with minimal goods, contrast with the Upper Paleolithic (approximately 50,000–12,000 years ago), where rituals evolved to include more elaborate grave goods, body orientations, and ochre applications, indicating intensified beliefs in post-mortem existence and communal mourning.^[186]^[187] Animistic beliefs, positing that animals, natural forces, and objects possess spirits or souls, are evoked through Upper Paleolithic art featuring therianthropic figures—hybrids blending human and animal forms—that likely represented interactions with a spirit world. Such depictions, including a 44,000-year-old hunting scene in Leang Bulu’ Sipong 4 cave on Sulawesi, Indonesia, show half-human, half-buffalo figures alongside therianthropes wielding spears, interpreted as shamans or ancestral spirits guiding human endeavors in a metaphysical realm. In European contexts, similar animal-human hybrids in caves like Chauvet (France, ~36,000 years ago) suggest totemistic elements, where clans identified with animal spirits for protection or power, bridging the material and supernatural. Venus figurines, small carved female forms from the Upper Paleolithic across Europe (e.g., Willendorf, Austria, ~25,000–30,000 years ago), further imply fertility cults within this animistic framework, with exaggerated breasts, hips, and vulvas symbolizing life-giving forces or earth mother deities to ensure reproductive success and communal survival.^[188]^[189] Shamanism, involving trance-induced journeys to spirit realms for healing or divination, finds support in the deliberate placement of Upper Paleolithic art in deep, dark cave sections inaccessible for daily use, evoking altered states of consciousness. Archaeologist Jean Clottes posits that these remote locations facilitated shamanic rituals, where practitioners entered trances to commune with animal spirits depicted on walls, as seen in the narrow, echoing chambers of Lascaux and Altamira caves.^[190] Evidence for trance induction includes potential use of psychotropic plants in later Upper Paleolithic Europe, with archaeological traces of poisonous and hallucinogenic substances like those from nightshade family plants or ephedra suggesting ritual ingestion to access visionary states, building on earlier medicinal plant knowledge.^[191]^[192] Although direct evidence from Spain around 13,000 years ago remains elusive, broader European pollen and residue analyses indicate such plants were incorporated into rituals by the Magdalenian period (~17,000–12,000 years ago).^[193] Recent research emphasizes the global scope of Paleolithic spirituality beyond Europe, with 2023–2024 studies reinterpreting non-European art and burials as inherently spiritual rather than merely decorative or functional. In the Levant, Early Upper Paleolithic (~38,000 years ago) engravings on a massive boulder deep in Manot Cave, Israel, alongside ochre and marine shell arrangements, point to collective rituals invoking symbolic protection or ancestral presence in a challenging environment.^[194] Similarly, in South Africa, 100,000-year-old ochre processing kits from Blombos Cave suggest early symbolic rituals tied to blood or life-force metaphors, paralleling animistic practices observed in later African hunter-gatherer traditions and expanding the timeline of spiritual expression.^[195] These findings underscore a diverse evolution of beliefs, from Middle Paleolithic mortuary simplicity to Upper Paleolithic shamanic complexity, rooted in human adaptation to existential uncertainties.

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Some present-day humans derive up to ~5% [1] of their ancestry from archaic Denisovans, an even larger proportion than the ~2% from Neanderthals.Missing: Homo sapiens<|separator|>
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[84]
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### Key Findings on Fire Evidence at Wonderwerk Cave
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[180]
Grave Markers : Middle and Early Upper Paleolithic Burials and the ...
For the Early Upper Paleolithic, it appears that males were buried more often than both females and juveniles. Adults appear to have accounted for at least ...
[181]
Hominin evolutionary thanatology from the mortuary to funerary realm
In some cases, corpses may have come to rest in natural (rather than deliberate) cuttings, such as the Neanderthal child from Roc de Marsal, France [40] or ...
[182]
Venus Figurines of the European Paleolithic: Symbols of Fertility or ...
Jan 3, 2012 · They may, instead, have symbolized the hope for survival and longevity, within well-nourished and reproductively successful communities. 1.
[183]
Perspective: Upper Paleolithic Figurines Showing Women with ...
Dec 1, 2020 · What is known about the Subject. Venus Figurines are the oldest art sculptures of humans known and tend to be of women who are obese or pregnant.Missing: portable | Show results with:portable
[184]
Dr Jean Clottes Paleolithic Cave Art of France - Bradshaw Foundation
Shamans will do so through trance. A shaman thus has a most important role as a mediator between the real world and the world of the spirits, as well as a ...
[185]
Paleomedicine and the Evolutionary Context of Medicinal Plant Use
Oct 9, 2020 · During the later Paleolithic, there is evidence for the use of poisonous and psychotropic plants, suggesting that Paleolithic humans built on ...
[186]
(PDF) Psychoactive Substances in Prehistoric Times: Examining the ...
Jan 2, 2015 · This paper aims to provide a comprehensive overview of the consumption of drug plants and fermented beverages in prehistoric times
[187]
The oldest archeological data evidencing the relationship of Homo ...
Jun 1, 2019 · Indicating a European use of ephedra as a psychoactive, a concentration of ephedra pollen was identified at the bottom of some amphorae dated ...Missing: Paleolithic | Show results with:Paleolithic
[188]
Early human collective practices and symbolism in the Early ... - PNAS
This study provides evidence indicating the practice of deep cave collective rituals in the Levant during the Early Upper Paleolithic (EUP) period.
[189]
Blood symbolism at the root of symbolic culture? African hunter ...
Blood/ochre/Pterocarpus are interchangeable in African hunter-gatherer ritual. A 'blood'-coded metaphor links women's reproduction to men's hunting.