This article presents a synthesis of recent developments in the study of human evolution over the past five years. It begins with an overview of hominin species nomenclature and diversity, followed by an examination of the proposed population bottleneck similar to 900,000 years ago. The discussion then turns to the ongoing debate regarding the Last Common Ancestor (LCA) of Homo sapiens, Neanderthals, and Denisovans, contextualized within patterns of hominin dispersal over the last million years. Key fossil discoveries, including Harbin, Homo luzonensis, Apidima 1, and Yunxian, are evaluated for their implications on evolutionary relationships. Finally, the paper explores recent findings related to hominin behaviour and considers their broader significance for understanding cognitive capacities in Middle to Late Pleistocene populations. Le pr & eacute;sent essai synth & eacute;tise les progr & egrave;s accomplis au cours des cinq derni & egrave;res ann & eacute;es en ce qui concerne l'& eacute;tude de l'& eacute;volution humaine. Il commence par une vue d'ensemble de la nomenclature et de la diversit & eacute; des esp & egrave;ces d'hominin & eacute;s, puis examine l'hypoth & egrave;se d'un goulet d'& eacute;tranglement de population il y a environ 900 000 ans. La discussion se porte ensuite sur le d & eacute;bat en cours & agrave; propos du dernier anc & ecirc;tre commun (DAC) d'Homo sapiens, des N & eacute;anderthaliens et des D & eacute;nisoviens, replac & eacute; dans le contexte des sch & eacute;mas de dispersion des hominin & eacute;s au cours du dernier million d'ann & eacute;es. Des d & eacute;couvertes cruciales, notamment celles de l'homme de Harbin, d'Homo luzonensis et des fossiles d'Apidima 1 et de Yunxian, ainsi que leurs implications sur les liens & eacute;volutionnaires, sont analys & eacute;es. Pour finir, l'article explore des r & eacute;sultats de travaux r & eacute;cents sur le comportement des hominin & eacute;s et ce qui en d & eacute;coule pour la compr & eacute;hension des capacit & eacute;s cognitives des populations du Pl & eacute;istoc & egrave;ne moyen & agrave; tardif.

In 1987, the academic conference ‘Origins and Dispersals of Modern Humans: Behavioural and Biological Perspectives’ was held in Cambridge, UK. Subsequently referred to as the ‘Human Revolution’ conference, this meeting brought together the most prominent academics working in the field of human origins, including archaeologists and palaeoanthropologists, as well as the geneticists who had recently published the first significant reconstruction of human evolution from mitochondrial DNA. The conference was particularly notable as the first platform to openly debate the fossil and revolutionary genetic evidence that pertained to two emerging theories of human origins, ‘Multiregionalism’ and ‘Recent African Origin’ – commonly known as ‘Out of Africa’. The importance of this conference was recognized at the time, and as such, BBC Radio 3 commissioned interviews with speakers and participants during the breaks; here we publish these interviews in full. Thirty-eight years on and the current major theories of human origins still trace back to those discussed at the Human Revolution conference. The article provides a fascinating historical perspective on scientific thought at the time and serves as a reminder of how quickly our current knowledge can be challenged and modified with the discovery of new fossil material or the rapid development of analytical techniques.

The site of Kozarnika in Bulgaria is a key, deep (~6m) archaeological sequence spanning the Lower, Middle and Upper Paleolithic of Europe. In its latter phases, it documents the transition from a Neanderthal-dominated Europe to one in which only modern humans are present. The process of this transition is of major importance to understanding the timeline of hominin dispersals, extinctions and replacement in Europe. We obtained 37 new AMS radiocarbon dates to build a robust chrono-stratigraphy for the site through sampling humanly modified bones, artifacts and human remains. We focussed particularly on dating the Initial Upper Paleolithic phase of the occupation, in geological layer 6/7, because of its potential link with early dispersing Homo sapiens populations and the presence of co-mingled human remains in the layer itself. The latter phases of human occupation at Kozarnika comprised five distinct periods, dated between about 50–26,000 years ago. We found that the 6/7 phase ranged from 48,500–44,050 cal BP, probably starting from about 49,960 cal BP. The stone tool evidence from the layer contains some Levallois pieces with echoes of the earlier Middle Paleolithic. One possible scenario is that both Neanderthals and early modern humans alternated in their use of the site during this period. Aside from the evidence from Apidima (Greece) which has been argued to evidence Homo sapiens at about 210,000 years ago (Harvati et al. 2019), Kozarnika is the earliest site found in southeastern Europe from the last 100,000 years which contains evidence for the presence of modern humans.

The year 2025 marked the ninetieth since a fossil hominin occipital bone was discovered in Swanscombe, southeast England. In subsequent years, its parietal bones were found, producing what remains the oldest partial cranium from Britain today. In the earliest analyses, it was interpreted as a descendant of the infamous fraudulent fossil Piltdown Man and evidence for the deep antiquity of Homo sapiens in Europe. Eventually, its affinity to Neanderthals was recognized, although its taxonomic attribution remains contentious. The evolving interpretation of Swanscombe over the past ninety years reflects the history of human origins research and highlights many of its challenges, including how factors such as geopolitics influence scientific interpretation. Today, in light of larger and more complete fossil assemblages, Swanscombe is sometimes a footnote in research; however, its age, location, and combination of morphological traits distinguish it among the Middle Pleistocene European fossils. Future comparisons to the similarly aged Sima de los Huesos assemblage in Spain would be particularly valuable for advancing our understanding of within-species diversity during the Middle Pleistocene. L'ann & eacute;e 2025 a marqu & eacute; le 90e anniversaire de la d & eacute;couverte d'un os occipital d'hominin & eacute; fossile & agrave; Swanscombe, dans le sud-est de l'Angleterre. Dans les ann & eacute;es qui ont suivi, les os pari & eacute;taux correspondants ont & eacute;t & eacute; retrouv & eacute;s. Cet ensemble constitue, aujourd'hui encore, le plus ancien cr & acirc;ne partiel de Grande-Bretagne. Les premi & egrave;res interpr & eacute;tations en ont fait un descendant du tristement c & eacute;l & egrave;bre Homme de Piltdown - d & eacute;masqu & eacute; depuis comme un faux - et une preuve de la tr & egrave;s ancienne pr & eacute;sence d'Homo sapiens en Europe. En d & eacute;finitive, son affinit & eacute; avec les N & eacute;anderthaliens a & eacute;t & eacute; reconnue, bien que sa place taxonomique reste controvers & eacute;e. L'& eacute;volution de l'interpr & eacute;tation de ce fossile au fil des 90 derni & egrave;res ann & eacute;es refl & egrave;te l'histoire de la recherche sur les origines de l'humanit & eacute; et met en lumi & egrave;re plusieurs des difficult & eacute;s auxquelles elle se heurte, notamment l'influence de facteurs tels que la g & eacute;opolitique sur l'interpr & eacute;tation scientifique. Aujourd'hui, eu & eacute;gard & agrave; d'autres assemblages fossiles plus importants et plus complets, le cr & acirc;ne de Swanscombe est parfois rel & eacute;gu & eacute; aux notes de bas de page des travaux de recherche. Pourtant, son & acirc;ge, son emplacement et sa combinaison de traits morphologiques le distinguent des autres fossiles du Pl & eacute;istoc & egrave;ne moyen europ & eacute;en. Il serait int & eacute;ressant, en particulier, de le comparer & agrave; l'assemblage de la Sima de los Huesos, en Espagne, qui date de la m & ecirc;me p & eacute;riode. Cela permettrait d'approfondir notre compr & eacute;hension de la diversit & eacute; intrasp & eacute;cifique au cours du Pl & eacute;istoc & egrave;ne moyen.

In 1987, the academic conference ‘Origins and Dispersals of Modern Humans: Behavioural and Biological Perspectives’ was held in Cambridge, UK. Subsequently referred to as the ‘Human Revolution’ conference, this meeting brought together the most prominent academics working in the field of human origins, including archaeologists and palaeoanthropologists, as well as the geneticists who had recently published the first significant reconstruction of human evolution from mitochondrial DNA. The conference was particularly notable as the first platform to openly debate the fossil and revolutionary genetic evidence that pertained to two emerging theories of human origins, ‘Multiregionalism’ and ‘Recent African Origin’ – commonly known as ‘Out of Africa’. The importance of this conference was recognized at the time, and as such, BBC Radio 3 commissioned interviews with speakers and participants during the breaks; here we publish these interviews in full. Thirty‐eight years on and the current major theories of human origins still trace back to those discussed at the Human Revolution conference. The article provides a fascinating historical perspective on scientific thought at the time and serves as a reminder of how quickly our current knowledge can be challenged and modified with the discovery of new fossil material or the rapid development of analytical techniques.

Our current understanding of the origins of Homo sapiens is limited, in part, by the fragmented fossil record from Late Pleistocene and early Holocene Africa. Here, we re-examine the Kabua 1 cranium, an enigmatic and little-studied Kenyan fossil discovered in the 1950s. We compare virtual reconstructions created previously by our team with a wide range of Middle Pleistocene to Early Holocene fossils and recent African H. sapiens crania to assess Kabua 1's morphological affinities. We also provide a conservative new Uranium-series minimum date of 64.4 +/- 5.4 ka. Our results highlight Kabua 1's broad similarity to derived H. sapiens, both recent and fossil. Its morphology nevertheless demonstrates some more basal affinities and thus contributes to our understanding of the depth of phenotypic diversity in Late Pleistocene African H. sapiens. Notre compr & eacute;hension actuelle des origines d'Homo sapiens est limit & eacute;e, en partie, par les lacunes des traces fossiles du Pl & eacute;istoc & egrave;ne tardif et de l'Holoc & egrave;ne pr & eacute;coce en Afrique. Les auteurs r & eacute;examinent ici le cr & acirc;ne Kabua 1, fossile & eacute;nigmatique et peu & eacute;tudi & eacute; qui fut d & eacute;couvert au Kenya dans les ann & eacute;es 1950. Afin d'& eacute;tudier ses affinit & eacute;s morphologiques, l'& eacute;quipe compare des reconstitutions en images de synth & egrave;se qu'elle a r & eacute;alis & eacute;es ant & eacute;rieurement & agrave; un large assortiment de fossiles datant du Pl & eacute;istoc & egrave;ne moyen & agrave; l'Holoc & egrave;ne pr & eacute;coce, ainsi qu'& agrave; des cr & acirc;nes d'H. sapiens africains r & eacute;cents. Les auteurs proposent & eacute;galement une nouvelle datation par les s & eacute;ries de l'uranium, conservatrice, de 64,4 +/- 5,4 ka au minimum. Les r & eacute;sultats mettent en lumi & egrave;re les larges similitudes entre Kabua 1 et les d & eacute;riv & eacute;s d'H. sapiens, r & eacute;cents aussi bien que fossiles. Toutefois, la morphologie de ce fossile montre certaines affinit & eacute;s plus basales et contribue ainsi & agrave; notre compr & eacute;hension de la vaste diversit & eacute; ph & eacute;notypique d'H. sapiens au Pl & eacute;istoc & egrave;ne tardif en Afrique.

Archaeological evidence suggests that dogs diverged from wolves during the Palaeolithic, more than 15,000 years ago1-7. The earliest unequivocal genetic evidence, however, is associated with dog remains from Mesolithic archaeological contexts approximately 10,900 years ago8,9. Here we generate both nuclear and mitochondrial genomes from canid remains at Pınarbaşı in Türkiye (15,800 years ago)10 and Gough's Cave in the UK (14,300 years ago)11, as well as from dogs excavated from two Mesolithic sites in Serbia (Padina between 11,500-7,900 years ago and Vlasac 8,900 years ago)12,13. Our analyses indicate that a genetically homogeneous dog population was already widely distributed across Europe and Anatolia during the Late Upper Palaeolithic (by at least 14,300 years ago). This finding suggests that dogs were exchanged among genetically and culturally distinct western Eurasian Late Palaeolithic human populations, namely the Magdalenian, Epigravettian and Anatolian hunter-gatherers10,14-16. Last, we identify a major influx of eastern Eurasian dog ancestry during the Mesolithic, concomitant with the movement of eastern hunter-gatherer populations into Europe14, which led to the establishment of the primary ancestry characteristics that define European dog populations today.

Our current understanding of the origins of Homo sapiens is limited, in part, by the fragmented fossil record from Late Pleistocene and early Holocene Africa. Here, we re‐examine the Kabua 1 cranium, an enigmatic and little‐studied Kenyan fossil discovered in the 1950s. We compare virtual reconstructions created previously by our team with a wide range of Middle Pleistocene to Early Holocene fossils and recent African H. sapiens crania to assess Kabua 1's morphological affinities. We also provide a conservative new Uranium‐series minimum date of 64.4 ± 5.4 ka. Our results highlight Kabua 1's broad similarity to derived H. sapiens , both recent and fossil. Its morphology nevertheless demonstrates some more basal affinities and thus contributes to our understanding of the depth of phenotypic diversity in Late Pleistocene African H. sapiens .

In 1987, the academic conference 'Origins and Dispersals of Modern Humans: Behavioural and Biological Perspectives' was held in Cambridge, UK. Subsequently referred to as the 'Human Revolution' conference, this meeting brought together the most prominent academics working in the field of human origins, including archaeologists and palaeoanthropologists, as well as the geneticists who had recently published the first significant reconstruction of human evolution from mitochondrial DNA. The conference was particularly notable as the first platform to openly debate the fossil and revolutionary genetic evidence that pertained to two emerging theories of human origins, 'Multiregionalism' and 'Recent African Origin' - commonly known as 'Out of Africa'. The importance of this conference was recognized at the time, and as such, BBC Radio 3 commissioned interviews with speakers and participants during the breaks; here we publish these interviews in full. Thirty-eight years on and the current major theories of human origins still trace back to those discussed at the Human Revolution conference. The article provides a fascinating historical perspective on scientific thought at the time and serves as a reminder of how quickly our current knowledge can be challenged and modified with the discovery of new fossil material or the rapid development of analytical techniques. La conf & eacute;rence universitaire << Origines et dispersions des humains modernes : points de vue comportementaux et biologiques >> s'est tenue en 1987 & agrave; Cambridge, au Royaume-Uni. Cette rencontre, que l'on appellera par la suite la conf & eacute;rence de la << r & eacute;volution humaine >>, r & eacute;unissait les chercheurs les plus & eacute;minents dans le domaine des origines humaines, arch & eacute;ologues et pal & eacute;ontologues notamment, mais aussi des g & eacute;n & eacute;ticiens qui venaient de publier la premi & egrave;re reconstitution significative de l'& eacute;volution humaine & agrave; partir de l'ADN mitochondrial. Cette conf & eacute;rence fut particuli & egrave;rement remarquable en cela qu'elle constitua le premier forum o & ugrave; furent discut & eacute;s les indices fossiles et les preuves g & eacute;n & eacute;tiques r & eacute;volutionnaires concernant deux th & eacute;ories & eacute;mergentes des origines humaines : l'hypoth & egrave;se africaine et le multir & eacute;gionalisme. Eu & eacute;gard & agrave; l'importance de l'& eacute;v & eacute;nement, BBC Radio 3 commanda des interviews d'intervenants et de participants qui furent enregistr & eacute;es pendant les pauses. Ces entretiens sont retranscrits in extenso dans le pr & eacute;sent article. Trente-huit ans plus tard, les grandes th & eacute;ories actuelles des origines humaines se r & eacute;clament encore des approches discut & eacute;es & agrave; la conf & eacute;rence sur la r & eacute;volution humaine. Cet article offre un point de vue historique fascinant sur la pens & eacute;e scientifique de l'& eacute;poque et nous rappelle que nos connaissances peuvent & ecirc;tre remises en question et modifi & eacute;es & agrave; tout moment par la d & eacute;couverte de nouveaux fossiles ou le d & eacute;veloppement rapide des techniques d'analyse.

A recently excavated Neanderthal skeleton from southern France has yielded DNA from a distinct lineage, different from other late Neanderthals. This suggests Neanderthals expanded and diversified about 120,000 years ago, and some of that diversity persisted in Europe until near the time of their extinction.

Assigning an age to the nearly complete cranium found in the Petralona Cave in Greece is of outstanding importance because this fossil has a key position in European human evolution. This topic has been debated since its discovery more than 60 years ago, highlighting the difficulties in applying physical dating methods to prehistoric samples. Previous results obtained on various types of samples yielded a large age range between about 170 and 700 ka, precluding any consensus on the age of the human fossil. On the other hand, the original stratigraphic position of the cranium also remains enigmatic, in spite of all the efforts provided by various researchers. Here, we present new U-series dates performed on the calcite that grew directly on the cranium, which is the only sample able to provide crucial information on the age of the fossil. The results yield a finite age suggesting that the Petralona cranium has a minimum age of 286 ± 9 ka. Other speleothems and calcitic coatings were sampled in three main locations in the cave; among them, samples came from the 'Mausoleum' where the cranium was supposedly found cemented to a wall. The data show that the calcite covering the cranium is not contemporaneous with that of the Mausoleum wall, despite what was previously thought. The different possibilities, depending on whether or not the cranium was attached to the wall, are discussed in the paper. From a morphological point of view, the Petralona hominin forms part of a distinct and more primitive group than Homo sapiens and Neanderthals, and the new age estimate provides further support for the coexistence of this population alongside the evolving Neanderthal lineage in the later Middle Pleistocene of Europe.

Archaeology and the branch of population genetics focusing on the human past have historically lived parallel lives, often having complicated encounters when it came to unravelling the origins and evolution of Homo sapiens. These interactions were proven invaluable to obtain a deeper and more complete understanding of our past. At the same time, they sometimes uncovered biases and misinterpretations, with serious consequences for our understanding of data, methods and, most importantly, the history of our species. Cavalli-Sforza pioneered a real multidisciplinary approach, bridging population genetics and statistics with archaeology, human origins, and other fields in the humanities, inspiring researchers from these fields and blazing a trail for today’s successful interactions and collaborations. His legacy showed that these interdisciplinary approaches are possible and of vital importance, and exposed areas that still need significant development today.

OBJECTIVE:This study follows up on our recent morphological analysis of the juvenile maxilla from Mugharet el'Aliya, Morocco. Although this specimen shows a reportedly archaic morphology, likely due to its large size, 3D shape analyses indicated affinities with early Homo sapiens. Here, we conducted an in-depth comparative investigation of the associated dentition to further clarify this individual's phylogenetic and taxonomic affinities. MATERIALS AND METHODS:Our analyses were based on three kinds of data: (a) external crown dimensions and non-metric features, analyzed via summary statistics; (b) CT scan data enabling the study of internal structures (enamel-dentine junction) via geometric morphometrics; and (c) high-resolution replicas of the external surface of the upper canine enabling the study of perikymata numbers via probability functions. The comparative samples included Middle Pleistocene (Chibanian) Europeans and Africans, Neanderthals, and early and later H. sapiens. RESULTS:Mugharet el'Aliya showed the greatest similarities in external and internal tooth morphology with early and later H. sapiens. Perikymata counts cluster the upper canine with H. sapiens. However, its canine and fourth premolar are megadont at a level generally atypical for H. sapiens. DISCUSSION:Our analyses of the dentition of the Mugharet el'Aliya individual support our previous findings on the morphological analysis of the maxilla, placing this fossil closest to H. sapiens. Our study further strengthens the evidence connecting fossils from the North African Aterian to those from Western Asia, especially Qafzeh. We also provide the first comparative analysis of a permanent upper canine from the Aterian fossil record.

1920s/30s excavation of a Middle Devensian sequence in the northern part of Kents Cavern recovered important Late Middle and Early Upper Palaeolithic archaeological material, including Britain's oldest known Homo sapiens remains. Questions remain about this material, including how it came to be in the cave. Judged by the recorded distribution of finds it may have entered via the Northeast Gallery. A previously unrecorded entrance into the cave from the Northeast Gallery was identified in 2014, and a column through the entrance's sedimentary fill was excavated during 2015–2016. The results of that work are reported here. The entrance retains an intact and well‐stratified Pleistocene sequence comparable to the ‘Cave Earth’ unit described previously inside the cave. The uppermost part of the newly recognised Northeast Gallery entrance sequence has been removed by historical excavation, with most of the remaining sediments spanning the Middle Devensian and earlier part of the Late Devensian. The sequence contains bone and pollen, and ancient mammalian DNA is preserved within the sediments. The base of the Northeast Gallery entrance sequence is formed of a clayey diamict comparable to the cave's ‘Breccia’ unit, a deposit currently understood as Middle Pleistocene, and previously identified only in Kents Cavern's southern chambers. Comparison of the excavated sequence with the historical record of the Vestibule excavation shows that the basal cave earth deposits in the Northeast Gallery entrance are significantly higher than those inside the cave. Although requiring further work to confirm, this suggests that the Northeast Gallery could have played a major role in the accumulation of material in the Vestibule during the Middle Devensian.

Diverse forms of Homo coexisted during the Middle Pleistocene. Whether these fossil humans represent different species or clades is debated. The ~1-million-year-old Yunxian 2 fossil from China is important for understanding the cladogenesis of Homo and the origin of Homo sapiens . In this study, we restored and reconstructed the distorted Yunxian 2 cranium using recently introduced technology. The results show that this cranium displays mosaic primitive and derived features. Morphometric and phylogenetic analyses suggest that it is an early member of the Asian H. longi clade, which includes the Denisovans and is the main part of the sister group to the H. sapiens clade. Both the H. sapiens and H. longi clades have deep roots extending beyond the Middle Pleistocene and probably experienced rapid early diversification. Yunxian 2 may preserve transitional features close to the origins of the two clades.

Fire-making is a uniquely human innovation that stands apart from other complex behaviours such as tool production, symbolic culture and social communication. Controlled fire use provided adaptive opportunities that had profound effects on human evolution. Benefits included warmth, protection from predators, cooking and creation of illuminated spaces that became focal points for social interaction1-3. Fire use developed over a million years, progressing from harvesting natural fire to maintaining and ultimately making fire4. However, determining when and how fire use evolved is challenging because natural and anthropogenic burning are hard to distinguish5-7. Although geochemical methods have improved interpretations of heated deposits, unequivocal evidence of deliberate fire-making has remained elusive. Here we present evidence of fire-making on a 400,000-year-old buried landsurface at Barnham (UK), where heated sediments and fire-cracked flint handaxes were found alongside two fragments of iron pyrite-a mineral used in later periods to strike sparks with flint. Geological studies show that pyrite is locally rare, suggesting it was brought deliberately to the site for fire-making. The emergence of this technological capability provided important social and adaptive benefits, including the ability to cook food on demand-particularly meat-thereby enhancing digestibility and energy availability, which may have been crucial for hominin brain evolution3.

Abstract Tracking the origins of new species and delimiting taxa across space and time present well-trodden sources of controversy for palaeoanthropology. Although biological diversity comes with frustratingly elusive boundaries, the task of describing and understanding diversity remains no less crucial, and palaeotaxonomy no more dispensable. This is epitomized by recent developments in discussions on our species’ origins and the extent to which Middle Pleistocene hominin forms represent distinct lineages. While it is tempting to think that progress in such debates is only hampered by the paucity of fossil and genomic data, we argue that problems also lie with unrealistic assumptions in theory. In particular, we examine ongoing discussions on whether Homo sapiens and Neanderthals deserve distinct species status as a means to advocate for the necessity of reframing speciation in palaeoanthropology in a more biologically plausible way. We argue that available palaeontological evidence is best interpreted under a framework that sees speciation as an evolutionary process that starts in space, thereby involving a geographical dimension, and progresses in time, thereby involving a diachronic dimension, with an incremental accumulation of relevant characters at different phases of the process. We begin by discussing evidence about species-level differentiation of H. sapiens and Neanderthals and analyse major sources of taxonomic disagreement, before illustrating the potential of this perspective in making progress on the earliest stages of H. sapiens speciation within Africa.

This study analyzes the effects of bioclimate and masticatory factors on the regional variability of human cranial forms across 150 ethnic groups worldwide. Morphometric variables were generated using principal component analysis applied to 3D homologous models. Relationships between cranial form and bioclimate (temperature and precipitation) and masticatory factors (infratemporal space) were tested considering sampling bias due to past population movements during the late Pleistocene and/or early- to mid-Holocene. Cranial size correlated with thermal conditions, consistent with Bergmann’s rule. The length/breadth proportion of the neurocranium aligned with Allen’s rule for thermal adaptation, while no relationship with masticatory stress was found. Facial form responded to either climate or masticatory conditions, although the primary factor was unclear due to the high correlation between stresses. However, masticatory stress was identified as an equally significant factor behind facial flatness in cold regions, else than the effect of Allen’s rule. High narrowness of nasal and orbital openings correlated significantly with cold temperatures and cranial size, suggesting not only functional but also allometric effect. This study demonstrated the complexity of environmental influences on cranial form diversity, nonetheless suggested reduction of selective pressure on cranial form caused by natural environmental stress due to the development of civilization.

Interview with Chris Stringer, who studies human origins at the Natural History Museum of London.

Diverse Middle Pleistocene forms of Homo coexisted in Africa, Europe, and Asia. It is very controversial whether these fossil humans represent different species or lineages. The ∼1 Ma old Yunxian 2 fossil from China is crucial for understanding the cladogenesis of Homo and the origin of Homo sapiens . Here, we restored and reconstructed the distorted Yunxian 2 cranium using new technology. The results show that this cranium displays mosaic features of plesiomorphy and apomorphy. Phylogenetic analyses and Bayesian tip-dating including the reconstructed Yunxian 2 suggest that it is an early member of the Asian ‘Dragon Man’ lineage, which probably includes the Denisovans, and is the sister group of the Homo sapiens lineage. Both the H. sapiens and Dragon Man lineages had deep roots extending beyond the Middle Pleistocene, and the basal position of the Yunxian fossil cranium suggests it represents a population lying close to the last common ancestor of the two lineages.One-Sentence Summary The newly-reconstructed Yunxian 2 cranium represents a basal member of the Dragon Man and Denisovan lineage, and probably lies close to the last common ancestor of that lineage and the lineage of H. sapiens .### Competing Interest StatementThe authors have declared no competing interest.