A new reconstruction of the Yunxian 2 skull, and its implications for the relationship between the 'Archaic Homo sapiens' of China and Modern Humans.

The Middle Pleistocene, or Chibanian (roughly 774 100 to 129 000 years ago), has produced a range of Hominin fossils, with a surprising range of morphological diversity. Recent finds from Callao Cave on Luzon Island in 2007 and Rising Star Chamber in South Africa in 2013, have led to the description of two new Human species, Homo luzonensis and Homo naledi respectively, from this age. 

China has produced a diverse assemblage of Chibanian and older Hominin fossils, often labelled as 'Archaic Homo sapiens', although their relationship to Modern Humans has been the subject of debate for a long time. The Yunxian 1 & 2 crania were discovered on on a terrace of the Hanjiang River in the Yunyang District (formerly Yunxian) of Shiyan City, in Hubei Province, China, in 1989 and 1990. These skulls have been dated to between 1.1 million and 940 000 years ago, and are considered important for the understanding of the genus Homo in China, and to a certain extent the world, although both are heavily distorted, hampering attempts to reconstruct their original morphology and therefore phylogenetic relationships. 

The Yunxian 2 skull. Although the least distorted of the two crania from Yunxian, it is still partially crushed, and therefore difficult to interpret. Garry Todd/Hubei Provincial Museum/Wikimedia Commons.

In a paper published in the journal Science on 25 September 2025, a team of scientists led by Xiaobo Feng of the School of History and Culture at Shanxi University, Hubei Polytechnic University, and the Institute of Yunxian Man Site and School of History, Culture and Tourism, at Hanjiang Normal University, Qiyu Yin of the Institute of Vertebrate Paleontology and Paleoanthropology of the Chinese Academy of Sciences, and the College of Earth and Planetary Sciences of the University of the Chinese Academy of Sciences, Feng Gao of the Yunnan Institute of Cultural Relics and Archaeology, and Dan Lu, also of the Institute of Vertebrate Paleontology and Paleoanthropology of the Chinese Academy of Sciences, and the College of Earth and Planetary Sciences of the University of the Chinese Academy of Sciences, present a new reconstruction of the Yunxian 2 skull, and discuss the implications of that reconstruction for the relationship between the 'Archaic Homo sapiens' of China and Modern Humans.

Our understanding of the relationship between Modern Humans and our extinct Hominin relatives is based almost entirely on the reconstruction of the anatomy of extinct species from fossil specimens, many of which are quite distorted. Of the three fossil crania discovered at Yunxian, one (Yunxian 3) is still under preparation, having been found in 2022. Both of the previously discovered specimens are distorted, with Yunxian 1 being quite badly crushed as well. Computed tomography scans of Yunxian 2 have suggested that most of the deformation is due to the fragmentation and displacement of parts of the skull, rather than actual warping of the material. Feng et al. built a digital reconstruction of this fossil, using software to move parts back into their original position, and filling in some gaps with data from Yunxian 1. 

The reconstructed cranium is large and long, with a braincase which appears flattened in lateral view. It is smaller than the Harbin Xuchang crania, approximately the same size as the as Kabwe, Petralona, Bodo, Jinniushan, and Sangiran 17 crania, and larger than the Jebel Irhoud 1, Dali, and Maba crania. The reconstruction still lacks small parts of the zygomatic arches and the central incisors, but is otherwise intact. It contains a number of traits associated with earlier members of the genus Homo, including a thick supraorbital torus, a broad basicranium and palate, a long and low vault in lateral view, a receding frontal con￾tour, and a rather flat parietal contour, and a high, anteriorly projecting face. However, it lacks other traits associated with early members of the genus, such as a strongly angulated occipital with a prominent transverse torus. The specimen lacks the occipital bun, forward projecting midface, and general rounded 'en bombe' shape of Neanderthal skulls, and the upper parietal expansion of Modern Humans. The zygomaxillary region is transversely flat and faces anteriorly, similar to the Harbin, Dali, Jinniushan, and Hualongdong crania from Asia, as well as specimens assigned to Homo antecessor from Europe, as wellas Modern Humans. Its cheek￾bones are large and high.

Reconstruction of the Yunxian 2 cranium in standard views. (A) to (F) Anterior, posterior, inferior, superior, left, and right views, respectively. Brown colour indicates the fossil bone. The zygomatic bone and the tip of the left maxilla, as indicated with dark brown, were grafted and reconstructed by incorporating elements of Yunxian 1. White colour indicates the reconstructed parts inferred from the fracture edge and Yunxian 1. Neutral grey indicates the bones crushed and covered by other bones and matrix. Scale bar is 5 cm. Feng et al. (2025).

The matrix filling the skull is dense, and does not produce a good contrast with the bone material in computed tomography scans, preventing analysis of the features of the interior of the skull. However, Feng et al. were able to calculate an endocranial capacity of 1143 cm². However, the frontal lobe appears low and narrow, little expanded from the situation in early Homo, and unlike the expanded frontal lobes of Neanderthals and Modern Humans. 

Reconstruction of the endocranial cast of Yunxian 2. (A to F) Anterior, posterior, inferior, superior, left, and right views, respectively. Scale bar is 5 cm. Feng et al. (2025).

The traits which can be measured suggest that morphologically, Yunxian 2 appears to be intermediate between early Homo species and specimens such as Home erectus, Homo ergaster, and the Kabwe and Petralona crania, and later Asian skulls such as Harbin, Dali, and Jinniushan. 

A phylogenetic analysis based upon the new reconstruction of Yunxian 2, found it grouped with a group of Asian specimens including Dali, Jinniushan, Xujiayao, and Hualongdong and the Xiahe and Penghu mandibles, which have together been referred to as the Homo longi clade, based upon the suggestion that the Xujiayao, Xuchang, Xiahe, and Penghu specimens, as well as the Denisovans should be grouped together as a new species, Homo longi (although Feng et al. did not recover the Xuchang specimens as part of this group). Feng et al.'s reconstruction found this clade to be the sister to Homo antecessor, with the Homo longi clade plus Homo antecessor being the sister clade to Homo sapiens. Yunxian 2 is the oldest member of the Homo longi clade as recovered by Feng et al., but not the earliest branching. 

Phylogeny and divergence time of 57 selected fossil operational taxonomic units from the genus Homo. The topology of the tree was the majority consensus of the most parsimonious trees from the parsimony analysis in TNT. The divergence time was inferred from the Bayesian tip-dating analysis in MrBayes 3.2. Branch lengths are proportional to the division age in thousands of years (Ka). Numbers at the internal nodes are the median ages, and the blue bars indicate the 95% highest posterior density interval of the node ages. The red half-brackets on the right indicate the ranges of the Neanderthal, Homo longi, and Homo sapiens clades. The numbers in red highlight the ages of division of the three clades. Yunxian is also highlighted in red. Feng et al (2025).

The term 'Denisovan' was coined for a group of highly fragmentary fossils from the Denisova Cave in the Altai Mountains of Russia. Because of the fragmentary nature of these fossils, it has been possible to determine little about their original morphology, but DNA has been recovered from the specimens, revealing a great deal of genetic information. Studies of mitochondrial DNA from these specimens has suggested that this group branched off from the lineage that led to modern Humans before the Neanderthal line. However, studies based upon nuclear DNA have suggested that Denisovans and Neanderthals were sister groups, with a common ancestor more recent than their last common ancestor with Modern Humans. While the limited amount of data available makes this hard to resolve, Feng et al. suggest that the Denisovans were probably members of the Homo longi clade, with those morphological traits known all consistent with membership of this group. 

The divergence between the ancestors of Neanderthals and those of Modern Humans was for a long time considered to have happened between 700 000 and 500 000 years ago. However, more recent studies incorporating DNA recovered from Neanderthal specimens have suggested a much earlier split. Feng et al.'s analysis suggests that the Homo longi clade emerged as a distinct lineage about 1.2 million years ago, with Homo sapiens first appearing about 1.02 million years ago, around the time when Yunxian 2 was alive. The Homo longi and Homo sapiens clades are recovered as having diverged about 1.32 million years ago, while the Neanderthal clade diverged from the ancestors of these two groups about 1.38 million years ago.

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