Denisovan mitochondrial DNA from dental calculus of the >146,000-year-old Harbin cranium.

Late Middle Pleistocene hominin teeth from Panxian Dadong, South China

Structural analysis of premolar roots in Middle Pleistocene hominins from China

Africa demonstrates a complex process of the hominin evolution with a series of adaptive radiations during several millions of years that led to diverse morphological forms. Recently, Hammer et al. (2011) and Harvati et al. (2011) provided integrated morphological and genetic evidence of interbreeding between modern humans and unknown archaic hominins in Africa as recently as 35,000 years ago. However, a genetic evidence of hybridization between hominin lineages during the Lower and Middle Pleistocene epochs is unknown and the direct retrieval of DNA from extinct lineages of African hominins remains elusive. The availability of both nuclear and mitochondrial genome sequences from modern humans, Neanderthals, and Denisovans allows collecting nuclear DNA sequences of mitochondrial origin (numts) inserted into the nuclear genome of the ancestral hominin lineages and drawing conclusions about the hominin evolution in the remote past. The mtDNA and numt analysis uncovered a deep division of mtDNA lineages that existed in African hominins in the Middle Pleistocene. The first cluster included the human and Neanderthal-like mtDNA sequences while the second consisted of DNA sequences that are known today as mtAncestor-1, a nuclear fossil of the mtDNA, and the Denisova mtDNA isolated from a bone and a tooth found in southern Siberia. The two groups initially diverged 610,000-1,110,000 years ago. Approximately 220,000 years after the primary split, the Denisova - mtAncestor-1 mtDNA lineages mixed with the mtDNA pool of an ancestral population of Neanderthals and modern humans. This admixture after the profound division is demonstrated by the transposition of the Denisova-like mtDNA sequence into the nuclear genome of an ancestor of Neanderthals and modern humans. This finding suggests the matrilineal genetic structure among the Middle Pleistocene hominins as well as the existence of gene flow between African hominin lineages. Through paleogenomic analyses, it is impossible to exclude the theory that population structure and gene flow in African hominins influenced the admixture pattern observed in the nuclear genomes of non-Africans.

Morphological description and evolutionary significance of 300 ka hominin facial bones from Hualongdong, China

Another look at the nose and the functional significance of the face and nasal mucous membrane for cooling the brain in fossil hominids

The aim of this study is to explore the root and root canal morphology of Homo fossil occupying China during the Middle Pleistocene period. Human occupation and evolutionary dynamics in East Asia during the Middle Pleistocene period is one of the most intriguing issues in paleoanthropology, with the coexistence of multiple lineages and regional morphs suggesting a complex population interaction scenario. Although premolar root and canal morphology has certain phylogenetic, taxonomic, and functional implications, its morphological diversity, possible evolutionary trend and characteristics regarding Middle Pleistocene hominins inhabiting East Asia are still insufficiently understood; where these populations fits within the Homo lineage (with respect to root and pulp canal structure) needs to be explored. Using microtomography, we directly observed and assessed the nonmetric variability of root and canal forms in maxillary and mandibular premolars of Chinese Middle Pleistocene Homo (N = 19), and compared our observed variations with Eurasian Early Pleistocene specimens from the Asia continent (N = 1) and Java (N = 2), as well as with Neanderthals (N = 28) and recent modern humans (N = 67). A total number of nine types of root-canal forms were recorded. As a whole, the Chinese Middle Pleistocene record shows an evolutionary trend toward a modern human-like condition (a reduction of root/canal number and a simplification of root surface structure). We documented primitive signals like high percentage of Tomes' root in lower premolars. A considerable occurrence of incompletely separated root branches and bifid root and canal apices, representing evolutionary transformation from multi-root to single-root condition was also noticed. The results were compared with previous publications on Early and Middle Pleistocene Homo in East Africa, North Africa, and Eurasia. This work provides new original data, incorporates the latest human fossil discoveries and suggests that analyzing the variation of premolar root structural organization, notably integrating together root/canal form and number, could possibly contribute to taxonomic and phylogenetic assessments. The mid-Middle Pleistocene populations, or "classic" Homo erectus, in our study show closer affinity to Early and Middle Pleistocene hominins in Eurasia, than to East African early Homo, which supports the suggestion that at least some of the Early Pleistocene hominin groups in Eurasia contribute to the later population; on the other hand, it is still difficult to clearly trace the evolutionary fate of those late Middle Pleistocene populations (roughly assigned as archaic Homo sapiens through a craniodental perspective). More comparable materials from the Early to Middle Pleistocene period as well as precise chronological framework is needed to further explore the evolutionary trends of archaic hominins in the Asian continent before the arrival of modern humans.

This study aimed to understand the ontogenetic and allometric relationships in scaling between the anterior and posterior openings of the cranial airways and facial size, in order to shed light on the mechanisms that might underlie the evolution of a large face and large airways in Middle Pleistocene hominins and Neandertals. Sizes were calculated from 3D landmarks measured on the facial skeleton and airway structures of 403 skulls from two ontogenetic series of H. sapiens and P. troglodytes, an adult sample of gorillas and 11 Middle Pleistocene hominins and Neandertals. RMA regression models were used to compare the patterns in scaling between the anterior and posterior airways in relation to overall facial size. Our results show that the size of the anterior airways correlates more positively with facial size than the size of the posterior airways. This ontogenetic mechanism could explain the large faces and noses in the Neandertal lineage despite the adverse effects of such a phenotype for respiratory air-conditioning in cold climates. A large facial size could be a developmentally constrained consequence of generating airways large enough to provide the necessary oxygen for high energy demand in this large-brained and heavy-bodied hominin lineage.

A unique assemblage of 28 hominin individuals, found in Sima de los Huesos in the Sierra de Atapuerca in Spain, has recently been dated to approximately 430,000 years ago. An interesting question is how these Middle Pleistocene hominins were related to those who lived in the Late Pleistocene epoch, in particular to Neanderthals in western Eurasia and to Denisovans, a sister group of Neanderthals so far known only from southern Siberia. While the Sima de los Huesos hominins share some derived morphological features with Neanderthals, the mitochondrial genome retrieved from one individual from Sima de los Huesos is more closely related to the mitochondrial DNA of Denisovans than to that of Neanderthals. However, since the mitochondrial DNA does not reveal the full picture of relationships among populations, we have investigated DNA preservation in several individuals found at Sima de los Huesos. Here we recover nuclear DNA sequences from two specimens, which show that the Sima de los Huesos hominins were related to Neanderthals rather than to Denisovans, indicating that the population divergence between Neanderthals and Denisovans predates 430,000 years ago. A mitochondrial DNA recovered from one of the specimens shares the previously described relationship to Denisovan mitochondrial DNAs, suggesting, among other possibilities, that the mitochondrial DNA gene pool of Neanderthals turned over later in their history.

Homo erectus and Middle Pleistocene hominins: Brain size, skull form, and species recognition

Homo erectus in Salkhit, Mongolia?

Late Middle Pleistocene hominin teeth from Tongzi, southern China

Morphological description and comparison of the dental remains from Atapuerca-Sima de los Huesos site (Spain)

Hominin diversity in East Asia during the Middle Pleistocene: A premolar endostructural perspective

The hominin teeth from the late Middle Pleistocene Hualongdong site, China.

In 1978, a nearly complete hominin fossil cranium was recovered from loess deposits at the site of Dali in Shaanxi Province, northwestern China. It was subsequently briefly described in both English and Chinese publications. Here we present a comprehensive univariate and nonmetric description of the specimen and provide comparisons with key Middle Pleistocene Homo erectus and non-erectus hominins from Eurasia and Africa. In both respects we find affinities with Chinese H. erectus as well as African and European Middle Pleistocene hominins typically referred to as Homo heidelbergensis. Specifically, the Dali specimen possesses a low cranial height, relatively short and arched parietal bones, an angled occipital bone, and a nonprominent articular tubercle relative to the preglenoid surface all of which distinguish it from Afro/European Middle Pleistocene Homo and align it with Asian H. erectus. At the same time, it displays a more derived morphology of the supraorbital torus and supratoral sulcus and a thinner tympanic plate than H. erectus, a relatively long upper (lambda-inion) occipital plane with a clear separation of inion and opisthocranion, and an absolute and relative increase in brain size, all of which align it with African and European Middle Pleistocene Homo. Finally, traits such as the form of the frontal keel and the relatively short, broad midface align Dali specifically with other Chinese specimens from the Middle Pleistocene and Late Pleistocene, including H. erectus, and differentiate these from the Afro/European specimens of this time period.