Hominin evolution and diversity: a comparison of earlier-Middle and later-Middle Pleistocene hominin fossil variation in China.

Historical views of Asia as an evolutionary 'backwater' are associated with the idea that Homo erectus experienced long periods of stasis and ultimately went extinct. However, recent discoveries of well-dated Middle Pleistocene hominin fossils in China have considerably challenged these ideas and provide sufficient data to propose a testable model that explains the patterning of variation in Middle Pleistocene China, and why it changed over time. A series of hominin fossil studies comparing earlier-Middle and later-Middle Pleistocene groups confirm that the expressions of certain traits shift around 300 ka. Fossils from the later Middle Pleistocene are more variable with a mix of archaic traits as well as ones that are common in Western Eurasian early Homo sapiens and Neanderthals. The period around 300 ka appears to have been a critical turning point for later-Middle Pleistocene morphological changes in China. It coincides with a phase of climatic instability in the Northern Hemisphere between Marine Isotope Stages 12 and 10 that would have led to changes in gene flow patterning, and regional population survival/extinction. This localized and testable model can be used for future explorations of hominin evolution in later Pleistocene eastern Eurasia. This article is part of the theme issue 'The impact of Chinese palaeontology on evolutionary research'.

Evolution of cranial capacity revisited: A view from the late Middle Pleistocene cranium from Xujiayao, China.

The Late Middle Pleistocene hominin fossils from the Xujiayao site in northern China have been closely studied in light of their morphological variability. However, all previous studies have focused on separated cranial fragments. Here, we report the first reconstruction of a fairly complete posterior cranium, Xujiayao 6 (XJY 6), confidently dated to ∼200-160 ka, which facilitated an assessment of its overall cranial size. XJY 6 was reconstructed from three of the original fragments-the PA1486 (No.7/XJY 6a) occipital bone, PA1490 (No.10/XJY 6b) right parietal bone, and PA1498 (No.17/XJY 15) left temporal bone-which originated from the same young adult individual. The XJY 6 endocranial capacity, estimated by measuring endocranial volume, was estimated using multiple regression formulae derived from ectocranial and endocranial measurements on select samples of Pleistocene hominins and recent modern humans. The results indicate that the larger pooled sample of both Pleistocene and recent modern humans was more robust for the endocranial capacity estimate. Based on the pooled sample using the ectocranial and endocranial measurements, we conservatively estimate the XJY 6 endocranial volume to be ∼1700 cm3 with a 95% confidence interval of 1555-1781 cm3. This is close to Xuchang 1, which dates to 125-105 ka and whose endocranial volume is ∼1800 cm3. Thus, XJY 6 provides the earliest evidence of a brain size that falls in the upper range of Neanderthals and modern Homo sapiens. XJY 6, together with Xuchang 1, Homo floresiensis, Homo luzonensis, and Homo naledi, challenge the general pattern that brain size gradually increases over geological time. This study also finds that hominin brain size expansion occurred at different rates across time and space.

Conceptual issues in hominin taxonomy: Homo heidelbergensis and an ethnobiological reframing of species.

Efforts to name and classify Middle Pleistocene Homo, often referred to as "Homo heidelbergensis" are hampered by confusing patterns of morphology but also by conflicting paleoanthropological ideologies that are embedded in approaches to hominin taxonomy, nomenclature, and the species concept. We deconstruct these issues to show how the field's search for a "real" species relies on strict adherence to pre-Darwinian essentialist naming rules in a post-typological world. We then examine Middle Pleistocene Homo through the framework of ethnobiology, which examines on how Indigenous societies perceive, classify, and name biological organisms. This research reminds us that across human societies, taxonomies function to (1) identify and classify organisms based on consensus pattern recognition and (2) construct a stable nomenclature for effective storage, retrieval and communication of information. Naming Middle Pleistocene Homo as a "real" species cannot be verified with the current data; and separating regional groups into distinct evolutionary lineages creates taxa that are not defined by readily perceptible or universally salient differences. Based on ethnobiological studies of this kind of patterning, referring to these hominins above the level of the species according to their generic category with modifiers (e.g., "European Middle Pleistocene Homo") is consistent with observed human capabilities for cognitive differentiation, is both necessary and sufficient given the current data, and will allow for the most clear communication across ideologies going forward.

A multivariate assessment of the Dali hominin cranium from China: Morphological affinities and implications for Pleistocene evolution in East Asia.

OBJECTIVES: A nearly complete hominin fossil cranium from Dali in Shaanxi Province, China was excavated in 1978. We update and expand on previous research by providing a multivariate analysis of the specimen relative to a large sample of Middle and Late Pleistocene hominins. MATERIALS AND METHODS: We apply principal components analysis, discriminant function analysis, and a method of assessing group membership based on a soft independent model of class analogy (SIMCA) to the study of Dali's cranial morphology. We evaluate Dali's affinities within the context of Middle and Late Pleistocene Homo patterns of craniofacial morphology. RESULTS: When just the facial skeleton is considered, Dali aligns with Middle Paleolithic H. sapiens and is clearly more derived than African or Eurasian Middle Pleistocene Homo. When just the neurocranium is considered, Dali is most similar to African and Eastern Eurasian but not Western European Middle Pleistocene Homo. When both sets of variables are considered together, Dali exhibits a unique morphology that is most closely aligned with the earliest H. sapiens from North Africa and the Levant. DISCUSSION: These results add perspective to our previous view of as Dali a "transitional" form between Chinese H. erectus and H. sapiens. Athough no taxonomic allocation is appropriate at this time for Dali, it appears to represent a population that played a more central role in the origin of Chinese H. sapiens. Dali's affinities can be understood in the context of Wu's Continuity with Hybridization scenario and a braided-stream network model of gene flow. Specifically, we propose that Pleistocene populations in China were shaped by periods of isolated evolutionary change within local lineages at certain times, and gene flow between local lineages or between Eastern and Western Eurasia, and Africa at other times, resulting in contributions being made in different capacities to different regions at different times.

A description of the geological context, discrete traits, and linear morphometrics of the Middle Pleistocene hominin from Dali, Shaanxi Province, 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.

Geometric variation of the frontal squama in the genus homo: frontal bulging and the origin of modern human morphology.

The majority of studies of frontal bone morphology in paleoanthropology have analyzed the frontal squama and the browridge as a single unit, mixing information from different functional elements. Taking into account that the bulging of the frontal bone is often described as a species-specific trait of Homo sapiens, in this article we analyze variation in the midsagittal profile of the genus Homo, focusing on the frontal squama alone, using landmark-based superimpositions and principal components analysis. Our results demonstrate that anatomically modern humans are definitely separated from extinct human taxa on the basis of frontal bulging. However, there is minor overlap among these groups, indicating that it is necessary to exercise caution when using this trait alone to make taxonomic inferences on individual specimens. Early modern humans do not show differences with recent modern humans, and "transitional" individuals such as Jebel Irhoud 1, Maba, and Florisbad, show modern-like frontal squama morphology. The bulging of the frontal squama in modern humans may represent a structural consequence of more general cranial changes, or it could be a response to changes in the morphology of the underlying prefrontal brain elements. A subtle difference between Neandertals and the Afro-European Middle Pleistocene Homo sample is associated with flattening at bregma in the former group, a result that merits further investigation.

The frontal bone in the genus Homo: a survey of functional and phylogenetic sources of variation.

The frontal bone is a useful aspect of the craniofacial skeleton to study in physical anthropology because it contains several characters considered to be important for both population- and species-level distinctions. These include forehead (frontal squama) inclination and supraorbital morphology. Because it lies at the interface between the anterior neurocranium and the upper face, it is also informative about the evolution of both of these regions of the skull. Previous research on frontal bone morphology can be grouped into two broad categories. One set of studies explored the relationship between craniofacial structure and function in an attempt to explain biological sources of variation in the torus development of various extant primate species, including modern humans. The second group of studies examined geographical and temporal patterns of variation in frontal morphology to make inferences about the phylogenetic relationship relationships among fossil hominin populations in the Pleistocene. This paper offers a review of both phylogenetic and functional studies of variation in frontal bone morphology, and synthesizes them to offer a comprehensive understanding of what the frontal bone can tell us about bio-behavioral and evolutionary differences both among extant and extinct members of the genus Homo.

A comparative study of frontal bone morphology among Pleistocene hominin fossil groups.

Features of the frontal bone that are conventionally used to distinguish among fossil hominin groups were quantitatively examined. Fifty-five fossil crania dating from the early to the late Pleistocene were analyzed. Using a modified pantograph, outlines of the frontal bone were collected along the midsagittal and two parasagittal planes. The profile from nasion to bregma, as well as two profiles above the medial and lateral sections of the orbit, respectively, extending from the orbital margin to the coronal suture were traced. The outlines were measured using Elliptical Fourier Function Analysis (EFFA), which enabled a quantification of aspects of the frontal bone that have historically been described primarily in nonmetric or linear terms. Four measurements were obtained: 1) overall morphology as expressed in the Fourier harmonic amplitudes; 2) maximum projection of the supraorbital torus at three points along the browridge (glabella and the medial and lateral aspects of the torus above the orbit); 3) maximum distance of the frontal squama from the frontal chord, capturing forehead curvature; and 4) nasion-bregma chord length. The results indicate that the midsagittal profile is significantly different among all Pleistocene groups in analyses that include both size and shape, as well as size-adjusted data. Homo erectus is significantly different from the late Pleistocene groups (Neandertals and early modern H. sapiens) in glabellar projection. Anatomically modern humans are significantly different from all other groups in both raw and size-standardized analyses of all three outlines that captured overall morphology, as well as forehead curvature and lateral supraorbital torus prominence, and middle Pleistocene Homo are significantly different in both medial and lateral overall parasagittal form. However, for the majority of analyses there were no significant differences among the Pleistocene archaic groups in supraorbital torus projection, frontal squama curvature, nasion-bregma chord length, or overall frontal bone morphology.

Is Central Asia the eastern outpost of the Neandertal range? A reassessment of the Teshik-Tash child.

Since its discovery in southeastern Uzbekistan in 1938, the Teshik-Tash child has been considered a Neandertal. Its affinity is important to studies of Late Pleistocene hominin growth and development as well as interpretations of the Central Asian Middle Paleolithic and the geographic distribution of Neandertals. A close examination of the original Russian monograph reveals the incompleteness of key morphologies associated with the cranial base and face and problems with the reconstruction of the Teshik-Tash cranium, making its Neandertal attribution less certain than previously assumed. This study reassesses the Neandertal status of Teshik-Tash 1 by comparing it to a sample of Neandertal, Middle and Upper Paleolithic modern humans, and recent human sub-adults. Separate examinations of the cranium and mandible are conducted using multinomial logistic regression and discriminant function analysis to assess group membership. Results of the cranial analysis group Teshik-Tash with Upper Paleolithic modern humans when variables are not size-standardized, while results of the mandibular analysis place the specimen with recent modern humans for both raw and size-standardized data. Although these results are influenced by limitations related to the incomplete nature of the comparative sample, they suggest that the morphology of Teshik-Tash 1 as expressed in craniometrics is equivocal. Although, further quantitative studies as well as additional sub-adult fossil finds from this region are needed to ascertain the morphological pattern of this specimen specifically, and Central Asian Middle Paleolithic hominins in general, these results challenge current characterizations of this territory as the eastern boundary of the Neandertal range during the Late Pleistocene.

Patterning of geographic variation in Middle Pleistocene Homo frontal bone morphology.

A quantitative assessment of the frontal bone morphology of a sample of Middle Pleistocene hominins was undertaken in order to address questions regarding their population structure and evolutionary history. Outline tracings of the frontal bones of forty-seven fossil crania were obtained, and size-standardized measurements were then computed using an Elliptical Fourier analysis of these tracings. Principal component scores of the Fourier harmonic amplitudes were derived and served as a quantitative representation of the morphology of the frontal bone. Morphological, geographical, and temporal distance matrices were then constructed between each pair of fossils. A partial Mantel matrix correlation test was performed between morphological and geographical distance matrices, controlling for temporal distance, in order to determine if the pattern of geographical differentiation in features of the frontal bone of mid-Pleistocene Homo followed that of an isolation-by-distance model of population structure. The results of the partial Mantel tests indicate that the overall patterning of differentiation in the features of the frontal bone cannot best be explained by a population structure shaped by isolation-by-distance. Additionally, various aspects of the frontal bone quantified here follow different patterns of geographical differentiation, suggesting that a mosaic pattern of evolution holds true for characters within one cranial region and not just for those between regions.

Impact of character correlation and variable groupings on modern human population tree resolution.

Relationships among modern human populations are often explored through the use of linear measurements taken on the cranium and expressed in the form of dendrograms. However, craniometric variables are strongly correlated and thereby violate the assumption of independence that most statistical analyses require. This study explores the relationship between differing methods of variable treatment and the statistical robustness of the outcomes they yield, as depicted in interpopulational trees of relatedness among modern humans. Three methods of grouping variables are examined. The first method leaves them ungrouped, the second groups variables on the basis of the developmental and/or functional complex of the cranium to which they are thought to belong, and the last method reduces variables by using principal components analysis. The strength of each of these methods is tested through the use of the Continuous Character Maximum Likelihood (CONTML) program in the PHYLIP phylogeny inference package. This program produces output in the form of trees, and the resolution of the branching topology is given as a log-likelihood value, with statistical confidence intervals supporting each branch placement on the tree. The results indicate that leaving variables ungrouped provides misleadingly strong results by failing to account for character correlation. Of the alternative two grouping methods, the covarying components method yields the best-resolved tree with stronger statistical support for its topology than the approach of grouping variables on the basis of their location on the cranium. Finally, the implications for interpreting population histories based on such methods are discussed.

An early modern human from the Pestera cu Oase, Romania.

The 2002 discovery of a robust modern human mandible in the Pestera cu Oase, southwestern Romania, provides evidence of early modern humans in the lower Danubian Corridor. Directly accelerator mass spectrometry radiocarbon (14C)-dated to 34,000-36,000 14C years B.P., the Oase 1 mandible is the oldest definite early modern human specimen in Europe and provides perspectives on the emergence and evolution of early modern humans in the northwestern Old World. The moderately long Oase 1 mandible exhibits a prominent tuber symphyseos and overall proportions that place it close to earlier Upper Paleolithic European specimens. Its symmetrical mandibular incisure, medially placed condyle, small superior medial pterygoid tubercle, mesial mental foramen, and narrow corpus place it closer to early modern humans among Late Pleistocene humans. However, its cross-sectional symphyseal orientation is intermediate between late archaic and early modern humans, the ramus is exceptionally wide, and the molars become progressively larger distally with exceptionally large third molars. The molar crowns lack derived Neandertal features but are otherwise morphologically undiagnostic. However, it has unilateral mandibular foramen lingular bridging, an apparently derived Neandertal feature. It therefore presents a mosaic of archaic, early modern human and possibly Neandertal morphological features, emphasizing both the complex population dynamics of modern human dispersal into Europe and the subsequent morphological evolution of European early modern humans.