Pleistocene Human Fossils Research Articles

Chronology of Late Pleistocene Humans in Eurasia: Results and Perspectives

A compilation of direct age determinations for Late Pleistocene human fossils in eastern Europe and Asia is presented in this paper, and current problems with the dating of hominids in these regions are discussed. Only 25 human finds (4 Neanderthals and 21 modern humans) have been directly dated from Pleistocene eastern Europe and Asia. Indirect dating of human remains (using presumably associated organics) often is insecure, especially when information about the exact provenance of human fossils is lacking. Continuation of direct dating of Late Pleistocene humans in Eurasia, primarily with the help of the accelerator mass spectrometry (AMS) 14C method, is therefore an urgent task.

Comparative analysis of the glabellar region morphology of the late Pleistocene Minatogawa crania: a three-dimensional approach

The Late Pleistocene human fossils recovered from the Minatogawa fissure in Okinawa Island are important in understanding the population history of the Japan archipelago. Based on high-resolution laser scanning, we conducted a three-dimensional (3-D) topographic evaluation of the glabellar and superciliary regions of the Minatogawa and Jomon crania. We aimed to address the question of whether the glabellar morphology of the Minatogawa crania can be seen as a possible ancestral condition of the Jomon people, or whether it more likely represents a distinct morphological pattern. In addition to comparative descriptions and evaluations of the visualized topography, we conducted principal components and cluster analyses using a data set of approximately 240 points located within a confined glabellofrontal area. Results indicate that the male Minatogawa I exhibits a distinct rhomboid to ovoid glabellosuperciliary prominence, in the form of an undifferentiated swollen structure that encompasses both glabellar and superciliary arch regions. Unlike the Jomon condition, this leads to a lack of concave inferolateral contour of the prominence and the virtually complete obliteration of bilateral superciliary expression. The Minatogawa II and IV female crania also exhibit the same morphological pattern at weaker levels. On the other hand, absolute glabellar projection of Minatogawa I is not as great as the developed Jomon examples. This indicates that it is the pattern that is distinctive of Minatogawa, rather than glabellar prominence itself.

Typicality probabilities of Late Pleistocene human fossils from East Asia, Southeast Asia, and Australia: implications for the Jomon population in Japan

The question of the origins of the Jomon population is examined by Campbell’s predictive approach, a method using ‘typicality probability’ which is currently the best way of comparing a single specimen with a reference population. Among the five Pleistocene fossils compared with a Late/Final Jomon sample from the Tohoku district, the analysis based on 13 cranial measurements revealed that Keilor from Australia was more likely a member of the Jomon population than Minatogawa I from Okinawa and Liujiang from southern China. Although, from a chronological viewpoint, it is not possible for Keilor to be an ancestor of the Jomon, these results suggest that the Australian Late Pleistocene population, or their ancestors, should also be considered when seeking the origins of the Jomon.

Evolution of middle-late Pleistocene human cranio-facial form: A 3-D approach

The classification and phylogenetic relationships of the middle Pleistocene human fossil record remains one of the most intractable problems in paleoanthropology. Several authors have noted broad resemblances between European and African fossils from this period, suggesting a single taxon ancestral to both modern humans and Neanderthals. Others point out ‘incipient’ Neanderthal features in the morphology of the European sample and have argued for their inclusion in the Neanderthal lineage exclusively, following a model of accretionary evolution of Neanderthals. We approach these questions using geometric morphometric methods which allow the intuitive visualization and quantification of features previously described qualitatively. We apply these techniques to evaluate proposed cranio-facial ‘incipient’ facial, vault, and basicranial traits in a middle-late Pleistocene European hominin sample when compared to a sample of the same time depth from Africa. Some of the features examined followed the predictions of the accretion model and relate the middle Pleistocene European material to the later Neanderthals. However, although our analysis showed a clear separation between Neanderthals and early/recent modern humans and morphological proximity between European specimens from OIS 7 to 3, it also shows that the European hominins from the first half of the middle Pleistocene still shared most of their cranio-facial architecture with their African contemporaries.

Into Eurasia: A geometric morphometric re-assessment of the Upper Cave (Zhoukoudian) specimens

Since the discovery of the human remains from the Upper Cave of Zhoukoudian in the 1930s there has been speculation over their affinities. In particular, the degree to which the three adult crania exhibit recent East Asian morphology, as well as their degree of within-group variability, has long been debated. Several early researchers described a resemblance to East Asian populations, but these findings have been for the most part rejected by more recent metric and non-metric analyses. Nevertheless, the Upper Cave specimens have not been classified conclusively into any recent modern human population to which they have been compared, and classification results differ for each cranium. Here, the question of the affinities of Upper Cave 101 and 103, the two better-preserved crania, is examined from the perspective of the Late Pleistocene human fossil record using the methodology of 3-D geometric morphometrics. The degree of morphological variation between the two specimens is also evaluated within the context of recent population variability. Neurocranial and facial morphology are analyzed separately so as to maximize comparative samples. Results show a morphological resemblance of the Upper Cave material to Upper Paleolithic Europeans. It is proposed that the Upper Cave specimens retain important aspects of modern human ancestral morphology, and possibly share a recent common ancestral population with Upper Paleolithic Europeans, in accordance with the Single Origin model of modern human origins.

Huanglong Cave: A Late Pleistocene human fossil site in Hubei Province, China

This contribution discusses recent paleoanthropological findings from Huanglong Cave, a Late Pleistocene human fossil site from Yunxi County, Hubei Province, China. Three excavations in the Huanglong Cave from 2004 to 2006 yielded seven human teeth, some stone and bone tools, possible burnt sediment and other evidence possibly related to hominin activities. Based on the presence of extinct faunas (20% of total taxa identified), the deposits dated to the Late Pleistocene. Electron spin resonance (ESR) and uranium-series (U-series) dating analyses on associated teeth and speleothems have resulted in divergent chronometric ages (ESR: 44–34 ka; U-series: 103–79 ka). Analysis indicates: (1) most of the morphological and metric features of the human teeth from Huanglong Cave fall within the range of variation of modern Chinese, but a few characters may still link them to more archaic hominins; (2) some activity-induced abrasion and other tooth use-marks were identified, including pronounced tooth chipping and interproximal grooves on the anterior teeth; (3) the sample of blackened deposit has a high carbon content (over 70%), experienced high temperatures, and likely was of cultural origin and not natural; (4) the mammal fossils represent the “Ailuropoda-Stegodon” faunal unit which lived in southern China throughout the Pleistocene. Synthesizing all of these findings, especially the human teeth that display modern human characteristics, Huanglong Cave will offer some new insights into various issues currently being debated in Late Pleistocene human evolutionary research.

Evidence of fire use of late Pleistocene humans from the Huanglong Cave, Hubei Province, China

Since 2004, three excavations have been carried out at a late Pleistocene human fossil site of Huanglong Cave in Yunxi County, Hubei Province of China, which unearthed seven human teeth, dozens of stone tools, mammal fossils and other evidence indicating human activities. During the third excavation in 2006, in the same layer as the human teeth, we found some patches of black materials embedded in the deposit. We doubted that this black deposit layer is the remains of burning or even human use of fire at the cave. To further explore the possibility of human fire use at the Huanglong Cave, we examined samples directly taken from the black deposit layer and compared them with samples taken from several places in the cave using three methods: micromorphology, element content determination and deposit temperature analysis. Our results indicate that the contents of carbon element in the black deposit reach 64.59%–73-29%. In contrast, contents of carbon element of the comparative samples from other parts in the cave are only 5.82%–9.49%. The micromorphology analysis of the black deposit samples reveals a plant structure like axial parenchyma, fibrocyte, uniseriate ray and vessel. High-temperature magnetic susceptibility measurements suggest that the stratum possibly underwent a high temperature in the nature. Based on these lab analyses, we are sure that the black layer in the Huanglong Cave is the remains of fire and combustion did occur in the cave 100000 years ago. Taking other evidence of human activities found in the Huanglong Cave into consideration, we believe that the evidence of fire from the Huanglong Cave was caused by the human activities of controlled use of fire.

The brain morphology of Homo Liujiang cranium fossil by three-dimensional computed tomography

The Liujiang cranium is the most complete and well-preserved late Pleistocene human fossils ever unearthed in south China. Because the endocranial cavity is filled with hard stone matrix, earlier studies focused only on the exterior morphology of the specimen using the traditional methods. In order to derive more information for the phyletic evaluation of the Liujiang cranium, high-resolution industrial computed tomography (CT) was used to scan the fossil, and the three-dimensional (3D) brain image was reconstructed. Compared with the endocasts of the hominin fossils (Hexian, Zhoukoudian, KNM-WT 15000, Sm 3, Kabwe, Brunn 3, Predmost) and modern Chinese, most morphological features of the Liujiang brain are in common with modern humans, including a round brain shape, bulged and wide frontal lobes, an enlarged brain height, a full orbital margin and long parietal lobes. A few differences exist between Liujiang and the modern Chinese in our sample, including a strong posterior projection of the occipital lobes, and a reduced cerebellar lobe. The measurement of the virtual endocast shows that the endocranial capacity of Liujiang is 1567 cc, which is in the range of Late Homo sapiens and much beyond the mean of modern humans. The brain morphology of Liujiang is assigned to Late Homo sapiens.

Late Pleistocene Human Skull from Hofmeyr, South Africa, and Modern Human Origins

The lack of Late Pleistocene human fossils from sub-Saharan Africa has limited paleontological testing of competing models of recent human evolution. We have dated a skull from Hofmeyr, South Africa, to 36.2 +/- 3.3 thousand years ago through a combination of optically stimulated luminescence and uranium-series dating methods. The skull is morphologically modern overall but displays some archaic features. Its strongest morphometric affinities are with Upper Paleolithic (UP) Eurasians rather than recent, geographically proximate people. The Hofmeyr cranium is consistent with the hypothesis that UP Eurasians descended from a population that emigrated from sub-Saharan Africa in the Late Pleistocene.

Huanglong cave, a new late pleistocene hominid site in Hubei Province, China

For the past 20 years the modern human origins debate has received a significant amount of attention in paleoanthropological research. Primarily supported by the evidence of earlier dates of anatomically modern human fossils and genetic studies, the “Out of Africa” hypothesis is based on the belief that the ancestor of all modern humans, including modern Chinese, came from Africa. The opposite hypothesis “Mutiregional evolution” proposes that continuous evolution occurred on a regional scale, for which human paleontology offers strong support. However, due to the paucity of hominid fossils in China between 100 and 50 ka, support to the latter hypothesis is currently weak. This is a report here of five human fossil teeth, and associated stone tools and mammal fossils from a newly discovered cave site, Huanglong Cave, located in Yunxi County, Hubei Province, China. Preliminary studies indicate: (1) the morphological features of the human fossils resemble those of late Pleistocene human fossils from China; (2) the stone tools display patterns of both the southern and northern Paleolithic cultures of China; (3) the mammal fossils represent the “Ailuropoda-Stegodon” faunal unit which lived in southern China throughout the Pleistocene. ESR and U-series dating on animal teeth and a stalagmite derived from the same layer as the human teeth indicate two possible ages: 103±1.6 ka and 44±12.5 ka. In addition to other evidence presented here, it is believed that hominid occupation of the cave was likely around 100 ka. If this age is further substantiated, Huanglong Cave will be the first late Pleistocene hominid fossil site in China where anatomically modern humans lived about 100 ka. The human fossils and other related materials from Huanglong Cave will provide important information for research on the origin of modern Chinese.

Comparaison de l'expression de certains caractères crâniens sur les hominidés chinois du Pléistocène récent et de l'Holocène (grotte supérieure de Zhoukoudian, sites de Longxian et de Yanqing)

The human remains recovered from “Grotte supérieure de Zhoukoudian” are the best-preserved Late Pleistocene human fossils in East Asia. For decades, as the representative of the Late Pleistocene human in East Asia, the Upper Cave skulls have been playing important role in the research of origins of modern Mongoloids and American Indians. With the advance of the origin and evolution of modern humans, more attention has been paid to the details and the mechanisms for the late Pleistocene human evolution and the formation of modern human populations. Both the origin and diversification of modern humans have been stressed. Some studies further trigger the debaters on the Upper Cave Man concerning its evolutionary level and its role in the formation of modern human populations in East Asia. To further explore these problems, we examined and compared 12 non-metric features on the 3 Late Pleistocene Upper Cave skulls and 162 Holocene individuals earthed from two archaeological sites of North China (Longxian and Yanqing). Our results indicate that 8 on the 12 features have different expression patterns between Upper Cave Man and recent Chinese leading the authors to believe that more primitive expressions appeared on the Upper Cave Man than on recent Chinese populations. Based on these findings, some problems on the intragroup variation in Late Pleistocene and Holocene populations are discussed.

Study of a temporal bone of Homo heildelbergensis

ConclusionsThe characteristic features of the Hh specimen conformed to those of other Pleistocene human fossils, indicating strong cranial structures and a heavy mandible. The mastoid was large and suggested a powerful sternocleidomastoid muscle. The inner ear and tympanic cavities were similar in size and orientation, suggesting that their functions were probably similar. Our observations suggest that the left ear of this Hh specimen was healthy. The large canaliculo–fenestral angle confirms that this ancestor was bipedal. It also strongly suggests that Hh individuals were predisposed to develop certain pathologies of the labyrinth capsule associated with bipedalism, in particular otosclerosis.ObjectiveWe studied a temporal bone of Homo heidelbergensis (Hh) in order to investigate the clinical and physiological implications of certain morphological features, especially those associated with the evolutionary reorganization of the inner ear.Material and methodsThe bone, found in a breach of a cave near Málaga in southern Spain, together with Middle–Upper Pleistocene faunal remains, is >300 000 years old. Four analytical methods were employed. A 3D high-resolution surface laser scan was used for anatomical measurements. For the sectional analysis of the middle and inner ears of Hh we used high-resolution CT, simultaneously studying a normal temporal bone from Homo sapiens sapiens (Hss). To study the middle and inner ear spaces we used 3D reconstruction CT preceded by an intra-bone air shielding technique. To examine the tympanic cavities and measure the canaliculo–fenestral angle, we used a special minimally invasive endoscopic procedure.ResultsThe surface, sectional and 3D CT examinations showed that the Hh specimen was generally more robust and larger than the Hss specimen. It had a large glenoid fossa. The external meatus was wide and deep. The middle ear, and especially the mastoid, was large and widely pneumatized. There were no appreciable differences in the position and size of the labyrinthine spaces and tympanic cavity. The dimensions of the semicircular canals were similar to those of the Hss specimen. Endoscopy revealed normal, healthy tympanic walls and an ossicle fragment in the atticum that probably belonged to the body of the malleus. The diameters of the fallopian duct and the tympanic opening of the Eustachian tube were large. The canaliculo–fenestral angle was ≈114°.

A preliminary report on the newly found Tianyuan Cave, a Late Pleistocene human fossil site near Zhoukoudian

The Tianyuan Cave is the only human fossil-bearing site containing rich mammalian fossils found in the last decades near Zhoukoudian. Up to now more than 34 specimens of the human body have been recovered, and the mammalian fossils can be put into 29 species. Cervids dominate the fauna, and carnivores are very rare. Based on the primary examination, the human fossils can be attributed to the speciesHomo sapiens. All the mammalian fossils, except one between tooth ofCrocuta belong to the species that still exist today. But some of them are the first records in fossil form north of the Yellow River, such asArctonyx andCapricornis. Based on the mammalian fauna study, it seems that the Tianyuan Cave can be correlated with the Upper Cave. Sixty-three percent of the species of the mammalian fauna from the Tianyuan Cave are also present in the Upper Cave. The characters of the deposits also share some similarities between the Tianyuan Cave and the Upper Cave; both of them are mainly composed of breccia without cement. The dating using the U-series method on deer tooth samples indicates that the geological age of the new site is around 25 thousand years B.P. This is the first discovery of human sites outside the core area of the Peking Man Site at Zhoukoudian, which throws new light onto this world famous site complex.

A preliminary report on the newly found Tianyuan Cave, a Late Pleistocene human fossil site near Zhoukoudian

A preliminary report on the newly found Tianyuan Cave, a Late Pleistocene human fossil site near Zhoukoudian

On the origin of modern humans in China

The Multiregional Evolution Hypothesis (MEH) and the Recent Out of Africa Hypothesis (ROAH) are two main hypotheses bearing on the origin of modern humans. In China, there are many common morphological features among Pleistocene human fossil skulls. These features and the morphological mosaic between Homo sapiens erectus and Homo sapiens sapiens of China indicate the continuity of human evolution in this region. There are a few skulls possessing one or two features that suggest gene flow from the West to China. Based on the evidence of continuity and gene flow, a new hypothesis, Continuity with Hybridization, was proposed in 1998 for characterizing human evolution in China. This hypothesis is also supported by the Paleolithic archaeological record of China. There are serious problems with the ROAH. For example, the dates of the last common ancestor of anatomically modern humans obtained, determined by different investigators using molecular clock estimates, vary considerably. Further, the important assumptions of the ROAH—a constant mutation rate and no loss of gene variants during human evolution—cannot be verified. The results of recent studies on ancient DNA could be interpreted in different ways, and could also be interpreted as supporting the MEH. Also, the Paleolithic artifacts from the Near East and China do not support the ROAH.

Isolation and the origin of the khoisan: Late pleistocene and early holocene human evolution at the southern end of Africa

The debate between the proponents of the Recent African Origin and the Multiregional Evolution models for the origin of anatomically modern humans has not been of great importance to the interpretation of late Pleistocene human fossils in South Africa. The main reason is that both models propose that evidence of anatomically moderns should happen in South Africa at an early date. A more important issue to the African context is whether or not those early transitions to modernity were accompanied by the development of the distinctive local populations known today as the Khoisan. Serogenetic evidence suggests a relative antiquity for the origins of living Khoisan peoples, but this may simply reflect the longer time that anatomically modern peoples have lived in the sub-continent. A search for features of distinctive Khoisan morphology in the early remains may be of more value as it might indicate the presence of a local adaptive pattern to the special environment conditions of the region.

The Atapuerca human fossils

After 20 years of research, the Atapuerca sites have provided a large amount of archaeological and palaeontological remains. Human fossils have been found in three sites: Gran Dolina, galeria and Sima de los Huesos. The Early Pleistocene human fossils from Gran Dolina have been ascribed to a new species,Homo antecessor, that represent the last common ancestor of Neandertals and modern humans. The Sima de los Huesos fossils and all the European Middle Pleistocene human fossils are the ancestors exclusively of the Neandertals, which evolved in Europe in conditions of geographic and genetic isolation.

The human cranial remains from Gran Dolina Lower Pleistocene site (Sierra de Atapuerca, Spain)

In this article we study the cranial remains of the late Lower Pleistocene human fossils from Gran Dolina (Sierra de Atapuerca, Spain), assigned to the new species Homo antecessor. The cranial remains belong to at least five individuals, both juveniles and adults. The most outstanding feature is the totally modern human morphology of the very complete face ATD6-69, representing the earliest occurrence of the modern face in the fossil record. The Gran Dolina fossils show in the face a suite of modern human apomorphies not found in earlier hominids nor in contemporary or earlier Homo erectus fossils. There are also traits in the Gran Dolina fossils shared with both Neandertals and modern humans, which reinforce the hypothesis that Neandertals and modern humans form a clade, and that the Gran Dolina fossils are a common ancestor to both lineages.

A modern human pattern of dental development in lower pleistocene hominids from Atapuerca-TD6 (Spain).

The study of life history evolution in hominids is crucial for the discernment of when and why humans have acquired our unique maturational pattern. Because the development of dentition is critically integrated into the life cycle in mammals, the determination of the time and pattern of dental development represents an appropriate method to infer changes in life history variables that occurred during hominid evolution. Here we present evidence derived from Lower Pleistocene human fossil remains recovered from the TD6 level (Aurora stratum) of the Gran Dolina site in the Sierra de Atapuerca, northern Spain. These hominids present a pattern of development similar to that of Homo sapiens, although some aspects (e.g., delayed M3 calcification) are not as derived as that of European populations and people of European origin. This evidence, taken together with the present knowledge of cranial capacity of these and other late Early Pleistocene hominids, supports the view that as early as 0.8 Ma at least one Homo species shared with modern humans a prolonged pattern of maturation.

Minatogawa Hominid Fossils and the Evolution of Late Pleistocene Humans in East Asia.

In order to understand the origins of Japanese people, this paper deals with the skull morphology of Late Pleistocene human fossils in Eastern Asia. After the Minatogawa 1 skull, the most complete Japanese Pleistocene fossil human skull now available, is described, a functional interpretation of the evidence is presented. The Minatogawa 1 skull exhibits many archaic features, which mainly relate to masticatory adaptation. When the Minatogawa 1 skull is compared with other fossil human skulls in East and Southeast Asia, it is found to exhibit morphological features slightly closer to the Wadjak 1 skull in Java than to the Zhoukoudian Upper Cave and Liujiang skulls in China. This suggests that there were no major human migrations between China and Japanese Islands in Late Pleistocene.