Pleistocene Hominins Research Articles

The genome of the offspring of a Neanderthal mother and a Denisovan father.

Neandertals and Denisovans are extinct groups of hominins that separated from each other more than 390,000 years ago1,2. Here we present the genome of “Denisova 11”, a bone fragment from Denisova Cave (Russia)3, and show that it comes from an individual who had a Neandertal mother and a Denisovan father. The father, whose genome bears traces of Neandertal ancestry, came from a population related to a later Denisovan found in the cave4–6. The mother came from a population more closely related to Neandertals who lived later in Europe2,7 than to an older Neandertal found in Denisova Cave8, suggesting that migrations of Neandertals between eastern and western Eurasia occurred sometime after ~120,000 years ago. The finding of a first-generation Neandertal-Denisovan offspring among the small number of archaic specimens sequenced to date suggests that mixing between Late Pleistocene hominin groups was common when they met.

Ribcage measurements indicate greater lung capacity in Neanderthals and Lower Pleistocene hominins compared to modern humans

Our most recent fossil relatives, the Neanderthals, had a large brain and a very heavy body compared to modern humans. This type of body requires high levels of energetic intake. While food (meat and fat consumption) is a source of energy, oxygen via respiration is also necessary for metabolism. We would therefore expect Neanderthals to have large respiratory capacities. Here we estimate the pulmonary capacities of Neanderthals, based on costal measurements and physiological data from a modern human comparative sample. The Kebara 2 male had a lung volume of about 9.04 l; Tabun C1, a female individual, a lung volume of 5.85 l; and a Neanderthal from the El Sidrón site, a lung volume of 9.03 l. These volumes are approximately 20% greater than the corresponding volumes of modern humans of the same body size and sex. These results show that the Neanderthal body was highly sensitive to energy supply.

The top of the Gran Dolina (Atapuerca, Spain) sequence: A zooarchaeological and occupational perspective

Middle Pleistocene unit TD10 of the Gran Dolina site is nearly four metres thick and is divided into four subunits (TD10.1, TD10.2, TD10.3, TD10.4). To date, the upper two subunits (TD10.1 and TD10.2) have been completely excavated and have been studied from zooarchaeological, taphonomic and occupational perspectives. The top of the sequence (Upper TD10.1), however, has not undergone these types of studies until now. In this paper we report the results of our analyses of the anatomical profiles, age, and the anthropogenic and carnivore-induced modifications in this assemblage. Methods employed to evaluate sequential scenarios (carnivore to hominin; hominin to carnivore; carnivore to hominin to carnivore) have led to contradictory results. We conclude that the formation of Upper TD10.1 is the product of the overlap of independent events (hominin only and carnivore only), with limited commensalism between the two agents. The type of accumulation is consistent with the characteristics of an accumulative palimpsest generated by different actors. Unlike those documented in the lower levels of TD10 (TD10.1BB and TD10.2BB), hominin occupations in this part of the sequence were very brief. This scenario completes the picture of the types of occupations that took place during the end of Middle Pleistocene at Gran Dolina. In short, level TD10 was the site of three types of occupation by Middle Pleistocene hominins: a kill/butchering site in TD10.2BB, a long-term residential camp in TD10.1BB, and finally, logistical and short-term occupations in Upper TD10.1.

Lithic artifacts and bone tools from the Lower Palaeolithic site Marathousa 1, Megalopolis, Greece: Preliminary results

The technological systems and subsistence strategies of Middle Pleistocene hominins in South-East Europe are insufficiently understood due to the scarcity of well-preserved, excavated assemblages. In this paper, we present first results from the study of the lithic and bone artifacts unearthed at the Lower Palaeolithic site Marathousa 1 (MAR-1), Megalopolis, Greece. The context of the site represents a depositional environment close to a lakeshore, where rapid burial in a very fine-grained matrix ensured extraordinary conditions for preservation. Lithic artifacts occur in spatial and stratigraphic association with remains of the elephant Palaeoloxodon antiquus as well as other mammals. Bones, including those of elephants, show clear anthropogenic flaking scars, cut-marks and fracture patterns indicating deliberate breakage and modification by early humans. The MAR-1 lithic assemblage is composed of small-sized debitage, retouched tools, a few small and exhausted cores, as well as a large number of debris and retouch products, such as chips and resharpening flakes. Currently, there are no indications of Acheulean bifacial debitage, large cutting tools are missing, and a key aspect of the material refers to its 'microlithic' character. The scarcity of cores and primary flakes indicates a fragmented reduction sequence and complex discard patterns that require further investigation. On the basis of the on-going analysis of lithic material from three field seasons, we discuss aspects of assemblage composition and the role of raw material types, the main technological and typological traits of the industry, as well as the potential contribution of the MAR-1 assemblage in broader discussions about Middle Pleistocene lithic techno-complexes and subsistence strategies in Eurasia. Finally, we briefly present a small sample of bone artifacts, which suggest that hominin exploitation of the animal carcasses was not restricted to marrow extraction and bone processing for nutritional needs, but included also the knapping of bones, potentially with the aim of using the knapped products as tools. The MAR-1 archaeological record compares well with other important Eurasian sites yielding ‘small tool assemblages’, such as Ficoncella, La Polledrara, Bilzingsleben, Schöningen and Vértesszőlős, some of which, like MAR-1, have provided evidence of elephant or other mega-fauna exploitation.

Palaeolithic occupation of the Anatolian High Plateau during a cold period: An MIS 6 aged artifact from the Avlamış Valley, Eskişehir, NW Turkey

AbstractIn the Avlamış Valley, 10 km north of Eskişehir (NW Anatolia), an undamaged triangular Levallois flake was encountered in a paleosol, 430 cm beneath the ground surface. The artifact has a minimal dorsal retouch on the right margin, and there is a light beige partial patina on the ventral surface. Based on the technological investigations, this artifact was made using the centripetal recurrent Levallois technique. An observation of the trench walls revealed the triple nature of the stratigraphy: an upper gravelly sand (Unit‐1), an underlying reddish‐brown pebbly paleosol (Unit‐2), and a lowermost dominantly pink, finer‐grained deposit (Unit‐3) where the artifact was retrieved. Optically stimulated luminescence ages indicate a strong influence of global marine isotopic stages (MIS) on the trench stratigraphy, with the deposit hosting the artifact dating to 148 ± 20 ka (MIS 6 cold period). The available pollen data from the same stratigraphic level verified an open steppe landscape with some arboreal plant cover during deposition. This is the first stratigraphically dated Middle Palaeolithic artifact from NW Anatolia, and one of the few in the whole country, thus igniting further discussion about the ways Pleistocene hominins adapted to cold and arid environmental conditions.

Metric and morphological comparison between the Arago (France) and Atapuerca-Sima de los Huesos (Spain) dental samples, and the origin of Neanderthals

The variability observed in the growing Middle Pleistocene hominin fossil record of Europe continues to trigger much debate on taxonomic issues and the biological processes that gave rise to Neanderthals. Here we present a metric and morphological comparative study of the dental samples recovered from the sites of Arago (southeast France) and Sima de los Huesos (SH) in the Sierra de Atapuerca (northern Spain). These sites are key to providing answers to these questions since they have yielded the largest hominin samples so far recovered for this time period. Despite the geographical proximity of the two sites and the contemporaneity of their hominin assemblages, we have observed remarkable metric and morphological differences between the teeth at Arago and SH. Whereas the SH teeth present an almost morphological identity with European Neanderthals, the Arago teeth exhibit a combination of plesiomorphic as well as some Neanderthal-derived features. In addition, the Arago crown dimensions are remarkably larger than those from SH, the differences being statistically significant for most variables. We hypothesize that during the Middle Pleistocene the European continent was settled at different points in time by hominin groups coming from Southwest Asia, probably from a common mother population evolving in this latter region. These first settlers can be identified by their more plesiomorphic morphology, whereas the most recent settlers are closer in appearance to Neanderthals. In addition, genetic processes such as isolation, genetic drift, directional adaptation or hybridization would have given rise to the puzzle we observe in the current fossil record.

A reassessment of the early archaeological record at Leang Burung 2, a Late Pleistocene rock-shelter site on the Indonesian island of Sulawesi.

This paper presents a reassessment of the archaeological record at Leang Burung 2, a key early human occupation site in the Late Pleistocene of Southeast Asia. Excavated originally by Ian Glover in 1975, this limestone rock-shelter in the Maros karsts of Sulawesi, Indonesia, has long held significance in our understanding of early human dispersals into ‘Wallacea’, the vast zone of oceanic islands between continental Asia and Australia. We present new stratigraphic information and dating evidence from Leang Burung 2 collected during the course of our excavations at this site in 2007 and 2011–13. Our findings suggest that the classic Late Pleistocene modern human occupation sequence identified previously at Leang Burung 2, and proposed to span around 31,000 to 19,000 conventional 14C years BP (~35–24 ka cal BP), may actually represent an amalgam of reworked archaeological materials. Sources for cultural materials of mixed ages comprise breccias from the rear wall of the rock-shelter–remnants of older, eroded deposits dated to 35–23 ka cal BP–and cultural remains of early Holocene antiquity. Below the upper levels affected by the mass loss of Late Pleistocene deposits, our deep-trench excavations uncovered evidence for an earlier hominin presence at the site. These findings include fossils of now-extinct proboscideans and other ‘megafauna’ in stratified context, as well as a cobble-based stone artifact technology comparable to that produced by late Middle Pleistocene hominins elsewhere on Sulawesi.

Supraorbital morphology and social dynamics in human evolution

Uniquely, with respect to Middle Pleistocene hominins, anatomically modern humans do not possess marked browridges, and have a more vertical forehead with mobile eyebrows that play a key role in social signalling and communication. The presence and variability of browridges in archaic Homo species and their absence in ourselves have led to debate concerning their morphogenesis and function, with two main hypotheses being put forward: that browridge morphology is the result of the spatial relationship between the orbits and the brain case; and that browridge morphology is significantly impacted by biting mechanics. Here, we virtually manipulate the browridge morphology of an archaic hominin (Kabwe 1), showing that it is much larger than the minimum required to fulfil spatial demands and that browridge size has little impact on mechanical performance during biting. As browridge morphology in this fossil is not driven by spatial and mechanical requirements alone, the role of the supraorbital region in social communication is a potentially significant factor. We propose that conversion of the large browridges of our immediate ancestors to a more vertical frontal bone in modern humans allowed highly mobile eyebrows to display subtle affiliative emotions.

Reassessment of Anthropic Modifications on the Early Pleistocene Hominin Specimen Stw53 (Sterkfontein, South Africa)

Linear marks were observed on the maxillary zygomatic process of the early Pleistocene hominin fossil Stw53 from the Sterkfontein cave site. The taxonomic allocation of this specimen is still under debate, as it has been attributed either to the genusHomoorAustralopithecus. Two alternative hypotheses have been proposed regarding the taphonomic agent responsible for the linear marks observed on this specimen. These modifications were initially assigned to anthropic cutmarks inflicted by stone tools, thus representing the earliest cutmarks discovered on hominin remains. Conversely, these linear marks were interpreted by others as non-anthropic modifications. In order to better understand the origin of the linear marks, a new taphonomic appraisal of these modifications was conducted using a multivariate statistical approach complemented by butchery and trampling experiments on extant mammal bones. This new study of Stw53 linear marks does not provide evidence for anthropic activities but, rather, supports the unintentional origin hypothesis.

Timing of the Saalian- and Elsterian glacial cycles and the implications for Middle \u2013 Pleistocene hominin presence in central Europe

By establishing a luminescence-based chronology for fluvial deposits preserved between the Elsterian- and Saalian tills in central Germany, we obtained information on the timing of both the Middle Pleistocene glacial cycles and early human appearance in central Europe. The luminescence ages illustrate different climatic driven fluvial aggradation periods during the Saalian glacial cycle spanning from 400–150 ka. The ages of sediments directly overlying the Elsterian till are approximately 400 ka and prove that the first extensive Fennoscandian ice sheet extension during the Quaternary correlates with MIS 12 and not with MIS 10. Furthermore, the 400 ka old fluvial units contain Lower Paleolithic stone artefacts that document the first human appearance in the region. In addition, we demonstrate that early MIS 8 is a potential date for the onset of the Middle Paleolithic in central Germany, as Middle Paleolithic stone artefacts are correlated with fluvial units deposited between 300 ka and 200 ka. However, the bulk of Middle Paleolithic sites date to MIS 7 in the region. The fluvial units preserved directly under the till of the southernmost Saalian ice yield an age of about 150 ka, and enable a correlation of the Drenthe stage to late MIS 6.

Hominin hand bone fossils from Sterkfontein Caves, South Africa (1998–2003 excavations)

We describe eleven hominin metacarpals and phalanges recovered from Jacovec Cavern and Member 4 of the Sterkfontein Formation between 1998 and 2003. Collectively, the fossils date in excess of 2.0 Ma, and are probably attributable to Australopithecus africanus and/or Australopithecus prometheus. When combined with results of previous studies on Australopithecus postcranial functional morphology, the new data presented here suggest that at least some late Pliocene and/or early Pleistocene hominins from Sterkfontein were arboreally adept. This finding accords with the reconstruction of the site's >2.0 Ma catchment area as well-vegetated and containing significant woody components. In addition, most of the new specimens described here evince morphologies that indicate the hands from which they derived lacked complete modern humanlike manual dexterity, which is integral to the manufacture and use of intentionally shaped stone tools. The absence of lithic artifacts from both stratigraphic units from which the fossils were excavated is consistent with this conclusion.

Hominin distribution and density patterns in Pleistocene China: Climatic influences

At the broad Early/Middle/Late Pleistocene level, the Chinese hominin fossil site distribution data may inform on hominin mobility/behavioral/interaction strategies. Hominins during the Early Pleistocene and well into the Middle Pleistocene appear to be more restricted to central and southern China. By the advent of the late Middle Pleistocene hominins are found regularly in northern China and by the Late Pleistocene, hominin fossils are found in all latitudes and longitudes encompassing modern day China, and beginning some time during marine isotope stage 3 appear in higher altitudinal regions (e.g., Qinghai-Tibetan Plateau) as well. During the Middle Pleistocene a number of sites are found well away from major riverways suggesting an ability to transport potable water or perhaps an ability to be able to locate smaller bodies of fresh water. Given the paucity of evidence for storage containers until relatively recently in time, it is perhaps not that surprising that many of the Middle Pleistocene open-air sites are reportedly found in or near lacustrine environments. Hominins had a more restricted range due to climatic variation during the Early and early Middle Pleistocene, but were able to more successfully adapt to changing climes beginning during the late Middle Pleistocene. In part, this is due to a range of behavioral changes that hominins developed during the Pleistocene.

A New Pleistocene Hominin Tracksite from the Cape South Coast, South Africa

A Late Pleistocene hominin tracksite has been identified in coastal aeolianite rocks on the Cape south coast of South Africa, an area of great significance for the emergence of modern humans. The tracks are in the form of natural casts and occur on the ceiling and side walls of a ten-metre long cave. Preservation of tracks is of variable quality. Up to forty hominin tracks are evident. Up to thirty-five hominin tracks occur on a single bedding plane, with potential for the exposure of further tracks. Five tracks are apparent on a second hominin track-bearing bedding plane. A number of individuals made the tracks while moving down a dune surface. A geological investigation at the site and stratigraphic comparison to published geochronological studies from this area suggest that the tracks are ~90 ka in age. If this is the case, the shoreline at the time would have been approximately 2 km distant. This is the first reported hominin tracksite from this time period. It adds to the relatively sparse global record of early hominin tracks, and represents the largest and best preserved archive of Late Pleistocene hominin tracks found to date. The tracks were probably made by Homo sapiens.

Coupled ESR and U-series dating of Middle Pleistocene hominin site Bailongdong cave, China

Abstract The fossil evidence of hominins in China is crucial to understand the human evolution and dispersal in Eurasia. However, the dating of Chinese Early and Middle Pleistocene hominin record remains a serious problem. In this study, we applied coupled ESR/U-series dating method on Bailongdong hominin cave site in Hubei Province, central China, which is a key area to study early human migration between South and North China. The U-series analyses show that the fossil teeth samples from three horizons are close or beyond equilibrium, which indicate very early uptake or potential loss of uranium. Single saturation exponential fitting of 10 dose points irradiated up to 4 kGy was used for DE determination except one by double saturation exponential fitting with 16 dose points up to 63 kGy. Calculated by two uranium migration models, we obtain a weighted mean age of 509 ± 16 ka for five fossil teeth from layer 2. Combined with cosmogenic nuclides 26Al/10Be burial dating carried out on the cave deposits beneath the fossil layers in previous study, the multi-methods dating study places Bailongdong site at the early stage of Middle Pleistocene, and pinpoints the deposition of the fossil remains to the time of marine isotope stages 13. This is in agreement with the fauna composition and sedimentary characteristics which show a subtropical-tropical warm forest-grass environment. The multi-methods dating of Bailongdong site by both cosmogenic and coupled ESR/U-series methods is a good example for the future chronological studies of early human sites in China.

The fossil teeth of the Peking Man

This study provides new original data, including the endostructure of most Zhoukoudian H. erectus teeth preserved to date, since the publication of Black in 1927 and Weidenreich in 1937. The new evidence ratifies the similarities of Zhoukoudian with other East Asian mid-Middle Pleistocene hominins such as Hexian and Yiyuan, and allows defining a dental pattern potentially characteristic of this population commonly referred to as classic H. erectus. Given the possible chronological overlaps of classic H. erectus with other archaic Homo, the characterization of this group becomes a key issue when deciphering the taxonomy and evolutionary scenario of the Middle Pleistocene hominins in East Asia. Internally, the most remarkable feature of Zhoukoudian teeth is the highly crenulated enamel-dentine junction (EDJ) and its imprint on the roof of the pulp cavity. So far, this “dendrite-like” EDJ has been found only in East Asia Middle Pleistocene hominins although a large group of samples were assessed, and it could be useful to dentally define classic H. erectus in China. The crenulated EDJ surface, together with the stout roots and the taurodontism could be a mechanism to withstand high biomechanical demand despite a general dentognathic reduction, particularly of the crowns, in these populations.

Shedding light on the Early Pleistocene of TD6 (Gran Dolina, Atapuerca, Spain): The technological sequence and occupational inferences.

This paper aims to update the information available on the lithic assemblage from the entire sequence of TD6 now that the most recent excavations have been completed, and to explore possible changes in both occupational patterns and technological strategies evidenced in the unit. This is the first study to analyse the entire TD6 sequence, including subunits TD6.3 and TD6.1, which have never been studied, along with the better-known TD6.2 Homo antecessor-bearing subunit. We also present an analysis of several lithic refits found in TD6, as well as certain technical features that may help characterise the hominin occupations. The archaeo-palaeontological record from TD6 consists of 9,452 faunal remains, 443 coprolites, 1,046 lithic pieces, 170 hominin remains and 91 Celtis seeds. The characteristics of this record seem to indicate two main stages of occupation. In the oldest subunit, TD6.3, the lithic assemblage points to the light and limited hominin occupation of the cave, which does, however, grow over the course of the level. In contrast, the lithic assemblages from TD6.2 and TD6.1 are rich and varied, which may reflect Gran Dolina cave’s establishment as a landmark in the region. Despite the occupational differences between the lowermost subunit and the rest of the deposit, technologically the TD6 lithic assemblage is extremely homogeneous throughout. In addition, the composition and spatial distribution of the 12 groups of lithic refits found in unit TD6, as well as the in situ nature of the assemblage demonstrate the high degree of preservation at the site. This may help clarify the nature of the Early Pleistocene hominin occupations of TD6, and raise reasonable doubt about the latest interpretations that support the ex situ character of the assemblage as a whole.

Morphology and structure of Homo erectus humeri from Zhoukoudian, Locality 1.

BackgroundRegional diversity in the morphology of the H. erectus postcranium is not broadly documented, in part, because of the paucity of Asian sites preserving postcranial fossils. Yet, such an understanding of the initial hominin taxon to spread throughout multiple regions of the world is fundamental to documenting the adaptive responses to selective forces operating during this period of human evolution.MethodsThe current study reports the first humeral rigidity and strength properties of East Asian H. erectus and places its diaphyseal robusticity into broader regional and temporal contexts. We estimate true cross-sectional properties of Zhoukoudian Humerus II and quantify new diaphyseal properties of Humerus III using high resolution computed tomography. Comparative data for African H. erectus and Eurasian Late Pleistocene H. sapiens were assembled, and new data were generated from two modern Chinese populations.ResultsDifferences between East Asian and African H. erectus were inconsistently expressed in humeral cortical thickness. In contrast, East Asian H. erectus appears to exhibit greater humeral robusticity compared to African H. erectus when standardizing diaphyseal properties by the product of estimated body mass and humeral length. East Asian H. erectus humeri typically differed less in standardized properties from those of side-matched Late Pleistocene hominins (e.g., Neanderthals and more recent Upper Paleolithic modern humans) than did African H. erectus, and often fell in the lower range of Late Pleistocene humeral rigidity or strength properties.DiscussionQuantitative comparisons indicate that regional variability in humeral midshaft robusticity may characterize H. erectus to a greater extent than presently recognized. This may suggest a temporal difference within H. erectus, or possibly different ecogeographical trends and/or upper limb loading patterns across the taxon. Both discovery and analysis of more adult H. erectus humeri are critical to further evaluating and potentially distinguishing between these possibilities.

Lower limb articular scaling and body mass estimation in Pliocene and Pleistocene hominins

Previous attempts to estimate body mass in pre-Holocene hominins have relied on prediction equations derived from relatively limited extant samples. Here we derive new equations to predict body mass from femoral head breadth and proximal tibial plateau breadth based on a large and diverse sample of modern humans (avoiding the problems associated with using diaphyseal dimensions and/or cadaveric reference samples). In addition, an adjustment for the relatively small femoral heads of non-Homo taxa is developed based on observed differences in hip to knee joint scaling. Body mass is then estimated for 214 terminal Miocene through Pleistocene hominin specimens. Mean body masses for non-Homo taxa range between 39 and 49 kg (39–45 kg if sex-specific means are averaged), with no consistent temporal trend (6–1.85 Ma). Mean body mass increases in early Homo (2.04–1.77 Ma) to 55–59 kg, and then again dramatically in Homo erectus and later archaic middle Pleistocene Homo, to about 70 kg. The same average body mass is maintained in late Pleistocene archaic Homo and early anatomically modern humans through the early/middle Upper Paleolithic (0.024 Ma), only declining in the late Upper Paleolithic, with regional variation. Sexual dimorphism in body mass is greatest in Australopithecus afarensis (log[male/female] = 1.54), declines in Australopithecus africanus and Paranthropus robustus (log ratio 1.36), and then again in early Homo and middle and late Pleistocene archaic Homo (log ratio 1.20–1.27), although it remains somewhat elevated above that of living and middle/late Pleistocene anatomically modern humans (log ratio about 1.15).

Correction: The lithic assemblages of Donggutuo, Nihewan basin: Knapping skills of Early Pleistocene hominins in North China.

[This corrects the article DOI: 10.1371/journal.pone.0185101.].

The duration of a Yellowstone super-eruption cycle and implications for the age of the Olduvai subchron

Small-volume rhyolitic eruptions preceding and following a caldera-forming eruption can provide insights into the tempo of eruption cycles and timing of magmatic recharge. In this contribution, high-precision 40Ar/39Ar eruption ages were obtained on the three effusive eruptions bracketing the Huckleberry Ridge Tuff, which comprise Yellowstone's first volcanic cycle. These dates are supplemented with detailed paleomagnetic and rock magnetic analyses to resolve discrepancies with previous reported stratigraphy. The Huckleberry Ridge Tuff (2.08 Ma) was preceded by an eruption at 2.14 Ma, and followed by eruptions at 1.98 and 1.95 Ma, all of which occurred during four distinct periods of geomagnetic instability within the Matuyama chron. The first volcanic cycle of Yellowstone has now been constrained to within a 200 kyr timespan, or half of the previously proposed duration, and similar to the duration of volcanic activity for caldera-forming systems in the Jemez Volcanic Field. The maximum duration for magmatic recharge for the first Yellowstone volcanic cycle is no greater than 100 kyr, and likely closer to 40 kyr. Furthermore, the combined 40Ar/39Ar eruption ages and paleomagnetic results provide polarity anchors for the Pre-Olduvai excursion and Olduvai subchron, which are often used as tie-points in studies of early Pleistocene hominin evolution.