Narrative cave art in Indonesia by 51,200 years ago.

Previous dating research indicated that the Indonesian island of Sulawesi is host to some of the oldest known rock art1-3. That work was based on solution uranium-series (U-series) analysis of calcite deposits overlying rock art in the limestone caves of Maros-Pangkep, South Sulawesi1-3. Here we use a novel application of this approach-laser-ablation U-series imaging-to re-date some of the earliest cave art in this karst area and to determine the age of stylistically similar motifs at other Maros-Pangkep sites. This method provides enhanced spatial accuracy, resulting in older minimum ages for previously dated art. We show that a hunting scene from Leang Bulu' Sipong 4, which was originally dated using the previous approach to a minimum of 43,900 thousand years ago (ka)3, has a minimum age of 50.2 ± 2.2 ka, and so is at least 4,040 years older than thought. Using the imaging approach, we also assign a minimum age of 53.5 ± 2.3 ka to a newly described cave art scene at Leang Karampuang. Painted at least 51,200 years ago, this narrative composition, which depicts human-like figures interacting with a pig, is now the earliest known surviving example of representational art, and visual storytelling, in the world3. Our findings show that figurative portrayals of anthropomorphic figures and animals have a deeper origin in the history of modern human (Homo sapiens) image-making than recognized to date, as does their representation in composed scenes.

Surgical amputation of a limb 31,000 years ago in Borneo.

The prevailing view regarding the evolution of medicine is that the emergence of settled agricultural societies around 10,000 years ago (the Neolithic Revolution) gave rise to a host of health problems that had previously been unknown among non-sedentary foraging populations, stimulating the first major innovations in prehistoric medical practices1,2. Such changes included the development of more advanced surgical procedures, with the oldest known indication of an 'operation' formerly thought to have consisted of the skeletal remains of a European Neolithic farmer (found in Buthiers-Boulancourt, France) whose left forearm had been surgically removed and then partially healed3. Dating to around 7,000 years ago, this accepted case of amputation would have required comprehensive knowledge of human anatomy and considerable technical skill, and has thus been viewed as the earliest evidence of a complex medical act3. Here, however, we report the discovery of skeletal remains of a young individual from Borneo who had the distal third of their left lower leg surgically amputated, probably as a child, at least 31,000 years ago. The individual survived the procedure and lived for another 6-9 years, before their remains were intentionally buried in Liang Tebo cave, which is located in East Kalimantan, Indonesian Borneo, in a limestone karst area that contains some of the world's earliest dated rock art4. This unexpectedly early evidence of a successful limb amputation suggests that at least some modern human foraging groups in tropical Asia had developed sophisticated medical knowledge and skills long before the Neolithic farming transition.

Two Late Pleistocene human femora from Trinil, Indonesia: Implications for body size and behavior in Southeast Asia.

Late Pleistocene hominin postcranial specimens from Southeast Asia are relatively rare. Here we describe and place into temporal and geographic context two partial femora from the site of Trinil, Indonesia, which are dated stratigraphically and via Uranium-series direct dating to ca. 37-32 ka. The specimens, designated Trinil 9 and 10, include most of the diaphysis, with Trinil 9 being much better preserved. Microcomputed tomography is used to determine cross-sectional diaphyseal properties, with an emphasis on midshaft anteroposterior to mediolateral bending rigidity (Ix/Iy), which has been shown to relate to both body shape and activity level in modern humans. The body mass of Trinil 9 is estimated from cortical area and reconstructed length using new equations based on a Pleistocene reference sample. Comparisons are carried out with a large sample of Pleistocene and Holocene East Asian, African, and European/West Asian femora. Our results show that Trinil 9 has a high Ix/Iy ratio, most consistent with a relatively narrow-bodied male from a mobile hunting-gathering population. It has an estimated body mass of 55.4 kg and a stature of 156 cm, which are small relative to Late Pleistocene males worldwide, but larger than the penecontemporaneous Deep Skull femur from Niah Cave, Malaysia, which is very likely female. This suggests the presence of small-bodied active hunter-gatherers in Southeast Asia during the later Late Pleistocene. Trinil 9 also contrasts strongly in morphology with earlier partial femora from Trinil dating to the late Early-early Middle Pleistocene (Femora II-V), and to a lesser extent with the well-known complete Femur I, most likely dating to the terminal Middle-early Late Pleistocene. Temporal changes in morphology among femoral specimens from Trinil parallel those observed in Homo throughout the Old World during the Pleistocene and document these differences within a single site.

Skeletal remains of a Pleistocene modern human (Homo sapiens) from Sulawesi.

Major gaps remain in our knowledge of the early history of Homo sapiens in Wallacea. By 70-60 thousand years ago (ka), modern humans appear to have entered this distinct biogeographical zone between continental Asia and Australia. Despite this, there are relatively few Late Pleistocene sites attributed to our species in Wallacea. H. sapiens fossil remains are also rare. Previously, only one island in Wallacea (Alor in the southeastern part of the archipelago) had yielded skeletal evidence for pre-Holocene modern humans. Here we report on the first Pleistocene human skeletal remains from the largest Wallacean island, Sulawesi. The recovered elements consist of a nearly complete palate and frontal process of a modern human right maxilla excavated from Leang Bulu Bettue in the southwestern peninsula of the island. Dated by several different methods to between 25 and 16 ka, the maxilla belongs to an elderly individual of unknown age and sex, with small teeth (only M1 to M3 are extant) that exhibit severe occlusal wear and related dental pathologies. The dental wear pattern is unusual. This fragmentary specimen, though largely undiagnostic with regards to morphological affinity, provides the only direct insight we currently have from the fossil record into the identity of the Late Pleistocene people of Sulawesi.

Oldest cave art found in Sulawesi.

Indonesia harbors some of the oldest known surviving cave art. Previously, the earliest dated rock art from this region was a figurative painting of a Sulawesi warty pig (Sus celebensis). This image from Leang Bulu' Sipong 4 in the limestone karsts of Maros-Pangkep, South Sulawesi, was created at least 43,900 years ago (43.9 ka) based on Uranium-series dating. Here, we report the Uranium-series dating of two figurative cave paintings of Sulawesi warty pigs recently discovered in the same karst area. The oldest, with a minimum age of 45.5 ka, is from Leang Tedongnge. The second image, from Leang Balangajia 1, dates to at least 32 ka. To our knowledge, the animal painting from Leang Tedongnge is the earliest known representational work of art in the world. There is no reason to suppose, however, that this early rock art is a unique example in Island Southeast Asia or the wider region.

Dating the skull from Broken Hill, Zambia, and its position in human evolution.

The cranium from Broken Hill (Kabwe) was recovered from cave deposits in 1921, during metal ore mining in what is now Zambia1. It is one of the best-preserved skulls of a fossil hominin, and was initially designated as the type specimen of Homo rhodesiensis, but recently it has often been included in the taxon Homo heidelbergensis2-4. However, the original site has since been completely quarried away, and-although the cranium is often estimated to be around 500 thousand years old5-7-its unsystematic recovery impedes its accurate dating and placement in human evolution. Here we carried out analyses directly on the skull and found a best age estimate of 299 ± 25 thousand years (mean ± 2σ). The result suggests that later Middle Pleistocene Africa contained multiple contemporaneous hominin lineages (that is, Homo sapiens8,9, H. heidelbergensis/H. rhodesiensis and Homo naledi10,11), similar to Eurasia, where Homo neanderthalensis, the Denisovans, Homo floresiensis, Homo luzonensis and perhaps also Homo heidelbergensis and Homo erectus12 were found contemporaneously. The age estimate also raises further questions about the mode of evolution of H. sapiens in Africa and whether H. heidelbergensis/H. rhodesiensis was a direct ancestor of our species13,14.

Earliest hunting scene in prehistoric art.

Humans seem to have an adaptive predisposition for inventing, telling and consuming stories1. Prehistoric cave art provides the most direct insight that we have into the earliest storytelling2-5, in the form of narrative compositions or 'scenes'2,5 that feature clear figurative depictions of sets of figures in spatial proximity to each other, and from which one can infer actions taking place among the figures5. The Upper Palaeolithic cave art of Europe hosts the oldest previously known images of humans and animals interacting in recognizable scenes2,5, and of therianthropes6,7-abstract beings that combine qualities of both people and animals, and which arguably communicated narrative fiction of some kind (folklore, religious myths, spiritual beliefs and so on). In this record of creative expression (spanning from about 40 thousand years ago (ka) until the beginning of the Holocene epoch at around 10 ka), scenes in cave art are generally rare and chronologically late (dating to about 21-14 ka)7, and clear representations of therianthropes are uncommon6-the oldest such image is a carved figurine from Germany of a human with a feline head (dated to about 40-39 ka)8. Here we describe an elaborate rock art panel from the limestone cave of Leang Bulu' Sipong 4 (Sulawesi, Indonesia) that portrays several figures that appear to represent therianthropes hunting wild pigs and dwarf bovids; this painting has been dated to at least 43.9 ka on the basis of uranium-series analysis of overlying speleothems. This hunting scene is-to our knowledge-currently the oldest pictorial record of storytelling and the earliest figurative artwork in the world.

Pre-Ribosomal RNA Processing in Human Cells: From Mechanisms to Congenital Diseases.

Ribosomal RNAs, the most abundant cellular RNA species, have evolved as the structural scaffold and the catalytic center of protein synthesis in every living organism. In eukaryotes, they are produced from a long primary transcript through an intricate sequence of processing steps that include RNA cleavage and folding and nucleotide modification. The mechanisms underlying this process in human cells have long been investigated, but technological advances have accelerated their study in the past decade. In addition, the association of congenital diseases to defects in ribosome synthesis has highlighted the central place of ribosomal RNA maturation in cell physiology regulation and broadened the interest in these mechanisms. Here, we give an overview of the current knowledge of pre-ribosomal RNA processing in human cells in light of recent progress and discuss how dysfunction of this pathway may contribute to the physiopathology of congenital diseases.

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

[This corrects the article DOI: 10.1371/journal.pone.0193025.].

Rare Late Pleistocene-early Holocene human mandibles from the Niah Caves (Sarawak, Borneo).

The skeletal remains of Late Pleistocene-early Holocene humans are exceptionally rare in island Southeast Asia. As a result, the identity and physical adaptations of the early inhabitants of the region are poorly known. One archaeological locality that has historically been important for understanding the peopling of island Southeast Asia is the Niah Caves in the northeast of Borneo. Here we present the results of direct Uranium-series dating and the first published descriptions of three partial human mandibles from the West Mouth of the Niah Caves recovered during excavations by the Harrissons in 1957. One of them (mandible E/B1 100") is somewhat younger than the 'Deep Skull' with a best dating estimate of c30-28 ka (at 2σ), while the other two mandibles (D/N5 42-48" and E/W 33 24-36") are dated to a minimum of c11.0-10.5 ka (at 2σ) and c10.0-9.0 ka (at 2σ). Jaw E/B1 100" is unusually small and robust compared with other Late Pleistocene mandibles suggesting that it may have been ontogenetically altered through masticatory strain under a model of phenotypic plasticity. Possible dietary causes could include the consumption of tough or dried meats or palm plants, behaviours which have been documented previously in the archaeological record of the Niah Caves. Our work suggests a long history back to before the LGM of economic strategies involving the exploitation of raw plant foods or perhaps dried and stored meat resources. This offers new insights into the economic strategies of Late Pleistocene-early Holocene hunter-gatherers living in, or adjacent to, tropical rainforests.

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.

U-series dating and classification of the Apidima 2 hominin from Mani Peninsula, Southern Greece.

Laser ablation U-series dating results on a human cranial bone fragment from Apidima, on the western cost of the Mani Peninsula, Southern Greece, indicate a minimum age of 160,000 years. The dated cranial fragment belongs to Apidima 2, which preserves the facial skeleton and a large part of the braincase, lacking the occipital bone. The morphology of the preserved regions of the cranium, and especially that of the facial skeleton, indicates that the fossil belongs to the Neanderthal clade. The dating of the fossil at a minimum age of 160,000 years shows that most of the Neanderthal traits were already present in the MIS 6 and perhaps earlier. This makes Apidima 2 the earliest known fossil with a clear Neanderthal facial morphology. Together with the nearby younger Neanderthal specimens from Lakonis and Kalamakia, the Apidima crania are of crucial importance for the evolution of Neanderthals in the area during the Middle to Late Pleistocene. It can be expected that systematic direct dating of the other human fossils from this area will elucidate our understanding of Neanderthal evolution and demise.

Early human symbolic behavior in the Late Pleistocene of Wallacea.

Wallacea, the zone of oceanic islands separating the continental regions of Southeast Asia and Australia, has yielded sparse evidence for the symbolic culture of early modern humans. Here we report evidence for symbolic activity 30,000-22,000 y ago at Leang Bulu Bettue, a cave and rock-shelter site on the Wallacean island of Sulawesi. We describe hitherto undocumented practices of personal ornamentation and portable art, alongside evidence for pigment processing and use in deposits that are the same age as dated rock art in the surrounding karst region. Previously, assemblages of multiple and diverse types of Pleistocene "symbolic" artifacts were entirely unknown from this region. The Leang Bulu Bettue assemblage provides insight into the complexity and diversification of modern human culture during a key period in the global dispersal of our species. It also shows that early inhabitants of Sulawesi fashioned ornaments from body parts of endemic animals, suggesting modern humans integrated exotic faunas and other novel resources into their symbolic world as they colonized the biogeographically unique regions southeast of continental Eurasia.

Revised stratigraphy and chronology for Homo floresiensis at Liang Bua in Indonesia.

Homo floresiensis, a primitive hominin species discovered in Late Pleistocene sediments at Liang Bua (Flores, Indonesia), has generated wide interest and scientific debate. A major reason this taxon is controversial is because the H. floresiensis-bearing deposits, which include associated stone artefacts and remains of other extinct endemic fauna, were dated to between about 95 and 12 thousand calendar years (kyr) ago. These ages suggested that H. floresiensis survived until long after modern humans reached Australia by ~50 kyr ago. Here we report new stratigraphic and chronological evidence from Liang Bua that does not support the ages inferred previously for the H. floresiensis holotype (LB1), ~18 thousand calibrated radiocarbon years before present (kyr cal. BP), or the time of last appearance of this species (about 17 or 13-11 kyr cal. BP). Instead, the skeletal remains of H. floresiensis and the deposits containing them are dated to between about 100 and 60 kyr ago, whereas stone artefacts attributable to this species range from about 190 to 50 kyr in age. Whether H. floresiensis survived after 50 kyr ago--potentially encountering modern humans on Flores or other hominins dispersing through southeast Asia, such as Denisovans--is an open question.

Spying on small wildlife sounds using affordable collar-mounted miniature microphones: an innovative method to record individual daylong vocalisations in chipmunks.

Technological advances can greatly benefit the scientific community by making new areas of research accessible. The study of animal vocal communication, in particular, can gain new insights and knowledge from technological improvements in recording equipment. Our comprehension of the acoustic signals emitted by animals would be greatly improved if we could continuously track the daily natural emissions of individuals in the wild, especially in the context of integrating individual variation into evolutionary ecology research questions. We show here how this can be accomplished using an operational tiny audio recorder that can easily be fitted as an on-board acoustic data-logger on small free-ranging animals. The high-quality 24 h acoustic recording logged on the spy microphone device allowed us to very efficiently collect daylong chipmunk vocalisations, giving us much more detailed data than the classical use of a directional microphone over an entire field season. The recordings also allowed us to monitor individual activity patterns and record incredibly long resting heart rates, and to identify self-scratching events and even whining from pre-emerging pups in their maternal burrow.

The chronostratigraphy of the Haua Fteah cave (Cyrenaica, northeast Libya).

The 1950s excavations by Charles McBurney in the Haua Fteah, a large karstic cave on the coast of northeast Libya, revealed a deep sequence of human occupation. Most subsequent research on North African prehistory refers to his discoveries and interpretations, but the chronology of its archaeological and geological sequences has been based on very early age determinations. This paper reports on the initial results of a comprehensive multi-method dating program undertaken as part of new work at the site, involving radiocarbon dating of charcoal, land snails and marine shell, cryptotephra investigations, optically stimulated luminescence (OSL) dating of sediments, and electron spin resonance (ESR) dating of tooth enamel. The dating samples were collected from the newly exposed and cleaned faces of the upper 7.5 m of the ∼14.0 m-deep McBurney trench, which contain six of the seven major cultural phases that he identified. Despite problems of sediment transport and reworking, using a Bayesian statistical model the new dating program establishes a robust framework for the five major lithostratigraphic units identified in the stratigraphic succession, and for the major cultural units. The age of two anatomically modern human mandibles found by McBurney in Layer XXXIII near the base of his Levalloiso-Mousterian phase can now be estimated to between 73 and 65 ka (thousands of years ago) at the 95.4% confidence level, within Marine Isotope Stage (MIS) 4. McBurney's Layer XXV, associated with Upper Palaeolithic Dabban blade industries, has a clear stratigraphic relationship with Campanian Ignimbrite tephra. Microlithic Oranian technologies developed following the climax of the Last Glacial Maximum and the more microlithic Capsian in the Younger Dryas. Neolithic pottery and perhaps domestic livestock were used in the cave from the mid Holocene but there is no certain evidence for plant cultivation until the Graeco-Roman period.

U-series and radiocarbon analyses of human and faunal remains from Wajak, Indonesia.

Laser ablation U-series dating results on human and faunal bone fragments from Wajak, Indonesia, indicate a minimum age of between 37.4 and 28.5 ka (thousands of years ago) for the whole assemblage. These are significantly older than previously published radiocarbon estimates on bone carbonate, which suggested a Holocene age for a human bone fragment and a late Pleistocene age for a faunal bone. The analysis of the organic components in the faunal material show severe degradation and a positive δ(13)C ratio indicate a high degree of secondary carbonatisation. This may explain why the thermal release method used for the original age assessments yielded such young ages. While the older U-series ages are not in contradiction with the morphology of the Wajak human fossils or Javanese biostratigraphy, they will require a reassessment of the evolutionary relationships of modern human remains in Southeast Asia and Oceania. It can be expected that systematic direct dating of human fossils from this area will lead to further revisions of our understanding of modern human evolution.

Confirmation of a late middle Pleistocene age for the Omo Kibish 1 cranium by direct uranium-series dating.

While it is generally accepted that modern humans evolved in Africa, the specific physical evidence for that origin remains disputed. The modern-looking Omo 1 skeleton, discovered in the Kibish region of Ethiopia in 1967, was controversially dated at ~130 ka (thousands of years ago) by U-series dating on associated Mollusca, and it was not until 2005 that Ar-Ar dating on associated feldspar crystals in pumice clasts provided evidence for an even older age of ~195 ka. However, questions continue to be raised about the age and stratigraphic position of this crucial fossil specimen. Here we present direct U-series determinations on the Omo 1 cranium. In spite of significant methodological complications, which are discussed in detail, the results indicate that the human remains do not belong to a later intrusive burial and are the earliest representative of anatomically modern humans. Given the more archaic morphology shown by the apparently contemporaneous Omo 2 calvaria, we suggest that direct U-series dating is applied to this fossil as well, to confirm its age in relation to Omo 1.