The demise of the giant ape Gigantopithecus blacki.

The largest ever primate and one of the largest of the southeast Asian megafauna, Gigantopithecus blacki1, persisted in China from about 2.0 million years until the late middle Pleistocene when it became extinct2-4. Its demise is enigmatic considering that it was one of the few Asian great apes to go extinct in the last 2.6 million years, whereas others, including orangutan, survived until the present5. The cause of the disappearance of G. blacki remains unresolved but could shed light on primate resilience and the fate of megafauna in this region6. Here we applied three multidisciplinary analyses-timing, past environments and behaviour-to 22 caves in southern China. We used 157 radiometric ages from six dating techniques to establish a timeline for the demise of G. blacki. We show that from 2.3 million years ago the environment was a mosaic of forests and grasses, providing ideal conditions for thriving G. blacki populations. However, just before and during the extinction window between 295,000 and 215,000 years ago there was enhanced environmental variability from increased seasonality, which caused changes in plant communities and an increase in open forest environments. Although its close relative Pongo weidenreichi managed to adapt its dietary preferences and behaviour to this variability, G. blacki showed signs of chronic stress and dwindling populations. Ultimately its struggle to adapt led to the extinction of the greatest primate to ever inhabit the Earth.

Why the geosciences are becoming increasingly vital to the interpretation of the human evolutionary record.

Advanced geoscience techniques are essential to contextualize fossils, artefacts and other archaeologically important material accurately and effectively. Their appropriate use will increase confidence in new interpretations of the fossil and archaeological record, providing important information about the life and depositional history of these materials and so should form an integral component of all human evolutionary studies. Many of the most remarkable recent finds that have transformed the field of human evolution are small and scarce, ranging in size from teeth to strands of DNA, recovered from complex sedimentary environments. Nevertheless, if properly analysed, they hold immense potential to rewrite what we know about the evolution of our species and our closest hominin ancestors.

Palaeoenvironments and hominin evolutionary dynamics in southeast Asia.

Secure environmental contexts are crucial for hominin interpretation and comparison. The discovery of a Denisovan individual and associated fauna at Tam Ngu Hao 2 (Cobra) Cave, Laos, dating back to 164-131 ka, allows for environmental comparisons between this (sub)tropical site and the Palearctic Denisovan sites of Denisova Cave (Russia) and Baishiya Karst Cave (China). Denisovans from northern latitudes foraged in a mix of forested and open landscapes, including tundra and steppe. Using stable isotope values from the Cobra Cave assemblage, we demonstrate that, despite the presence of nearby canopy forests, the Denisovan individual from Cobra Cave primarily consumed plants and/or animals from open forests and savannah. Using faunal evidence and proxy indicators of climates, results herein highlight a local expansion of rainforest at ~ 130 ka, raising questions about how Denisovans responded to this local climate change. Comparing the diet and habitat of the archaic hominin from Cobra Cave with those of early Homo sapiens from Tam Pà Ling Cave (46-43 ka), Laos, it appears that only our species was able to exploit rainforest resources.

Preliminary description of a late Middle Pleistocene mammalian fauna prior to the extinction of Gigantopithecus blacki from the Yixiantian Cave, Guangxi ZAR, South China.

In recent years, nearly 20 cave sites with rich assemblages of mammalian fossils have been found and excavated in the Chongzuo area, Guangxi Zhuang Autonomous Region, China. Their ages are distributed throughout the entire Pleistocene Epoch. These discoveries have greatly facilitated our understanding of the evolution of the Stegodon-Ailuropoda fauna and the environmental context of human evolution in southern China. Here, we present a preliminary report on a diverse late Middle Pleistocene mammalian fauna from the Yixiantian Cave in southern China, which is a typical representative of the Stegodon-Ailuropoda fauna (sensu lato). The fossil mammals are represented by isolated dental remains only. In 2010 and 2011, two seasons of systematic excavations at the Yixiantian Cave yielded a total of 4,958 identifiable mammalian teeth. They were identified as belonging to 37 species and 6 orders of mammals. In addition, the tooth type of all the teeth representing each species was also determined where possible. A single fragmentary molar was identified as belonging to Gigantopithecus blacki, indicating that its population had declined sharply at this time and was on the brink of extinction. Description of the Yixiantian fauna will not only help better characterize the composition of the Stegodon-Ailuropoda fauna during the late Middle Pleistocene, but also clarify our understanding of the paleoenvironmental context at a time just prior to the extinction of G. blacki.

Early presence of Homo sapiens in Southeast Asia by 86-68 kyr at Tam Pà Ling, Northern Laos.

The timing of the first arrival of Homo sapiens in East Asia from Africa and the degree to which they interbred with or replaced local archaic populations is controversial. Previous discoveries from Tam Pà Ling cave (Laos) identified H. sapiens in Southeast Asia by at least 46 kyr. We report on a recently discovered frontal bone (TPL 6) and tibial fragment (TPL 7) found in the deepest layers of TPL. Bayesian modeling of luminescence dating of sediments and U-series and combined U-series-ESR dating of mammalian teeth reveals a depositional sequence spanning ~86 kyr. TPL 6 confirms the presence of H. sapiens by 70 ± 3 kyr, and TPL 7 extends this range to 77 ± 9 kyr, supporting an early dispersal of H. sapiens into Southeast Asia. Geometric morphometric analyses of TPL 6 suggest descent from a gracile immigrant population rather than evolution from or admixture with local archaic populations.

A Middle Pleistocene Denisovan molar from the Annamite Chain of northern Laos.

The Pleistocene presence of the genus Homo in continental Southeast Asia is primarily evidenced by a sparse stone tool record and rare human remains. Here we report a Middle Pleistocene hominin specimen from Laos, with the discovery of a molar from the Tam Ngu Hao 2 (Cobra Cave) limestone cave in the Annamite Mountains. The age of the fossil-bearing breccia ranges between 164-131 kyr, based on the Bayesian modelling of luminescence dating of the sedimentary matrix from which it was recovered, U-series dating of an overlying flowstone, and U-series-ESR dating of associated faunal teeth. Analyses of the internal structure of the molar in tandem with palaeoproteomic analyses of the enamel indicate that the tooth derives from a young, likely female, Homo individual. The close morphological affinities with the Xiahe specimen from China indicate that they belong to the same taxon and that Tam Ngu Hao 2 most likely represents a Denisovan.

Speleological and environmental history of Lida Ajer cave, western Sumatra.

Some of the earliest evidence for the presence of modern humans in rainforests has come from the fossil deposits of Lida Ajer in Sumatra. Two human teeth from this cave were estimated to be 73-63 thousand years old, which is significantly older than some estimates of modern human migration out of Africa based on genetic data. The deposits were interpreted as being associated with a rainforest environment based largely on the presence of abundant orangutan fossils. As well as the main fossil-bearing chamber, fossil-bearing passages are present below a sinkhole, although the relationship between the different fossil deposits has only been tenuously established. Here, we provide significant new sedimentological, geochronological and palaeoecological data aimed at reconstructing the speleological and environmental history of the cave and the clastic and fossil deposits therein. Our data suggest that the Lida Ajer fossils were deposited during Marine Isotope Stage 4, with fossils from the lower passages older than the main fossil chamber. Our use of stable carbon and oxygen isotope analyses of mammalian tooth enamel demonstrates that early humans probably occupied a closed-canopy forest very similar to those present in the region today, although the fossil orangutans may have occupied a slightly different niche. This article is part of the theme issue 'Tropical forests in the deep human past'.

Trophic ecology of a Late Pleistocene early modern human from tropical Southeast Asia inferred from zinc isotopes.

Tam Pà Ling, a cave site in northeastern Laos, has yielded the earliest skeletal evidence of Homo sapiens in mainland Southeast Asia. The reliance of Pleistocene humans in rainforest settings on plant or animal resources is still largely unstudied, mainly due to poor collagen preservation in fossils from tropical environments precluding stable nitrogen isotope analysis, the classical trophic level proxy. However, isotopic ratios of zinc (Zn) in bioapatite constitute a promising proxy to infer trophic and dietary information from fossil vertebrates, even under adverse tropical taphonomic conditions. Here, we analyzed the zinc isotope composition (66Zn/64Zn expressed as δ66Zn value) in the enamel of two teeth of the Late Pleistocene (63-46 ka) H. sapiens individual (TPL1) from Tam Pà Ling, as well as 76 mammal teeth from the same site and the nearby Nam Lot cave. The human individual exhibits relatively low enamel δ66Zn values (+0.24‰) consistent with an omnivorous diet, suggesting a dietary reliance on both plant and animal matter. These findings offer direct evidence of the broad utilization of resources from tropical rainforests by one of the earliest known anatomically modern humans in Southeast Asia.

A multi-proxy approach to exploring Homo sapiens' arrival, environments and adaptations in Southeast Asia.

The capability of Pleistocene hominins to successfully adapt to different types of tropical forested environments has long been debated. In order to investigate environmental changes in Southeast Asia during a critical period for the turnover of hominin species, we analysed palaeoenvironmental proxies from five late Middle to Late Pleistocene faunas. Human teeth discoveries have been reported at Duoi U'Oi, Vietnam (70-60 ka) and Nam Lot, Laos (86-72 ka). However, the use of palaeoproteomics allowed us to discard the latter, and, to date, no human remains older than ~ 70 ka are documented in the area. Our findings indicate that tropical rainforests were highly sensitive to climatic changes over that period, with significant fluctuations of the canopy forests. Locally, large-bodied faunas were resilient to these fluctuations until the cooling period of the Marine Isotope Stage 4 (MIS 4; 74-59 ka) that transformed the overall biotope. Then, under strong selective pressures, populations with new phenotypic characteristics emerged while some other species disappeared. We argue that this climate-driven shift offered new foraging opportunities for hominins in a novel rainforest environment and was most likely a key factor in the settlement and dispersal of our species during MIS 4 in SE Asia.

Archaeology and art in context: Excavations at the Gunu Site Complex, Northwest Kimberley, Western Australia.

The Kimberley region of Western Australia is one of the largest and most diverse rock art provenances in the world, with a complex stylistic sequence spanning at least 16 ka, culminating in the modern art-making of the Wunumbal people. The Gunu Site Complex, in the remote Mitchell River region of the northwest Kimberley, is one of many local expressions of the Kimberley rock art sequence. Here we report excavations at two sites in this complex: Gunu Rock, a sand sheet adjacent to rock art panels; and Gunu Cave, a floor deposit within an extensive rockshelter. Excavations at Gunu Rock provide evidence for two phases of occupation, the first from 7-8 to 2.7 ka, and the second from 1064 cal BP. Excavations at Gunu Rock provide evidence for occupation from the end of the second phase to the recent past. Stone for tools in the early phase were procured from a variety of sources, but quartz crystal reduction dominated the second occupation phase. Small quartz crystals were reduced by freehand percussion to provide small flake tools and blanks for manufacturing small points called nguni by the Wunambal people today. Quartz crystals were prominent in historic ritual practices associated with the Wanjina belief system. Complex methods of making bifacially-thinned and pressure flaked quartzite projectile points emerged after 2.7 ka. Ochre pigments were common in both occupation phases, but evidence for occupation contemporaneous with the putative age of the oldest rock art styles was not discovered in the excavations. Our results show that developing a complete understanding of rock art production and local occupation patterns requires paired excavations inside and outside of the rockshelters that dominate the Kimberley.

Last appearance of Homo erectus at Ngandong, Java, 117,000-108,000 years ago.

Homo erectus is the founding early hominin species of Island Southeast Asia, and reached Java (Indonesia) more than 1.5 million years ago1,2. Twelve H. erectus calvaria (skull caps) and two tibiae (lower leg bones) were discovered from a bone bed located about 20 m above the Solo River at Ngandong (Central Java) between 1931 and 19333,4, and are of the youngest, most-advanced form of H. erectus5-8. Despite the importance of the Ngandong fossils, the relationship between the fossils, terrace fill and ages have been heavily debated9-14. Here, to resolve the age of the Ngandong evidence, we use Bayesian modelling of 52 radiometric age estimates to establish-to our knowledge-the first robust chronology at regional, valley and local scales. We used uranium-series dating of speleothems to constrain regional landscape evolution; luminescence, 40argon/39argon (40Ar/39Ar) and uranium-series dating to constrain the sequence of terrace evolution; and applied uranium-series and uranium series-electron-spin resonance (US-ESR) dating to non-human fossils to directly date our re-excavation of Ngandong5,15. We show that at least by 500 thousand years ago (ka) the Solo River was diverted into the Kendeng Hills, and that it formed the Solo terrace sequence between 316 and 31 ka and the Ngandong terrace between about 140 and 92 ka. Non-human fossils recovered during the re-excavation of Ngandong date to between 109 and 106 ka (uranium-series minimum)16 and 134 and 118 ka (US-ESR), with modelled ages of 117 to 108 thousand years (kyr) for the H. erectus bone bed, which accumulated during flood conditions3,17. These results negate the extreme ages that have been proposed for the site and solidify Ngandong as the last known occurrence of this long-lived species.

Macro-charcoal accumulation in floodplain wetlands: Problems and prospects for reconstruction of fire regimes and environmental conditions.

Floodplain wetland ecosystems respond dynamically to flooding, fire and geomorphological processes. We employed a combined geomorphological and environmental proxy approach to assess allochthonous and autochthonous macro-charcoal accumulation in the Macquarie Marshes, Australia, with implications for the reconstruction of fire regimes and environmental conditions in large, open-system wetlands. After accounting for fluvial macro-charcoal flux (1.05 ± 0.32 no. cm-2 a-1), autochthonous macro-charcoal in ~1 m deep sediment profiles spanning ~1.7 ka were highly variable and inconsistent between cores and wetlands (concentrations from 0 to 438 no. cm-3, mean accumulation rates from 0 to 3.86 no. cm-2 a-1). A positive correlation existed between the number of recent fires, satellite-observed ignition points, and macro-charcoal concentrations at the surface of the wetlands. Sedimentology, geochemistry, and carbon stable isotopes (δ13C range -15 to -25 ‰) were similar in all cores from both wetlands and varied little with depth. Application of macro-charcoal and other environmental proxy techniques is inherently difficult in large, dynamic wetland systems due to variations in charcoal sources, sediment and charcoal deposition rates, and taphonomic processes. Major problems facing fire history reconstruction using macro-charcoal records in these wetlands include: (1) spatial and temporal variations in fire activity and ash and charcoal products within the wetlands, (2) variations in allochthonous inputs of charcoal from upstream sources, (3) tendency for geomorphic dynamism to affect flow dispersal and sediment and charcoal accumulation, and (4) propensity for post-depositional modification and/or destruction of macro-charcoal by flooding and taphonomic processes. Recognition of complex fire-climate-hydrology-vegetation interactions is essential. High-resolution, multifaceted approaches with reliable geochronologies are required to assess spatial and temporal patterns of fire and to reconstruct in order to interpret wetland fire regimes.

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.].

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.

Into the Past: A Step Towards a Robust Kimberley Rock Art Chronology.

The recent establishment of a minimum age estimate of 39.9 ka for the origin of rock art in Sulawesi has challenged claims that Western Europe was the locus for the production of the world's earliest art assemblages. Tantalising excavated evidence found across northern Australian suggests that Australia too contains a wealth of ancient art. However, the dating of rock art itself remains the greatest obstacle to be addressed if the significance of Australian assemblages are to be recognised on the world stage. A recent archaeological project in the northwest Kimberley trialled three dating techniques in order to establish chronological markers for the proposed, regional, relative stylistic sequence. Applications using optically-stimulated luminescence (OSL) provided nine minimum age estimates for fossilised mudwasp nests overlying a range of rock art styles, while Accelerator Mass Spectrometry radiocarbon (AMS 14C) results provided an additional four. Results confirm that at least one phase of the northwest Kimberley rock art assemblage is Pleistocene in origin. A complete motif located on the ceiling of a rockshelter returned a minimum age estimate of 16 ± 1 ka. Further, our results demonstrate the inherent problems in relying solely on stylistic classifications to order rock art assemblages into temporal sequences. An earlier than expected minimum age estimate for one style and a maximum age estimate for another together illustrate that the Holocene Kimberley rock art sequence is likely to be far more complex than generally accepted with different styles produced contemporaneously well into the last few millennia. It is evident that reliance on techniques that produce minimum age estimates means that many more dating programs will need to be undertaken before the stylistic sequence can be securely dated.

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.

Early modern humans and morphological variation in Southeast Asia: fossil evidence from Tam Pa Ling, Laos.

Little is known about the timing of modern human emergence and occupation in Eastern Eurasia. However a rapid migration out of Africa into Southeast Asia by at least 60 ka is supported by archaeological, paleogenetic and paleoanthropological data. Recent discoveries in Laos, a modern human cranium (TPL1) from Tam Pa Ling's cave, provided the first evidence for the presence of early modern humans in mainland Southeast Asia by 63-46 ka. In the current study, a complete human mandible representing a second individual, TPL 2, is described using discrete traits and geometric morphometrics with an emphasis on determining its population affinity. The TPL2 mandible has a chin and other discrete traits consistent with early modern humans, but it retains a robust lateral corpus and internal corporal morphology typical of archaic humans across the Old World. The mosaic morphology of TPL2 and the fully modern human morphology of TPL1 suggest that a large range of morphological variation was present in early modern human populations residing in the eastern Eurasia by MIS 3.

First discovery of Pleistocene orangutan (Pongo sp.) fossils in Peninsular Malaysia: biogeographic and paleoenvironmental implications.

Nine isolated fossil Pongo teeth from two cave sites in Peninsular Malaysia are reported. These are the first fossil Pongo specimens recorded in Peninsular Malaysia and represent significant southward extensions of the ancient Southeast Asian continental range of fossil Pongo during two key periods of the Quaternary. These new records from Peninsular Malaysia show that ancestral Pongo successfully passed the major biogeographical divide between mainland continental Southeast Asia and the Sunda subregion before 500 ka (thousand years ago). If the presence of Pongo remains in fossil assemblages indicates prevailing forest habitat, then the persistence of Pongo at Batu Caves until 60 ka implies that during the Last Glacial Phase sufficient forest cover persisted in the west coast plain of what is now Peninsular Malaysia at least ten millennia after a presumed corridor of desiccation had extended to central and east Java. Ultimately, environmental conditions of the peninsula during the Last Glacial Maximum evidently became inhospitable for Pongo, causing local extinction. Following post-glacial climatic amelioration and reforestation, a renewed sea barrier prevented re-colonization from the rainforest refugium in Sumatra, accounting for the present day absence of Pongo in apparently hospitable lowland evergreen rainforest of Peninsular Malaysia. The new teeth provide further evidence that Pongo did not undergo a consistent trend toward dental size reduction over time.

Anatomically modern human in Southeast Asia (Laos) by 46 ka.

Uncertainties surround the timing of modern human emergence and occupation in East and Southeast Asia. Although genetic and archeological data indicate a rapid migration out of Africa and into Southeast Asia by at least 60 ka, mainland Southeast Asia is notable for its absence of fossil evidence for early modern human occupation. Here we report on a modern human cranium from Tam Pa Ling, Laos, which was recovered from a secure stratigraphic context. Radiocarbon and luminescence dating of the surrounding sediments provide a minimum age of 51-46 ka, and direct U-dating of the bone indicates a maximum age of ~63 ka. The cranium has a derived modern human morphology in features of the frontal, occipital, maxillae, and dentition. It is also differentiated from western Eurasian archaic humans in aspects of its temporal, occipital, and dental morphology. In the context of an increasingly documented archaic-modern morphological mosaic among the earliest modern humans in western Eurasia, Tam Pa Ling establishes a definitively modern population in Southeast Asia at ~50 ka cal BP. As such, it provides the earliest skeletal evidence for fully modern humans in mainland Southeast Asia.