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.

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.

Early hominin biogeography in Island Southeast Asia.

Island Southeast Asia covers Eurasia's tropical expanse of continental shelf and active subduction zones. Cutting between island landmasses, Wallace's Line separates Sunda and the Eastern Island Arc (the Arc) into distinct tectonic and faunal provinces. West of the line, on Sunda, Java Island yields many fossils of Homo erectus. East of the line, on the Arc, Flores Island provides one skeleton and isolated remains of Homo floresiensis. Luzon Island in the Philippines has another fossil hominin. Sulawesi preserves early hominin archeology. This insular divergence sets up a unique regional context for early hominin dispersal, isolation, and extinction. The evidence is reviewed across three Pleistocene climate periods. Patterns are discussed in relation to the pulse of global sea-level shifts, as well as regional geo-tectonics, catastrophes, stegodon dispersal, and paleogenomics. Several patterns imply evolutionary processes typical of oceanic islands. Early hominins apparently responded to changing island conditions for a million-and-a-half years, likely becoming extinct during the period in which Homo sapiens colonized the region.

Evolutionary trend in dental size in Gigantopithecus blacki revisited.

Previous analyses of dental size in Gigantopithecus blacki indicated marked sexual dimorphism and a trend towards increasing size through time. These studies were based on a sample of over 700 teeth from five localities excavated prior to 1990. Since then, 12 additional cave sites have been discovered in southern China, yielding hundreds of isolated teeth of G. blacki. Most of these sites are well dated by a combination of biochronology and absolute dating methods, so we now have a much better understanding of the chronology of G. blacki. Here, we reexamine the degree of sexual dimorphism and the question of dental size increase through time in G. blacki based on the expanded collections now available. Our results show that sexual dimorphism is not as marked as indicated in previous studies and confirm earlier analyses suggesting that the postcanine teeth of G. blacki tend to become larger through time from the beginning of the Early Pleistocene to the Middle Pleistocene.

Structure and composition of the Trinil femora: functional and taxonomic implications.

The original hominin femur (Femur I) and calotte discovered at Trinil, Java by Eugene Dubois in 1891/1892 played a key role in the early history of human paleontology by purportedly demonstrating the contemporaneity of archaic cranial form with modern human erect (bipedal) posture. On this basis, both specimens were subsequently assigned to Pithecanthropus erectus, later transferred to Homo erectus. However, chronological and phylogenetic links between the two have been questioned from the beginning. Four additional hominin partial femora (Femora II-V) from Trinil were subsequently described but have played a relatively minor part in evolutionary scenarios. Here we present the results of a new analysis of structural and density characteristics of the Trinil femora obtained using computed tomography. Trinil Femur I shows none of the characteristics typical of early Homo femora from elsewhere in Asia or Africa, including a relatively long neck, increased mediolateral bending rigidity of the mid-proximal shaft, or a low position of minimum mediolateral breath on the shaft. In contrast, Femora II-V all demonstrate features that are more consistent with this pattern. In addition, material density distributions within the specimens imply more recent and less complete fossilization of Femur I than Femora II-V. Thus, it is very likely that Trinil Femur I derives from a much more recent time period than the calotte, while the less famous and less complete Femora II-V may represent H. erectus at Trinil. The morphological variation within the Trinil femora can be attributed to broader changes in pelvic morphology occurring within the Homo lineage between the Early and late Middle Pleistocene.

Continuity of mammalian fauna over the last 200,000 y in the Indian subcontinent.

Mammalian extinction worldwide during the Late Pleistocene has been a major focus for Quaternary biochronology and paleoecology. These extinctions have been variably attributed to the impacts of climate change and human interference. However, until relatively recently, research has been largely restricted to the Americas, Europe, and Australasia. We present the oldest Middle-Late Pleistocene stratified and numerically dated faunal succession for the Indian subcontinent from the Billasurgam cave complex. Our data demonstrate continuity of 20 of 21 identified mammalian taxa from at least 100,000 y ago to the present, and in some cases up to 200,000 y ago. Comparison of this fossil record to contemporary faunal ranges indicates some geographical redistribution of mammalian taxa within India. We suggest that, although local extirpations occurred, the majority of taxa survived or adapted to substantial ecological pressures in fragmented habitats. Comparison of the Indian record with faunal records from Southeast and Southwest Asia demonstrates the importance of interconnected mosaic habitats to long-term faunal persistence across the Asian tropics. The data presented here have implications for mammalian conservation in India today, where increasing ecological circumscription may leave certain taxa increasingly endangered in the most densely populated region of the world.

Giant lizards occupied herbivorous mammalian ecospace during the Paleogene greenhouse in Southeast Asia.

Mammals dominate modern terrestrial herbivore ecosystems, whereas extant herbivorous reptiles are limited in diversity and body size. The evolution of reptile herbivory and its relationship to mammalian diversification is poorly understood with respect to climate and the roles of predation pressure and competition for food resources. Here, we describe a giant fossil acrodontan lizard recovered with a diverse mammal assemblage from the late middle Eocene Pondaung Formation of Myanmar, which provides a historical test of factors controlling body size in herbivorous squamates. We infer a predominately herbivorous feeding ecology for the new acrodontan based on dental anatomy, phylogenetic relationships and body size. Ranking body masses for Pondaung Formation vertebrates indicates that the lizard occupied a size niche among the larger herbivores and was larger than most carnivorous mammals. Paleotemperature estimates of Pondaung Formation environments based on the body size of the new lizard are approximately 2-5°C higher than modern. These results indicate that competitive exclusion and predation by mammals did not restrict body size evolution in these herbivorous squamates, and elevated temperatures relative to modern climates during the Paleogene greenhouse may have resulted in the evolution of gigantism through elevated poikilothermic metabolic rates and in response to increases in floral productivity.

New 1.5 million-year-old Homo erectus maxilla from Sangiran (Central Java, Indonesia).

Sangiran (Solo Basin, Central Java, Indonesia) is the singular Homo erectus fossil locale for Early Pleistocene Southeast Asia. Sangiran is the source for more than 80 specimens in deposits with (40)Ar/(39)Ar ages of 1.51-0.9 Ma. In April 2001, we recovered a H. erectus left maxilla fragment (preserving P(3)- M(2)) from the Sangiran site of Bapang. The find spot lies at the base of the Bapang Formation type section in cemented gravelly sands traditionally called the Grenzbank Zone. Two meters above the find spot, pumice hornblende has produced an (40)Ar/(39)Ar age of 1.51 ± 0.08 Ma. With the addition of Bpg 2001.04, Sangiran now has five H. erectus maxillae. We compare the new maxilla with homologs representing Sangiran H. erectus, Zhoukoudian H. erectus, Western H. erectus (pooled African and Georgian specimens), and Homo habilis. Greatest contrast is with the Zhoukoudian maxillae, which appear to exhibit a derived pattern of premolar-molar relationships compared to Western and Sangiran H. erectus. The dental patterns suggest distinct demic origins for the earlier H. erectus populations represented at Sangiran and the later population represented at Zhoukoudian. These two east Asian populations, separated by 5000 km and nearly 800 k.yr., may have had separate origins from different African/west Eurasian populations.

Way out of Africa: Early Pleistocene paleoenvironments inhabited by Homo erectus in Sangiran, Java.

A sequence of paleosols in the Solo Basin, Central Java, Indonesia, documents the local and regional environments present when Homo erectus spread through Southeast Asia during the early Pleistocene. The earliest human immigrants encountered a low-relief lake-margin landscape dominated by moist grasslands with open woodlands in the driest landscape positions. By 1.5 Ma, large streams filled the lake and the landscape became more riverine in nature, with riparian forests, savanna, and open woodland. Paleosol morphology and carbon isotope values of soil organic matter and pedogenic carbonates indicate a long-term shift toward regional drying or increased duration of the annual dry season through the early Pleistocene. This suggests that an annual dry season associated with monsoon conditions was an important aspect of the paleoclimate in which early humans spread from Africa to Southeast Asia.

Assessing mandibular shape variation within Gigantopithecus using a geometric morphometric approach.

This study provides a survey of mandibular shape in a sample of extant hominoids (Pan, Gorilla, Pongo, and Hylobates), as well as extinct Asian and Eurasian taxa (Ouranopithecus, Sivapithecus, and Gigantopithecus) in order to compare overall shape similarity. Results presented call into question differences in mandible shape recently used to distinguish Gigantopithecus giganteus from Gigantopithecus blacki and to justify resurrecting a different generic designation, "Indopithecus," for the former. It is concluded that while the two large-bodied Asian taxa may have been adapted to slightly different dietary niches with different geographic and temporal ranges, the unique mandibular/dental characters that the two taxa share should not be viewed as independent evolutionary developments.

The first skull of the earliest giant panda.

Fossils of the giant panda Ailuropoda (Order Carnivora, Family Ursidae) are largely isolated teeth, mandibles, and a few rare skulls, known from the late Pliocene to late Pleistocene in China and Southeast Asia. Much of this material represents a Pleistocene chronospecies, Ailuropoda baconi, an animal larger than the living giant panda, Ailuropoda melanoleuca. The earliest certain record of Ailuropoda is the late Pliocene chronospecies, Ailuropoda microta, smaller than either A. baconi or A. melanoleuca, and previously known only from teeth and a few mandibles from karst caves in south China. Here, we report the discovery of the first skull of A. microta, establishing its cranial anatomy and demonstrating that the specialized cranial and dental adaptations of Ailuropoda for durophagous feeding behavior centered on bamboo were already evident in this late Pliocene species. The skull from Jinyin cave (Guangxi) and dental remains from other karst localities in southeastern China show that Ailuropoda microta occupied south China from approximately 2 to 2.4 Myr ago after a marked global climatic deterioration. Dental and basicranial anatomy indicate a less specialized morphology early in the history of the lineage and support derivation of the giant panda from the Miocene Asian ursid Ailurarctos.

Mapping and taphonomic analysis of the Homo erectus loci at Locality 1 Zhoukoudian, China.

From a detailed analysis of published and unpublished sources, we constructed a digitized three-dimensional, stratigraphically-controlled excavation grid of Zhoukoudian Locality 1 in order to assess the spatial relationships of the excavated materials. All 15 fossil Homo erectus loci were mapped on the grid. Meter cubes were used in excavation starting in 1934, and Loci H through O, established between 1934 and 1937, were mapped to within 1 m(3)vertical and horizontal provenience. Loci A through G, established between 1921 and 1933, were excavated in the northernmost part of Locality 1 by unmapped quarrying, but their stratigraphic levels were recorded. We could localize Loci A through G on the grid system by utilizing locations of remaining walls, stratigraphic sections, excavation reports, excavation maps, and photographs. Loci contained skeletal elements of Homo erectus individuals scattered over areas of the cave floor of up to 9 m in diameter. Scoring of taphonomic damage on the Homo erectus sample, as observed on casts and originals, demonstrates that 67% of the hominid sample shows bite marks or other modifications ascribed to large mammalian carnivores, particularly the large Pleistocene cave hyena, Pachycrocuta brevirostris. Virtually all of the remaining Homo erectus skeletal assemblage shows breakage consistent with this taphonomic pattern of fragmentation. Bioturbation by digging carnivores is the most likely explanation for a fragment of Homo erectus Skull XI discovered 1 m below its other conjoined portions in Locus L. Carbon on all the Homo erectus fossils from Locus G, a circumscribed area of 1-meter diameter, earlier taken to indicate burning, cooking, and cannibalism, is here interpreted as detrital carbon deposited under water, perhaps the result of hyaenid caching behavior. Locus G records the close stratigraphic and horizontal association of stone artifacts with Homo erectus and other vertebrate skeletal elements, an association that is seen at other loci as well. Layer 4 of the excavation contains equid cranial bone previously interpreted to have been burned while fresh. We here document that Locus B Homo erectus, including Skull I, is stratigraphically associated with this evidence, but at some 10-12 m distance. Even though the presence of wood-stoked fires and hearths is not supported by geochemical results, evidence of fire at Locality 1 in the form of burned bone is confirmed. Contextual relationships of fossil skeletal elements, relationships of carnivore damage and stone tool cutmarks on bone, and evidence of the burning of fresh bone associated with Homo erectus and stone tools support a model of transient hominid scavenging aided by the use of fire at the large hyenid den that became Zhoukoudian Locality 1. Although the original excavation catalogue from Locality 1, as well as a significant number of fossils and stone artifacts, were lost during World War II, catalogue numbers on the many surviving specimens can be used to locate fossils and artifacts within the three-dimensional grid provided in this paper.

Chronology of primate discoveries in Myanmar: influences on the anthropoid origins debate.

The history of primate paleontology in Asia is long and complex, beginning with the first discoveries of fossil primates on the Indian subcontinent in the early 1830's. The first Eocene mammals from Asia were collected in Myanmar and described in 1916, while the first primates, Pondaungia and Amphipithecus, were described in 1927 and 1937, respectively, both from the Pondaung Formation in Myanmar. For the next 60 years, these two Pondaung taxa remained as the only known Eocene primates from Myanmar and one of the few records of Eocene primates from all of Asia. Taxonomically, Pondaungia and Amphipithecus were linked with a number of different groups, including archaic, hoofed ungulates (condylarths), adapiform primates, omomyid primates, and anthropoids. While no consensus existed, Pondaungia and Amphipithecus were most commonly compared with anthropoids. Beginning in the late 1990s, new primates were discovered in Myanmar, including smaller-bodied forms such as Bahinia and Myanmarpithecus. Also, new and better specimens of the larger-bodied Pondaungia and Amphipithecus began to appear, including the first cranial and postcranial fragments. Evaluations based on these new specimens, especially the postcrania, indicate that the two larger-bodied Myanmar taxa are adapiform primates that show their closest affinities to North American notharctines. The smaller-bodied taxa remain enigmatic, but may share their closest affinities with North American and Asian omomyid primates and Asian Tarsius. None of the known Asian primate taxa appear closely related to African anthropoids, which suggests that true anthropoids did not reach Asia until the latest Oligocene or earliest Miocene. These facts make an Asian origin for Anthropoidea unlikely. Additional and earlier evidence from both Asia and Africa is required before the ultimate origin of anthropoids can be determined. It appears possible that true anthropoids were an ancient radiation that may have been part of a Gondwanan (southern hemisphere) community that is, at present, poorly sampled and little understood.