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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., Competing Interests: The authors have the following interests: The Kandiwal Aboriginal Corporation, an incorporated body encompassing members of the Kandiwal community (about 45 people) provided support during field work and knowledge about sites in their country, under a research agreement with the ARC (as a Linkage Partner). Slingair and Heliwork Pty Ltd provided a discount on air travel across the northwest Kimberley under a research agreement with the ARC (as a Linkage Partner). There are no patents, products in development or marketed products to declare. This does not alter our adherence to all the PLoS One policies on sharing data and materials, as detailed online in the guide for authors.

Homo erectus is the founding early hominin species of Island Southeast Asia, and reached Java (Indonesia) more than 1.5 million years ago 1,2 . 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 1933 3,4 , and are of the youngest, most-advanced form of H. erectus 5-8 . Despite the importance of the Ngandong fossils, the relationship between the fossils, terrace fill and ages have been heavily debated 9-14 . 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, 40 argon/ 39 argon ( 40 Ar/ 39 Ar) 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 Ngandong 5,15 . 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 conditions 3,17 . 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.

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

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.

Genetic evidence for anatomically modern humans (AMH) out of Africa before 75 thousand years ago (ka) and in island southeast Asia (ISEA) before 60 ka (93-61 ka) predates accepted archaeological records of occupation in the region. Claims that AMH arrived in ISEA before 60 ka (ref. 4) have been supported only by equivocal or non-skeletal evidence. AMH evidence from this period is rare and lacks robust chronologies owing to a lack of direct dating applications, poor preservation and/or excavation strategies and questionable taxonomic identifications. Lida Ajer is a Sumatran Pleistocene cave with a rich rainforest fauna associated with fossil human teeth. The importance of the site is unclear owing to unsupported taxonomic identification of these fossils and uncertainties regarding the age of the deposit, therefore it is rarely considered in models of human dispersal. Here we reinvestigate Lida Ajer to identify the teeth confidently and establish a robust chronology using an integrated dating approach. Using enamel-dentine junction morphology, enamel thickness and comparative morphology, we show that the teeth are unequivocally AMH. Luminescence and uranium-series techniques applied to bone-bearing sediments and speleothems, and coupled uranium-series and electron spin resonance dating of mammalian teeth, place modern humans in Sumatra between 73 and 63 ka. This age is consistent with biostratigraphic estimations, palaeoclimate and sea-level reconstructions, and genetic evidence for a pre-60 ka arrival of AMH into ISEA. Lida Ajer represents, to our knowledge, the earliest evidence of rainforest occupation by AMH, and underscores the importance of reassessing the timing and environmental context of the dispersal of modern humans out of Africa.

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., Competing Interests: The authors have the following interests: The Kandiwal Aboriginal Corporation, an incorporated body encompassing members of the Kandiwal community (about 45 people) provided support during field work and knowledge about sites in their country, under a research agreement with the ARC (as a Linkage Partner). Slingair and Heliwork Pty Ltd provided a discount on air travel across the northwest Kimberley under a research agreement with the ARC (as a Linkage Partner). There are no patents, products in development or marketed products to declare. This does not alter our adherence to all the PLOS ONE policies on sharing data and materials, as detailed online in the guide for authors.

Recent excavations at the early Middle Pleistocene site of Mata Menge in the So'a Basin of central Flores, Indonesia, have yielded hominin fossils attributed to a population ancestral to Late Pleistocene Homo floresiensis. Here we describe the age and context of the Mata Menge hominin specimens and associated archaeological findings. The fluvial sandstone layer from which the in situ fossils were excavated in 2014 was deposited in a small valley stream around 700 thousand years ago, as indicated by (40)Ar/(39)Ar and fission track dates on stratigraphically bracketing volcanic ash and pyroclastic density current deposits, in combination with coupled uranium-series and electron spin resonance dating of fossil teeth. Palaeoenvironmental data indicate a relatively dry climate in the So'a Basin during the early Middle Pleistocene, while various lines of evidence suggest the hominins inhabited a savannah-like open grassland habitat with a wetland component. The hominin fossils occur alongside the remains of an insular fauna and a simple stone technology that is markedly similar to that associated with Late Pleistocene H. floresiensis.

The evolutionary origin of Homo floresiensis, a diminutive hominin species previously known only by skeletal remains from Liang Bua in western Flores, Indonesia, has been intensively debated. It is a matter of controversy whether this primitive form, dated to the Late Pleistocene, evolved from early Asian Homo erectus and represents a unique and striking case of evolutionary reversal in hominin body and brain size within an insular environment. The alternative hypothesis is that H. floresiensis derived from an older, smaller-brained member of our genus, such as Homo habilis, or perhaps even late Australopithecus, signalling a hitherto undocumented dispersal of hominins from Africa into eastern Asia by two million years ago (2 Ma). Here we describe hominin fossils excavated in 2014 from an early Middle Pleistocene site (Mata Menge) in the So'a Basin of central Flores. These specimens comprise a mandible fragment and six isolated teeth belonging to at least three small-jawed and small-toothed individuals. Dating to ~0.7 Ma, these fossils now constitute the oldest hominin remains from Flores. The Mata Menge mandible and teeth are similar in dimensions and morphological characteristics to those of H. floresiensis from Liang Bua. The exception is the mandibular first molar, which retains a more primitive condition. Notably, the Mata Menge mandible and molar are even smaller in size than those of the two existing H. floresiensis individuals from Liang Bua. The Mata Menge fossils are derived compared with Australopithecus and H. habilis, and so tend to support the view that H. floresiensis is a dwarfed descendent of early Asian H. erectus. Our findings suggest that hominins on Flores had acquired extremely small body size and other morphological traits specific to H. floresiensis at an unexpectedly early time.

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.

Sulawesi is the largest and oldest island within Wallacea, a vast zone of oceanic islands separating continental Asia from the Pleistocene landmass of Australia and Papua (Sahul). By one million years ago an unknown hominin lineage had colonized Flores immediately to the south, and by about 50 thousand years ago, modern humans (Homo sapiens) had crossed to Sahul. On the basis of position, oceanic currents and biogeographical context, Sulawesi probably played a pivotal part in these dispersals. Uranium-series dating of speleothem deposits associated with rock art in the limestone karst region of Maros in southwest Sulawesi has revealed that humans were living on the island at least 40 thousand years ago (ref. 5). Here we report new excavations at Talepu in the Walanae Basin northeast of Maros, where in situ stone artefacts associated with fossil remains of megafauna (Bubalus sp., Stegodon and Celebochoerus) have been recovered from stratified deposits that accumulated from before 200 thousand years ago until about 100 thousand years ago. Our findings suggest that Sulawesi, like Flores, was host to a long-established population of archaic hominins, the ancestral origins and taxonomic status of which remain elusive.

Archaeologists have long been puzzled by the appearance in Europe ∼40-35 thousand years (kyr) ago of a rich corpus of sophisticated artworks, including parietal art (that is, paintings, drawings and engravings on immobile rock surfaces) and portable art (for example, carved figurines), and the absence or scarcity of equivalent, well-dated evidence elsewhere, especially along early human migration routes in South Asia and the Far East, including Wallacea and Australia, where modern humans (Homo sapiens) were established by 50 kyr ago. Here, using uranium-series dating of coralloid speleothems directly associated with 12 human hand stencils and two figurative animal depictions from seven cave sites in the Maros karsts of Sulawesi, we show that rock art traditions on this Indonesian island are at least compatible in age with the oldest European art. The earliest dated image from Maros, with a minimum age of 39.9 kyr, is now the oldest known hand stencil in the world. In addition, a painting of a babirusa ('pig-deer') made at least 35.4 kyr ago is among the earliest dated figurative depictions worldwide, if not the earliest one. Among the implications, it can now be demonstrated that humans were producing rock art by ∼40 kyr ago at opposite ends of the Pleistocene Eurasian world.

The carpals from the Homo floresiensis type specimen (LB1) lack features that compose the shared, derived complex of the radial side of the wrist in Neandertals and modern humans. This paper comprises a description and three-dimensional morphometric analysis of new carpals from at least one other individual at Liang Bua attributed to H. floresiensis: a right capitate and two hamates. The new capitate is smaller than that of LB1 but is nearly identical in morphology. As with capitates from extant apes, species of Australopithecus, and LB1, the newly described capitate displays a deeply-excavated nonarticular area along its radial aspect, a scaphoid facet that extends into a J-hook articulation on the neck, and a more radially-oriented second metacarpal facet; it also lacks an enlarged palmarly-positioned trapezoid facet. Because there is no accommodation for the derived, palmarly blocky trapezoid that characterizes Homo sapiens and Neandertals, this individual most likely had a plesiomorphically wedge-shaped trapezoid (like LB1). Morphometric analyses confirm the close similarity of the new capitate and that of LB1, and are consistent with previous findings of an overall primitive articular geometry. In general, hamate morphology is more conserved across hominins, and the H. floresiensis specimens fall at the far edge of the range of variation for H. sapiens in a number of metrics. However, the hamate of H. floresiensis is exceptionally small and exhibits a relatively long, stout hamulus lacking the oval-shaped cross-section characteristic of human and Neandertal hamuli (variably present in australopiths). Documentation of a second individual with primitive carpal anatomy from Liang Bua, along with further analysis of trapezoid scaling relative to the capitate in LB1, refutes claims that the wrist of the type specimen represents a modern human with pathology. In total, the carpal anatomy of H. floresiensis supports the hypothesis that the lineage leading to the evolution of this species originated prior to the cladogenetic event that gave rise to modern humans and Neandertals., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

This paper describes in detail the external morphology of LB1/1, the nearly complete and only known cranium of Homo floresiensis. Comparisons were made with a large sample of early groups of the genus Homo to assess primitive, derived, and unique craniofacial traits of LB1 and discuss its evolution. Principal cranial shape differences between H. floresiensis and Homo sapiens are also explored metrically. The LB1 specimen exhibits a marked reductive trend in its facial skeleton, which is comparable to the H. sapiens condition and is probably associated with reduced masticatory stresses. However, LB1 is craniometrically different from H. sapiens showing an extremely small overall cranial size, and the combination of a primitive low and anteriorly narrow vault shape, a relatively prognathic face, a rounded oval foramen that is greatly separated anteriorly from the carotid canal/jugular foramen, and a unique, tall orbital shape. Whereas the neurocranium of LB1 is as small as that of some Homo habilis specimens, it exhibits laterally expanded parietals, a weak suprameatal crest, a moderately flexed occipital, a marked facial reduction, and many other derived features that characterize post-habilis Homo. Other craniofacial characteristics of LB1 include, for example, a relatively narrow frontal squama with flattened right and left sides, a marked frontal keel, posteriorly divergent temporal lines, a posteriorly flexed anteromedial corner of the mandibular fossa, a bulbous lateral end of the supraorbital torus, and a forward protruding maxillary body with a distinct infraorbital sulcus. LB1 is most similar to early Javanese Homo erectus from Sangiran and Trinil in these and other aspects. We conclude that the craniofacial morphology of LB1 is consistent with the hypothesis that H. floresiensis evolved from early Javanese H. erectus with dramatic island dwarfism. However, further field discoveries of early hominin skeletal remains from Flores and detailed analyses of the finds are needed to understand the evolutionary history of this endemic hominin species., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Previous excavations at Mata Menge and Boa Lesa in the Soa Basin of Flores, Indonesia, recovered stone artefacts in association with fossilized remains of the large-bodied Stegodon florensis florensis. Zircon fission-track ages from these sites indicated that hominins had colonized the island by 0.88 +/- 0.07 million years (Myr) ago. Here we describe the contents, context and age of Wolo Sege, a recently discovered archaeological site in the Soa Basin that has in situ stone artefacts and that lies stratigraphically below Mata Menge and immediately above the basement breccias of the basin. We show using (40)Ar/(39)Ar dating that an ignimbrite overlying the artefact layers at Wolo Sege was erupted 1.02 +/- 0.02 Myr ago, providing a new minimum age for hominins on Flores. This predates the disappearance from the Soa Basin of 'pygmy' Stegodon sondaari and Geochelone spp. (giant tortoise), as evident at the nearby site of Tangi Talo, which has been dated to 0.90 +/- 0.07 Myr ago. It now seems that this extirpation or possible extinction event and the associated faunal turnover were the result of natural processes rather than the arrival of hominins. It also appears that the volcanic and fluvio-lacustrine deposits infilling the Soa Basin may not be old enough to register the initial arrival of hominins on the island.

Excavations at Liang Bua, a limestone cave on the island of Flores, East Indonesia, have yielded a well-dated archaeological and faunal sequence spanning the last 95k.yr., major climatic fluctuations, and two human species -H. floresiensis from 95 to 17k.yr.(1), and modern humans from 11k.yr. to the present. The faunal assemblage comprises well-preserved mammal, bird, reptile and mollusc remains, including examples of island gigantism in small mammals and the dwarfing of large taxa. Together with evidence from Early-Middle Pleistocene sites in the Soa Basin, it confirms the long-term isolation, impoverishment, and phylogenetic continuity of the Flores faunal community. The accumulation of Stegodon and Komodo dragon remains at the site in the Pleistocene is attributed to Homo floresiensis, while predatory birds, including an extinct species of owl, were largely responsible for the accumulation of the small vertebrates. The disappearance from the sequence of the two large-bodied, endemic mammals, Stegodon florensis insularis and Homo floresiensis, was associated with a volcanic eruption at 17 ka and precedes the earliest evidence for modern humans, who initiated use of mollusc and shell working, and began to introduce a range of exotic animals to the island. Faunal introductions during the Holocene included the Sulawesi warty pig (Sus celebensis) at about 7ka, followed by the Eurasian pig (Sus scrofa), Long-tailed macaque, Javanese porcupine, and Masked palm civet at about 4ka, and cattle, deer, and horse - possibly by the Portuguese within historic times. The Holocene sequence at the site also documents local faunal extinctions - a result of accelerating human population growth, habitat loss, and over-exploitation.

Excavations at Liang Bua, on the Indonesian island of Flores, have yielded a stratified sequence of stone artifacts and faunal remains spanning the last 95k.yr., which includes the skeletal remains of two human species, Homo sapiens in the Holocene and Homo floresiensis in the Pleistocene. This paper summarizes and focuses on some of the evidence for Homo floresiensis in context, as presented in this Special Issue edition of the Journal of Human Evolution and elsewhere. Attempts to dismiss the Pleistocene hominins (and the type specimen LB1 in particular) as pathological pygmy humans are not compatible with detailed analyses of the skull, teeth, brain endocast, and postcranium. We initially concluded that H. floresiensis may have evolved by insular dwarfing of a larger-bodied hominin species over 880k.yr. or more. However, recovery of additional specimens and the numerous primitive morphological traits seen throughout the skeleton suggest instead that it is more likely to be a late representative of a small-bodied lineage that exited Africa before the emergence of Homo erectus sensu lato. Homo floresiensis is clearly not an australopithecine, but does retain many aspects of anatomy (and perhaps behavior) that are probably plesiomorphic for the genus Homo. We also discuss some of the other implications of this tiny, endemic species for early hominin dispersal and evolution (e.g., for the "Out of Africa 1" paradigm and more specifically for colonizing Southeast Asia), and we present options for future research in the region.

Bones of the lower extremity have been recovered for up to nine different individuals of Homo floresiensis - LB1, LB4, LB6, LB8, LB9, LB10, LB11, LB13, and LB14. LB1 is represented by a bony pelvis (damaged but now repaired), femora, tibiae, fibulae, patellae, and numerous foot bones. LB4/2 is an immature right tibia lacking epiphyses. LB6 includes a fragmentary metatarsal and two pedal phalanges. LB8 is a nearly complete right tibia (shorter than that of LB1). LB9 is a fragment of a hominin femoral diaphysis. LB10 is a proximal hallucal phalanx. LB11 includes pelvic fragments and a fragmentary metatarsal. LB13 is a patellar fragment, and LB14 is a fragment of an acetabulum. All skeletal remains recovered from Liang Bua were extremely fragile, and some were badly damaged when they were removed temporarily from Jakarta. At present, virtually all fossil materials have been returned, stabilized, and hardened. These skeletal remains are described and illustrated photographically. The lower limb skeleton exhibits a uniquely mosaic pattern, with many primitive-like morphologies; we have been unable to find this combination of ancient and derived (more human-like) features in either healthy or pathological modern humans, regardless of body size. Bilateral asymmetries are slight in the postcranium, and muscle markings are clearly delineated on all bones. The long bones are robust, and the thickness of their cortices is well within the ranges seen in healthy modern humans. LB1 is most probably a female based on the shape of her greater sciatic notch, and the marked degree of lateral iliac flaring recalls that seen in australopithecines such as "Lucy" (AL 288-1). The metatarsus has a human-like robusticity formula, but the proximal pedal phalanges are relatively long and robust (and slightly curved). The hallux is fully adducted, but we suspect that a medial longitudinal arch was absent.

This study examines trends in stone tool reduction technology at Liang Bua, Flores, Indonesia, where excavations have revealed a stratified artifact sequence spanning 95k.yr. The reduction sequence practiced throughout the Pleistocene was straightforward and unchanging. Large flakes were produced off-site and carried into the cave where they were reduced centripetally and bifacially by four techniques: freehand, burination, truncation, and bipolar. The locus of technological complexity at Liang Bua was not in knapping products, but in the way techniques were integrated. This reduction sequence persisted across the Pleistocene/Holocene boundary with a minor shift favoring unifacial flaking after 11ka. Other stone-related changes occurred at the same time, including the first appearance of edge-glossed flakes, a change in raw material selection, and more frequent fire-induced damage to stone artifacts. Later in the Holocene, technological complexity was generated by "adding-on" rectangular-sectioned stone adzes to the reduction sequence. The Pleistocene pattern is directly associated with Homo floresiensis skeletal remains and the Holocene changes correlate with the appearance of Homo sapiens. The one reduction sequence continues across this hominin replacement.

Liang Bua, in Flores, Indonesia, was formed as a subterranean chamber over 600ka. From this time to the present, a series of geomorphic events influenced the structure of the cave and cave deposits, creating a complex stratigraphy. Within these deposits, nine main sedimentary units have been identified. The stratigraphic relationships between these units provide the evidence needed to reconstruct the geomorphic history of the cave. This history was dominated by water action, including slope wash processes, channel formation, pooling of water, and flowstone precipitation, which created waterfalls, cut-and-fill stratigraphy, large pools of water, and extensive flowstone cappings. The reconstructed sequence of events over the last 190k.yr. has been summarized by a series of time slices that demonstrate the nature of the occupational environment in Liang Bua. The earliest artifacts at the site, dated to approximately 190ka, testify to hominin presence in the area, but the reconstructions suggest that occupation of the cave itself may not have been possible until after approximately 100ka. At approximately 95ka, channel erosion of a basal unit, which displays evidence of deposition in a pond environment, created a greater relief on the cave floor, and formed remanent areas of higher ground that later became a focus for hominin occupation from 74-61ka by the west wall and in the center of the cave, and from approximately 18-17ka by the east wall. These zones have been identified according to the sloping nature of the stratigraphy and the distribution of artifacts, and their locations have implications for the archaeological interpretation of the site.

Several bones of the upper extremity were recovered during excavations of Late Pleistocene deposits at Liang Bua, Flores, and these have been attributed to Homo floresiensis. At present, these upper limb remains have been assigned to six different individuals - LB1, LB2, LB3, LB4, LB5, and LB6. Several of these bones are complete or nearly so, but some are quite fragmentary. All skeletal remains recovered from Liang Bua were extremely fragile, but have now been stabilized and hardened in the laboratory in Jakarta. They are now curated in museum-quality containers at the National Research and Development Centre for Archaeology in Jakarta, Indonesia. These skeletal remains are described and illustrated photographically. The upper limb presents a unique mosaic of derived (human-like) and primitive morphologies, the combination of which is never found in either healthy or pathological modern humans.

Excavations at Liang Bua, Flores, Indonesia, have yielded evidence for an endemic human species, Homo floresiensis, a population that occupied the cave between approximately 95-17ka. This discovery has major implications for early hominin evolution and dispersal in Africa and Asia, attracting worldwide interest. This preface describes the rationale for the excavations in historical, geographical, and wider research contexts, as well as the methods used. It also introduces the other papers on aspects of Liang Bua research that feature in this edition of the Journal of Human Evolution.

The rapidly changing landscape of the eastern Indonesian archipelago has evolved at a pace dictated by its tropical climate and its geological and tectonic history. This has produced accelerated karstification, flights of alluvial terraces, and complex, multi-level cave systems. These cave systems sometimes contain a wealth of archaeological evidence, such as the almost complete skeleton of Homo floresiensis found at the site of Liang Bua in western Flores, but this information can only be understood in the context of the geomorphic history of the cave, and the more general geological, tectonic, and environmental histories of the river valley and region. Thus, a reconstruction of the landscape history of the Wae Racang valley using speleothems, geological structure, tectonic uplift, karst, cave, and terrace development, provides the necessary evidence to determine the formation, age, evolution, and influences on the site. This evidence suggests that Liang Bua was formed as two subterranean chambers approximately 600ka, but could not be occupied until approximately 190ka when the Wae Racang wandered to the southern side of the valley, exposing the chamber and depositing alluvial deposits containing artifacts. During the next approximately 190k.yr., the chambers coalesced and evolved into a multi-level and interconnected cave that was subjected to channel erosion and pooling events by the development of sinkholes. The domed morphology of the front chamber accumulated deep sediments containing well stratified archaeological and faunal remains, but ponded water in the chamber further prevented hominin use of the cave until approximately 100ka. These chambers were periodically influenced by river inundation and volcanic activity, whereas the area outside the cave was greatly influenced by glacial phases, which changed humid forest environments into grassland environments. This combined evidence has important implications for the archaeological interpretation of the site.

The announcement of a new species, Homo floresiensis, a primitive hominin that survived until relatively recent times is an enormous challenge to paradigms of human evolution. Until this announcement, the dominant paradigm stipulated that: 1) only more derived hominins had emerged from Africa, and 2) H. sapiens was the only hominin since the demise of Homo erectus and Homo neanderthalensis. Resistance to H. floresiensis has been intense, and debate centers on two sets of competing hypotheses: 1) that it is a primitive hominin, and 2) that it is a modern human, either a pygmoid form or a pathological individual. Despite a range of analytical techniques having been applied to the question, no resolution has been reached. Here, we use cladistic analysis, a tool that has not, until now, been applied to the problem, to establish the phylogenetic position of the species. Our results produce two equally parsimonious phylogenetic trees. The first suggests that H. floresiensis is an early hominin that emerged after Homo rudolfensis (1.86Ma) but before H. habilis (1.66Ma, or after 1.9Ma if the earlier chronology for H. habilis is retained). The second tree indicates H. floresiensis branched after Homo habilis.

Earlier observations of the virtual endocast of LB1, the type specimen for Homo floresiensis, are reviewed, extended, and interpreted. Seven derived features of LB1's cerebral cortex are detailed: a caudally-positioned occipital lobe, lack of a rostrally-located lunate sulcus, a caudally-expanded temporal lobe, advanced morphology of the lateral prefrontal cortex, shape of the rostral prefrontal cortex, enlarged gyri in the frontopolar region, and an expanded orbitofrontal cortex. These features indicate that LB1's brain was globally reorganized despite its ape-sized cranial capacity (417cm(3)). Neurological reorganization may thus form the basis for the cognitive abilities attributed to H. floresiensis. Because of its tiny cranial capacity, some workers think that LB1 represents a Homo sapiens individual that was afflicted with microcephaly, or some other pathology, rather than a new species of hominin. We respond to concerns about our earlier study of microcephalics compared with normal individuals, and reaffirm that LB1 did not suffer from this pathology. The intense controversy about LB1 reflects an older continuing dispute about the relative evolutionary importance of brain size versus neurological reorganization. LB1 may help resolve this debate and illuminate constraints that governed hominin brain evolution.

Background: The largest living lizard species, Varanus komodoensis Ouwens 1912, is vulnerable to extinction, being restricted to a few isolated islands in eastern Indonesia, between Java and Australia, where it is the dominant terrestrial carnivore. Understanding how large-bodied varanids responded to past environmental change underpins long-term management of V. komodoensis populations., Methodology/principal Findings: We reconstruct the palaeobiogeography of Neogene giant varanids and identify a new (unnamed) species from the island of Timor. Our data reject the long-held perception that V. komodoensis became a giant because of insular evolution or as a specialist hunter of pygmy Stegodon. Phyletic giantism, coupled with a westward dispersal from mainland Australia, provides the most parsimonious explanation for the palaeodistribution of V. komodoensis and the newly identified species of giant varanid from Timor. Pliocene giant varanid fossils from Australia are morphologically referable to V. komodoensis suggesting an ultimate origin for V. komodoensis on mainland Australia (>3.8 million years ago). Varanus komodoensis body size has remained stable over the last 900,000 years (ka) on Flores, a time marked by major faunal turnovers, extinction of the island's megafauna, the arrival of early hominids by 880 ka, co-existence with Homo floresiensis, and the arrival of modern humans by 10 ka. Within the last 2000 years their populations have contracted severely., Conclusions/significance: Giant varanids were once a ubiquitous part of Subcontinental Eurasian and Australasian faunas during the Neogene. Extinction played a pivotal role in the reduction of their ranges and diversity throughout the late Quaternary, leaving only V. komodoensis as an isolated long-term survivor. The events over the last two millennia now threaten its future survival.

Homo floresiensis is an endemic hominin species that occupied Liang Bua, a limestone cave on Flores in eastern Indonesia, during the Late Pleistocene epoch. The skeleton of the type specimen (LB1) of H. floresiensis includes a relatively complete left foot and parts of the right foot. These feet provide insights into the evolution of bipedalism and, together with the rest of the skeleton, have implications for hominin dispersal events into Asia. Here we show that LB1's foot is exceptionally long relative to the femur and tibia, proportions never before documented in hominins but seen in some African apes. Although the metatarsal robusticity sequence is human-like and the hallux is fully adducted, other intrinsic proportions and pedal features are more ape-like. The postcranial anatomy of H. floresiensis is that of a biped, but the unique lower-limb proportions and surprising combination of derived and primitive pedal morphologies suggest kinematic and biomechanical differences from modern human gait. Therefore, LB1 offers the most complete glimpse of a bipedal hominin foot that lacks the full suite of derived features characteristic of modern humans and whose mosaic design may be primitive for the genus Homo. These new findings raise the possibility that the ancestor of H. floresiensis was not Homo erectus but instead some other, more primitive, hominin whose dispersal into southeast Asia is still undocumented.

The Punung Fauna is a key component in the biostratigraphic sequence of Java. It represents the most significant faunal turnover on the island in the last 1.5 million years, when Stegodon and other archaic mammal species characteristic of earlier Faunal stages were replaced by a fully modern fauna that included rainforest-dependent species such as Pongo pygmaeus (orangutan). Here, we report the first numerical ages for the Punung Fauna obtained by luminescence and uranium-series dating of the fossil-bearing deposits and associated flowstones. The Punung Fauna contained in the dated breccia is of early Last Interglacial age (between 128+/-15 and 118+/-3 ka). This result has implications for the age of the preceding Ngandong Fauna, including Homo erectus remains found in the Ngandong Terrace, and for the timing of Homo sapiens arrival in Southeast Asia, in view of claims for a modern human tooth associated with the Punung breccia.

The holotype of Homo floresiensis, diminutive hominins with tiny brains living until 12,000 years ago on the island of Flores, is a partial skeleton (LB1) that includes a partial clavicle (LB1/5) and a nearly complete right humerus (LB1/50). Although the humerus appears fairly modern in most regards, it is remarkable in displaying only 110 degrees of humeral torsion, well below modern human average values. Assuming a modern human shoulder configuration, such a low degree of humeral torsion would result in a lateral set to the elbow. Such an elbow joint would function more nearly in a frontal than in a sagittal plane, and this is certainly not what anyone would have predicted for a tool-making Pleistocene hominin. We argue that Homo floresiensis probably did not have a modern human shoulder configuration: the clavicle was relatively short, and we suggest that the scapula was more protracted, resulting in a glenoid fossa that faced anteriorly rather than laterally. A posteriorly directed humeral head was therefore appropriate for maintaining a normally functioning elbow joint. Similar morphology in the Homo erectus Nariokotome boy (KNM-WT 15000) suggests that this shoulder configuration may represent a transitional stage in pectoral girdle evolution in the human lineage.

Whether the Late Pleistocene hominin fossils from Flores, Indonesia, represent a new species, Homo floresiensis, or pathological modern humans has been debated. Analysis of three wrist bones from the holotype specimen (LB1) shows that it retains wrist morphology that is primitive for the African ape-human clade. In contrast, Neandertals and modern humans share derived wrist morphology that forms during embryogenesis, which diminishes the probability that pathology could result in the normal primitive state. This evidence indicates that LB1 is not a modern human with an undiagnosed pathology or growth defect; rather, it represents a species descended from a hominin ancestor that branched off before the origin of the clade that includes modern humans, Neandertals, and their last common ancestor.

Because the cranial capacity of LB1 (Homo floresiensis) is only 417 cm(3), some workers propose that it represents a microcephalic Homo sapiens rather than a new species. This hypothesis is difficult to assess, however, without a clear understanding of how brain shape of microcephalics compares with that of normal humans. We compare three-dimensional computed tomographic reconstructions of the internal braincases (virtual endocasts that reproduce details of external brain morphology, including cranial capacities and shape) from a sample of 9 microcephalic humans and 10 normal humans. Discriminant and canonical analyses are used to identify two variables that classify normal and microcephalic humans with 100% success. The classification functions classify the virtual endocast from LB1 with normal humans rather than microcephalics. On the other hand, our classification functions classify a pathological H. sapiens specimen that, like LB1, represents an approximately 3-foot-tall adult female and an adult Basuto microcephalic woman that is alleged to have an endocast similar to LB1's with the microcephalic humans. Although microcephaly is genetically and clinically variable, virtual endocasts from our highly heterogeneous sample share similarities in protruding and proportionately large cerebella and relatively narrow, flattened orbital surfaces compared with normal humans. These findings have relevance for hypotheses regarding the genetic substrates of hominin brain evolution and may have medical diagnostic value. Despite LB1's having brain shape features that sort it with normal humans rather than microcephalics, other shape features and its small brain size are consistent with its assignment to a separate species.

In the Soa Basin of central Flores, eastern Indonesia, stratified archaeological sites, including Mata Menge, Boa Lesa and Kobatuwa (Fig. 1), contain stone artefacts associated with the fossilized remains of Stegodon florensis, Komodo dragon, rat and various other taxa. These sites have been dated to 840-700 kyr bp (thousand years before present). The authenticity of the Soa Basin artefacts and their provenance have been demonstrated by previous work, but to quell lingering doubts, here we describe the context, attributes and production modes of 507 artefacts excavated at Mata Menge. We also note specific similarities, and apparent technological continuity, between the Mata Menge stone artefacts and those excavated from Late Pleistocene levels at Liang Bua cave, 50 km to the west. The latter artefacts, dated to between 95-74 and 12 kyr ago, are associated with the remains of a dwarfed descendent of S. florensis, Komodo dragon, rat and a small-bodied hominin species, Homo floresiensis, which had a brain size of about 400 cubic centimetres. The Mata Menge evidence negates claims that stone artefacts associated with H. floresiensis are so complex that they must have been made by modern humans (Homo sapiens).

Homo floresiensis was recovered from Late Pleistocene deposits on the island of Flores in eastern Indonesia, but has the stature, limb proportions and endocranial volume of African Pliocene Australopithecus. The holotype of the species (LB1), excavated in 2003 from Liang Bua, consisted of a partial skeleton minus the arms. Here we describe additional H. floresiensis remains excavated from the cave in 2004. These include arm bones belonging to the holotype skeleton, a second adult mandible, and postcranial material from other individuals. We can now reconstruct the body proportions of H. floresiensis with some certainty. The finds further demonstrate that LB1 is not just an aberrant or pathological individual, but is representative of a long-term population that was present during the interval 95-74 to 12 thousand years ago. The excavation also yielded more evidence for the depositional history of the cave and for the behavioural capabilities of H. floresiensis, including the butchery of Stegodon and use of fire.

The brain of Homo floresiensis was assessed by comparing a virtual endocast from the type specimen (LB1) with endocasts from great apes, Homo erectus, Homo sapiens, a human pygmy, a human microcephalic, specimen number Sts 5 (Australopithecus africanus), and specimen number WT 17000 (Paranthropus aethiopicus). Morphometric, allometric, and shape data indicate that LB1 is not a microcephalic or pygmy. LB1's brain/body size ratio scales like that of an australopithecine, but its endocast shape resembles that of Homo erectus. LB1 has derived frontal and temporal lobes and a lunate sulcus in a derived position, which are consistent with capabilities for higher cognitive processing.

Excavations at Liang Bua, a large limestone cave on the island of Flores in eastern Indonesia, have yielded evidence for a population of tiny hominins, sufficiently distinct anatomically to be assigned to a new species, Homo floresiensis. The finds comprise the cranial and some post-cranial remains of one individual, as well as a premolar from another individual in older deposits. Here we describe their context, implications and the remaining archaeological uncertainties. Dating by radiocarbon (14C), luminescence, uranium-series and electron spin resonance (ESR) methods indicates that H. floresiensis existed from before 38,000 years ago (kyr) until at least 18 kyr. Associated deposits contain stone artefacts and animal remains, including Komodo dragon and an endemic, dwarfed species of Stegodon. H. floresiensis originated from an early dispersal of Homo erectus (including specimens referred to as Homo ergaster and Homo georgicus) that reached Flores, and then survived on this island refuge until relatively recently. It overlapped significantly in time with Homo sapiens in the region, but we do not know if or how the two species interacted.

Currently, it is widely accepted that only one hominin genus, Homo, was present in Pleistocene Asia, represented by two species, Homo erectus and Homo sapiens. Both species are characterized by greater brain size, increased body height and smaller teeth relative to Pliocene Australopithecus in Africa. Here we report the discovery, from the Late Pleistocene of Flores, Indonesia, of an adult hominin with stature and endocranial volume approximating 1 m and 380 cm3, respectively--equal to the smallest-known australopithecines. The combination of primitive and derived features assigns this hominin to a new species, Homo floresiensis. The most likely explanation for its existence on Flores is long-term isolation, with subsequent endemic dwarfing, of an ancestral H. erectus population. Importantly, H. floresiensis shows that the genus Homo is morphologically more varied and flexible in its adaptive responses than previously thought.