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.

Modern human teeth unearthed from below the ∼128,000-year-old level at Punung, Java: A case highlighting the problem of recent intrusion in cave sediments.

The emergence of modern humans in the eastern edge of the Eurasian Continent is debated between two major models: early (∼130-70 ka) and late (∼50 ka) dispersal models. The former view is grounded mainly on the claims that several cave sites in Southeast Asia and southern China yielded modern human fossils of those early ages, but such reports have been disputed for the lack of direct dating of the human remains and insufficient documentation of stratigraphy and taphonomy. By tracing possible burial process and conducting direct dating for an early Late Pleistocene paleontological site of Punung III, East Java, we here report a case that demonstrates how unexpected intrusion of recent human remains into older stratigraphic levels could occur in cave sediments. This further highlights the need of direct dating and taphonomic assessment before accepting either model. We also emphasize that the state of fossilization of bones and teeth is a useful guide for initial screening of recent intrusion and should be reported particularly when direct dating is unavailable. Additionally, we provide a revised stratigraphy and faunal list of Punung III, a key site that defines the tropical rainforest Punung Fauna during the early Late Pleistocene of the region.

Homo floresiensis-like fossils from the early Middle Pleistocene of Flores.

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.

Earliest hominin occupation of Sulawesi, Indonesia.

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.

Age and context of the oldest known hominin fossils from Flores.

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.

Hominins on Flores, Indonesia, by one million years ago.

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.

High-resolution record of the Matuyama-Brunhes transition constrains the age of Javanese Homo erectus in the Sangiran dome, Indonesia.

A detailed paleomagnetic study conducted in the Sangiran area, Java, has provided a reliable age constraint on hominid fossil-bearing formations. A reverse-to-normal polarity transition marks a 7-m thick section across the Upper Tuff in the Bapang Formation. The transition has three short reversal episodes and is overlain by a thick normal polarity magnetozone that was fission-track dated to the Brunhes chron. This pattern closely resembles another high-resolution Matuyama-Brunhes (MB) transition record in an Osaka Bay marine core. In the Sangiran sediments, four successive transitional polarity fields lie just below the presumed main MB boundary. Their virtual geomagnetic poles cluster in the western South Pacific, partly overlapping the transitional virtual geomagnetic poles from Hawaiian and Canary Islands' lavas, which have a mean (40)Ar/(39)Ar age of 776 ± 2 ka. Thus, the polarity transition is unambiguously the MB boundary. A revised correlation of tuff layers in the Bapang Formation reveals that the hominid last occurrence and the tektite level in the Sangiran area are nearly coincident, just below the Upper Middle Tuff, which underlies the MB transition. The stratigraphic relationship of the tektite level to the MB transition in the Sangiran area is consistent with deep-sea core data that show that the meteorite impact preceded the MB reversal by about 12 ka. The MB boundary currently defines the uppermost horizon yielding Homo erectus fossils in the Sangiran area.

Early stone technology on Flores and its implications for Homo floresiensis.

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).

New reconstruction and morphological description of a Homo erectus cranium: skull IX (Tjg-1993.05) from Sangiran, Central Java.

Skull IX (Tjg-1993.05) was unearthed from the upper stratigraphic zone (Bapang-AG levels) of the hominin-bearing sequence in Sangiran. This remarkably complete cranial specimen of Homo erectus from the early Pleistocene of Java preserves substantial portions of the vault and face. However, the distortion present in the original reconstruction has hampered detailed documentation of its morphological characteristics. We here report a new reconstruction of Skull IX that successfully recovers the original morphology and significantly differs from previous reconstructions. Detailed morphological description and the results of initial comparative analyses based on this new reconstruction are provided. The endocranial volume of Skull IX was measured as 870 cc using micro-CT data. The neurocranium of Skull IX is slightly smaller than the so far recorded smallest cranium from this zone, suggesting this individual was female. In most, but not all, aspects of the cranial vault form, details of the external surface structures, and facial morphology, Skull IX exhibits numerous similarities to the other Bapang-AG H. erectus specimens, indicating that it belonged to the Bapang-AG H. erectus population. Drawing on the expanded fossil sample of this chronoregional H. erectus group, we discuss their evolutionary status, degree of sexual dimorphism, and facial morphological variation in Afro-Asian earlier Homo specimens.

Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area.

The chronology of the World Heritage Site of Sangiran in Indonesia is crucial for the understanding of human dispersals and settlement in Asia in the Early Pleistocene (before 780,000 years ago). It has been controversial, however, especially regarding the timing of the earliest hominin migration into the Sangiran region. We use a method of combining fission-track and uranium-lead dating and present key ages to calibrate the lower (older) Sangiran hominin-bearing horizons. We conclude that the first appearance datum for the Sangiran hominins is most likely ~1.3 million years ago and less than 1.5 million years ago, which is markedly later than the dates that have been widely accepted for the past two decades.

Brief communication: "Pathological" deformation in the skull of LB1, the type specimen of Homo floresiensis.

If the holotype of Homo floresiensis, LB1, suffered from a severe developmental pathology, this could undermine its status as the holotype of a new species. One of the proposed pathological indicators that still remains untested is asymmetric distortion in the skull of LB1 (Jacob et al.: Proc Natl Acad Sci USA 103 (2006) 13421-13426). Here, we present evidence that LB1 exhibits antemortem craniofacial deformities that are consistent with posterior deformational (positional) plagiocephaly. This is a relatively common condition in modern people with no serious associated health problems and does not represent a severe developmental abnormality in LB1.

Cranial morphology of Javanese Homo erectus: new evidence for continuous evolution, specialization, and terminal extinction.

Our current knowledge of the evolution of Homo during the early to middle Pleistocene is far from complete. This is not only because of the small number of fossil samples available, but also due to the scarcity of standardized datasets which are reliable in terms of landmark identification, interobserver error, and other distorting factors. This study aims to accurately describe the cranial morphological changes of H. erectus in Java using a standardized set of measurements taken by the authors from 18 adult crania from Sangiran, Trinil, Sambungmacan, and Ngandong. The identification of some obscure landmarks was aided by the use of micro-CT imaging. While recent studies tend to emphasize evolutionary conservatism in Javanese H. erectus, our results reinforce the theory that chronologically later groups experienced distinct morphological changes in a number of cranial traits. Some of these changes, particularly those related to brain size expansion, are similar to those observed for the genus Homo as a whole, whereas others are apparently unique specializations restricted to Javanese H. erectus. Such morphological specializations in Java include previously undescribed anteroposterior lengthening of the midcranial base and an anterior shift of the posterior temporal muscle, which might have influenced the morphology of the angular torus and supramastoid sulcus. Analyses of morphological variation indicate that the three crania from Sambungmacan variously fill the morphological gap between the chronologically earlier (Bapang-AG, Bapang Formation above the Grenzbank zone in Sangiran) and later (Ngandong) morphotypes of Java. At least one of the Bapang-AG crania, Sangiran 17, also exhibits a few characteristics which potentially indicate evolution toward the Ngandong condition. These strongly suggest the continuous, gradual morphological evolution of Javanese H. erectus from the Bapang-AG to Ngandong periods. The development of some unique features in later Javanese H. erectus supports the hypothesis that this Javanese lineage went extinct without making significant contributions to the ancestry of modern humans.

Hominid mandibular remains from Sangiran: 1952-1986 collection.

Eight hominid mandibular and associated dental remains discovered between 1952-1986 from the Early Pleistocene deposits of Sangiran, Central Java, are described. Although the specimens are surface finds, their original stratigraphic positions can be reasonably inferred on the basis of coincidental sources of information. These specimens significantly increase the dento-gnathic sample available for intensive morphological investigation of the earliest Javanese hominids [Kaifu et al., 2005].

Historical evidence of the 1936 Mojokerto skull discovery, East Java.

To resolve ambiguities in the literature, we detail the discovery history of the Mojokerto child's skull (Perning 1), employing letters, maps, photographs, reports, and newspaper accounts not previously used for this purpose. Andoyo, an experienced vertebrate-fossil collector with the Geological Survey of the Netherlands Indies, found the skull on February 13, 1936, while collecting for Johan Duyfjes, who had mapped the field area geologically. On February 18-19 Andoyo sent the fossil and a 1:25,000-topographic map showing the discovery point to Survey headquarters. The locality lies between Perning and Sumbertengu villages, approximately 10km northeast of Mojokerto city, East Java. G.H. Ralph von Koenigswald, Survey paleontologist, identified the specimen as Pithecanthropus and then named it Homo modjokertensis (it is now accepted as Homo erectus). Unfortunately he confused the discovery record in a March 28 newspaper article by characterizing the skull as a "surface find" [Dutch: oppervlaktevondst] while also attributing it to ancient beds. von Koenigswald probably had insufficient basis for either assertion, having not yet talked to Andoyo or Duyfjes. Eugene Dubois challenged von Koenigswald on the "surface-find" issue, Andoyo was consulted, and Duyfjes went to the site. Duyfjes and von Koenigswald then published scientific papers stating that the skull was unearthed 1m deep from a hill-slope outcrop of conglomeratic sandstone in Duyfjes' Pucangan formation. A cross section by Andoyo, which may show the Mojokerto site, also indicates a skull at 1m depth in conglomeratic sandstone. Photographs taken in 1936-1938 show a shallow pit at a single field location that fits Duyfjes' site description and is identified as the Mojokerto-skull site in 1940-1943 publications. By WWII the scientific community accepted the skull as an early hominid. Although von Koenigswald's "surface-find" comment remains a source of doubt in the record, we consider in situ discovery for the Mojokerto skull to be very likely.

Taxonomic affinities and evolutionary history of the Early Pleistocene hominids of Java: dentognathic evidence.

Temporal changes, within-group variation, and phylogenetic positions of the Early Pleistocene Javanese hominids remain unclear. Recent debate focused on the age of the oldest Javanese hominids, but the argument so far includes little morphological basis for the fossils. To approach these questions, we analyzed a comprehensive dentognathic sample from Sangiran, which includes most of the existing hominid mandibles and teeth from the Early Pleistocene of Java. The sample was divided into chronologically younger and older groups. We examined morphological differences between these chronological groups, and investigated their affinities with other hominid groups from Africa and Eurasia. The results indicated that 1) there are remarkable morphological differences between the chronologically younger and older groups of Java, 2) the chronologically younger group is morphologically advanced, showing a similar degree of dentognathic reduction to that of Middle Pleistocene Chinese H. erectus, and 3) the chronologically older group exhibits some features that are equally primitive as or more primitive than early H. erectus of Africa. These findings suggest that the evolutionary history of early Javanese H. erectus was more dynamic than previously thought. Coupled with recent discoveries of the earliest form of H. erectus from Dmanisi, Georgia, the primitive aspects of the oldest Javanese hominid remains suggest that hominid groups prior to the grade of ca. 1.8-1.5 Ma African early H. erectus dispersed into eastern Eurasia during the earlier Early Pleistocene, although the age of the Javanese hominids themselves is yet to be resolved. Subsequent periods of the Early Pleistocene witnessed remarkable changes in the Javanese hominid record, which are ascribed either to significant in situ evolution or replacement of populations.

Homo erectus calvarium from the Pleistocene of Java.

A Homo erectus calvarium [Sambungmacan 4 (Sm 4)] was recovered from Pleistocene sediments at Sambungmacan in central Java. Micro-computed tomography analysis shows a modern human-like cranial base flexion associated with a low platycephalic vault, implying that the evolution of human cranial globularity was independent of cranial base flexion. The overall morphology of Sm 4 is intermediate between that of earlier and later Javanese Homo erectus; apparent morphological specializations are more strongly expressed in the latter. This supports the hypothesis that later Pleistocene Javanese populations were substantially isolated and made minimal contributions to the ancestry of modern humans.