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Pleistocene vertebrate faunas have long been known from the island of Timor but have rarely been studied. In particular, dwarf elephant-like proboscideans such as Stegodon were once widespread across the island, including to the Atambua Basin, but the cause of their extinction is unknown. Here, we document the fossil Pleistocene fauna of this basin with their palaeoenvironmental context. We also investigate the possibility of human-Stegodon chronological overlap. Our methods include field survey, excavation, pollen analysis, and U-series dating. Palaeoenvironmental data based on geomorphology and hydrological information show a transition from marine to fluvial depositional conditions in the basin during Pleistocene. Pollen data supports this interpretation and shows a clear shift from mangrove to terrestrial vegetation types. Palaeoclimatic conditions after the transition are characterized by a high humidity with high-flow water across the basin. Fossils of vertebrate faunas are restricted to fluvially-derived sandstone units. Meanwhile lithic artefacts were recorded widely across the basin, in the from of surface finds. There is so far no example of direct association of lithic artefacts with extinct Pleistocene faunas. Direct dating of the vertebrate fossils suggest they came from the Middle Pleistocene or earlier, and consequently pre-date the oldest current records of modern human existence on the island ~45,000 years ago.

Objectives Gigantopithecus blacki, the largest hominoid known, is one of the representative Pleistocene mammals in southern China and northern Southeast Asia. Here we investigate the feeding ecology of G. blacki in its core habitat (Guangxi, Southern China) during the early Early Pleistocene, which was the early period in its evolution. Materials and methods The stable isotopic (C, O) analysis of tooth enamel of the fauna associated with G. blacki (n = 58), including the largest number of G. blacki teeth (n = 12) to date from the Liucheng Gigantopithecus Cave (~2 Ma), Guangxi, China, is undertaken. Results The δ13 C values of Liucheng fauna range from -12.9 to -19.0‰ with an average of -16.1 ± 1.3‰ (n = 58) and the δ18 O values range from -4.3 to -9.6‰ with an average of -6.9 ± 1.2‰ (n = 58). The δ13 C values of G. blacki range from -15.9‰ to -17.0‰ with an average of -16.5 ± 0.4‰ (n = 12), and the δ18 O values vary from -5.9‰ to -7.5‰ with an average of -6.6 ± 0.5‰ (n = 12). Conclusions The isotopic data show Guangxi was characterized by closed C3 forest and humid climate in the early Early Pleistocene. Niche partitioning is found among G. blacki, Sinomastodon, Ailuropoda and Stegodon, the typical megafauna in South China in the early Early Pleistocene. This could be one of the important factors for them to co-exist until the Middle Pleistocene. Smallest isotopic variations of G. blacki are found compared with those of contemporary animals, indicating a conservative foraging ecology i.e., limited foraging area and/or narrow dietary flexibility. Furthermore, the more confined foraging ecology of G. blacki is also seen in comparison with fossil and extant large-bodied primates. However, the unique dietary pattern of G. blacki does not seem to have hindered its survival. The environment in Guangxi during the early Early Pleistocene offered the suitable conditions for G. blacki to become one of the typical species in the faunal assemblages.

A new Pleistocene site in the Pabbi Hills of Pakistan, and deposits to the east near Bhimbar provide important additions to the Upper Siwalik fauna of South Asia. The mammalian fauna of 10 species from Panjan Sher Shahana at the southwestern tip of the Pabbi Hills is a rich mammalian assemblage dating to the early to Middle Pleistocene of the Upper Siwalik Subgroup. The Panjan Sher Shahana assemblage shows strong resemblance to other Pleistocene faunas of the Siwalik Group from the Indian subcontinent. The new site is dominated by Bovidae and Equus. Among megafauna, two species of Elephas, commonly found in the Upper Siwaliks, are associated with the uncommon Stegodon in Pabbi Hills. These and other elements of the Panjan Sher Shahana fauna are treated briefly here. The Late Neogene deposits of Bhimbar, 30 km east of Jhelum, appear to represent a similar succession equivalent to the Pleistocene (“Pinjor”) fauna and produce important new elements, including a rare record of the bamboo rat Anepsirhizomys opdykei.

Stegodont fossils have been recovered from many paleoanthropological sites in South China and Southeast Asia, marking them as one of the most important constituents in Pleistocene mammalian faunas of Asia. Yet very limited taphonomic research has been conducted so far concerning the context of their present in paleoanthropological sites. Recently, a large number of stegodont fossils have been found at the Yumidong Cave site in Wushan County, Chongqing Municipality, providing us with opportunities not only for morphological observation, but also taphonomic analysis. The studied materials include 94 cranial remains (jaws and teeth) and 54 remains of the postcrania from both the upper and lower cultural layers at Yumidong, which make up about 18.5% of the total identified specimens. Morphologically, the ridge-crest formulae, the number of mammillae and the measurements of most these specimens conform to Stegodon orientalis . Taphonomically, traces of natural activities, such as water transportation and bite marks of animals, are very few on the surface of these Stegodon remains; meanwhile, these fossils are unearthed associated with paleolithic tools in the Yumidong Cave site, among which at least three bone tools are made of incisors of Stegodon orientalis . These observations indicate that ancient human activity was the dominant cause in the accumulation of this Stegodon fossil assemblage. Due to the peculiarity of tooth replacement and attrition of these proboscideans, we are able to attribute most of each individual specimen, except two fragments of M2/3, as belonging to a distinct individual animal. The analysis of mortality age profiles shows that neonate and juvenile individuals account for more than 85% among the Stegodon fossils, indicating a type of selective mortality of immature individuals. In the schematic typology of living and extinct proboscidean age profiles, the type B of Haynes is dominated by the youngest age group which may result from hunting either by carnivores or humans. The skeletal element profiles show that the cranial and foot elements are the most numerous category. This phenomenon is always considered as a resultant taphonomic effect of scavenger behavior. But given the unique anatomic structure of proboscidean, hunting behavior cannot be ruled out here; in other words, the ancient human might hunt immature stegodonts but take a strategy of transporting the cranial and foot parts of carcass to their base camp, for these parts of stegodont contain more fat than other animals and are easy to carry. Irrespective of scavenging or hunting, the ancient humans which occupied the Yumidong Cave site appear to have preferred neonate and juvenile stegodonts rather than the adults, which were not easy to hunt. Alternatively, meat from neonate and juvenile Stegodon may have had better taste and nutritional value. Perhaps, a combination of these two factors may have motivated the highly selective exploitation of juvenile Stegodon by early humans at the Yumidong site.

The Middle Pleistocene fluvial channel site of the Upper Fossil-bearing Interval at Mata Menge in the So’a Basin, Flores, Indonesia, has yielded the earliest fossil evidence for Homo floresiensis in association with stone artefacts and fossils of highly endemic insular fauna, including Stegodon, giant rats, crocodiles, Komodo dragons, and various birds. A preliminary taphonomic review of the fossil material here aimed to provide additional context for the hominin remains in this bonebed. Analysis was performed on two subsets of material from the same fluvial sandstone layer. Subset 1 comprised all material from two adjacent one-metre square quadrants (n = 91), and Subset 2 all Stegodon long limb bones excavated from the same layer (n = 17). Key analytical parameters included species and skeletal element identification; fossil size measurements and fragmentation; weathering stages; bone fracture characteristics; and other biological and geological bone surface modifications. Analysis of Subset 1 material identified a highly fragmented assemblage with a significant bias towards Stegodon. A large portion of these bones were likely fractured by trampling prior to entering the fluvial channel and were transported away from the death-site, undergoing surface modification causing rounding. Subset 2 material was less likely to have been transported far based on its limited susceptibility to fluvial transport. There was no significant difference in weathering for the long limb bones and fragments, with the highest portion exhibiting Stage 2 weathering, indicating that prior to final burial, all material was exposed to prolonged periods of surface exposure. Approximately 10% of all material have characteristics of fracturing on fresh bone, contributing to the taphonomic context for this bonebed; however insufficient evidence was found for anthropogenic modification.

The new fossil teeth of Stegolophodon (Proboscidea, Mammalia) were discovered from the terrestrial sediments of the Irrawaddy Formation, in Kyauksaungsan and Tebingan areas, Magway Region, central Myanmar. A new Stegolophodon species is established in this paper. Stegolophodon iravaticus shares the characters of the advanced Stegolophodon and the primitive Stegodon. The M3 of S. iravaticus is related to that of the molar of Sand Pit in Tha Chang, Thailand and that of from Dhedari, Pakistan which is included in the Stegolophodon group 6 of Saegusa et al. (2005).

Liang Bua, the type locality of Homo floresiensis, is a limestone cave located in the western part of the Indonesian island of Flores. The relatively continuous stratigraphic sequence of the site spans the past ∼190 kyr and contains ∼275,000 taxonomically identifiable vertebrate skeletal elements, ∼80% of which belong to murine rodent taxa (i.e., rats). Six described genera are present at Liang Bua (Papagomys, Spelaeomys, Hooijeromys, Komodomys, Paulamys, and Rattus), one of which, Hooijeromys, is newly recorded in the site deposits, being previously known only from Early to Middle Pleistocene sites in central Flores. Measurements of the proximal femur (n = 10,212) and distal humerus (n = 1186) indicate five murine body size classes ranging from small (mouse-sized) to giant (common rabbit-sized) are present. The proportions of these five classes across successive stratigraphic units reveal two major changes in murine body size distribution due to significant shifts in the abundances of more open habitat-adapted medium-sized murines versus more closed habitat-adapted smaller-sized ones. One of these changes suggests a modest increase in available open habitats occurred ∼3 ka, likely the result of anthropogenic changes to the landscape related to farming by modern human populations. The other and more significant change occurred ∼60 ka suggesting a rapid shift from more open habitats to more closed conditions at this time. The abrupt reduction of medium-sized murines, along with the disappearance of H. floresiensis, Stegodon florensis insularis (an extinct proboscidean), Varanus komodoensis (Komodo dragon), Leptoptilos robustus (giant marabou stork), and Trigonoceps sp. (vulture) at Liang Bua ∼60–50 ka, is likely the consequence of these animals preferring and tracking more open habitats to elsewhere on the island. If correct, then the precise timing and nature of the extinction of H. floresiensis and its contemporaries must await new discoveries at Liang Bua or other as yet unexcavated sites on Flores.

Elephas maximus and Stegodon orientalis were two keystone proboscideans in southern Asia that coexisted mainly after the Middle Pleistocene in many regions. The long-term paleoecology and possible foraging competition of these two species have not been intensively investigated yet. Here, we applied stable isotope (C, O) analysis to the tooth enamel of coexisting Elephas maximus, Stegodon orientalis, and other associated mammalian species in Quzai Cave, southern China, dated to the early Late Pleistocene, to explore their paleoenvironmental context and foraging ecology. The δ13C values of Elephas maximus were widely distributed between −17.9‰ and −11.9‰ (n = 10), while Stegodon orientalis δ13C values ranged from −16.7‰ to −14.7‰ (n = 7). These results suggest that Elephas maximus was possibly a mixed feeder with a broader range of dietary resources than Stegodon orientalis, which probably browsed on a narrower range of plant resources in more densely forested landscape. A chronological comparison (from 8 Ma to recent) of published δ13C data for these two species from Asia showed that none of them were dietary specialists. However, Elephas had a more flexible foraging ecology and a stronger ability to exploit abrasive grasses than Stegodon. The niche partitioning and perceived different foraging behaviors of Stegodon and Elephas might have reduced the level of interspecific competition and allowed them to coexist during the Pleistocene. Moreover, the high-level of ecological flexibility of Elephas might have helped them to survive until the present day, while Stegodon eventually went extinct by the terminal Pleistocene ∼12 ka. An extensive comparison and evaluation of the δ13C data from fossil mammals in mainland southeast Asia during the Early to Late Pleistocene suggests that southern China was dominated by C3 vegetation throughout the Pleistocene, in contrast with the evidence of C4 biomes in neighboring Laos, Cambodia, and Thailand. Southern China experienced relatively stable environments during the Pleistocene, which can be attributed to the wide range of mountainous regions that acted as ecological refugia from human interference and climatic fluctuations, and allowed the preservation of high biodiversity. The isotopic data we present here provides new evidence about the ecological complexity of mainland southeast Asia and elucidates the need for more systematic research to investigate extinction models and ecological conservation in this region.

Flores represents a unique insular environment with an extensive record of Pleistocene fossil remains and stone artefacts. In the So'a Basin of central Flores these include endemic Stegodon, Komodo dragons, giant tortoises, rats, birds and hominins, and lithic artefacts that can be traced back to at least one million years ago (1 Ma). This comprehensive review presents important new data regarding the dating and faunal sequence of the So'a Basin, including the site of Mata Menge where Homo floresiensis-like fossils dating to approximately 0.7 Ma were discovered in 2014. By chemical fingerprinting key silicic tephra originating from local and distal eruptive sources we have now established basin-wide tephrostratigraphic correlations, and, together with new numerical ages, present an update of the chronostratigraphy of the So'a Basin, with major implications for the faunal sequence. These results show that a giant tortoise and the diminutive proboscidean Stegodon sondaari last occurred at the site of Tangi Talo ∼1.3 Ma, and not 0.9 Ma as previously thought. We also present new data suggesting that the disappearance of giant tortoise and S. sondaari from the sedimentary record occurred before, and/or was coincident with, the earliest hominin arrival, as evidenced by the first records of lithic artefacts occurring directly below the 1 Ma Wolo Sege Tephra. Artefacts become common in the younger layers, associated with a distinct fauna characterized by the medium-sized Stegodon florensis and giant rat Hooijeromys nusatenggara. Furthermore, we describe a newly discovered terrace fill, which extends the faunal record of Stegodon in the So'a Basin to the Late Pleistocene. Our evidence also suggests that the paleoenvironment of the So'a Basin became drier around the time of the observed faunal transition and arrival of hominins on the island, which could be related to an astronomically-forced climate response at the onset of the Mid-Pleistocene Transition (MPT; ∼1.25 Ma) leading to increased aridity and monsoonal intensity. publishedVersion

In a limestone karst cave (Sanhe Cave) in Nongguan Mountains in Chongzuo, Guangxi, China, we unearthed in the sediments 82 mammal fossils in a Gigantopithecus fauna of 1.4 mya, including fossils of Gigantopithecus blacki, Ailuropoda milanoleuca wulingshanensis, Megatapirus augustus, Rhinoceros sinensis, and Stegodon preorientalis, the ancestor species of the white-headed langur, named Chongzuo langur (Trachypithecus chongzuosis), and the ancestor species of the Sumatran striped rabbit (Nesolagus sp.). The animals were buried during the middle phase of the Early Pleistocene Epoch, which coincided with the first extreme glaciations in the Quaternary Period. The glaciations caused the sea level to drop, and the continental shelf was thus exposed. As a result, the coastal areas surrounding Java Island and Sumatra Island became land, which connected with the Malay Peninsula, Indochina Peninsula, and the coastal lowlands around Beibu Gulf in southern Guangxi. This provided land passages for Javan Langurs (Trachypithecus sp.) to disperse northward, enter the karst hill district on the south bank of Zuo Jiang River, and gradually evolve into the present-day white-headed langur (Trachypithecus leucocephalus).

Liang Bua, the type site of Homo floresiensis, is a limestone cave on the Indonesian island of Flores with sedimentary deposits currently known to range in age from about 190 thousand years (ka) ago to the present. Recent revision of the stratigraphy and chronology of this depositional sequence suggests that skeletal remains of H. floresiensis are between ∼100 and 60 ka old, while cultural evidence of this taxon occurs until ∼50 ka ago. Here we examine the compositions of the faunal communities and stone artifacts, by broad taxonomic groups and raw materials, throughout the ∼190 ka time interval preserved in the sequence. Major shifts are observed in both the faunal and stone artifact assemblages that reflect marked changes in paleoecology and hominin behavior, respectively. Our results suggest that H. floresiensis and Stegodon florensis insularis, along with giant marabou stork (Leptoptilos robustus) and vulture (Trigonoceps sp.), were likely extinct by ∼50 ka ago. Moreover, an abrupt and statistically significant shift in raw material preference due to an increased use of chert occurs ∼46 thousand calibrated radiocarbon (14C) years before present (ka cal. BP), a pattern that continues through the subsequent stratigraphic sequence. If an increased preference for chert does, in fact, characterize Homo sapiens assemblages at Liang Bua, as previous studies have suggested (e.g., Moore et al., 2009), then the shift observed here suggests that modern humans arrived on Flores by ∼46 ka cal. BP, which would be the earliest cultural evidence of modern humans in Indonesia.

Several canid fossils, which were originally discovered and described in the early 20th century, are known from Early and earliest Middle Pleistocene of Java (Indonesia). Here we revise the taxonomy and relative age of the Javanese canid fossils in light of recent developments in the taxonomy and phylogeny of Canidae, and new insights in the evolution of island mammals. Results show that Cuon was absent during the Early and earliest Middle Pleistocene while the large-sized Xenocyon (previously Megacyon) was present in the Early Pleistocene and replaced by the small-sized Xenocyon (previously Mececyon) during the earliest Middle Pleistocene. The latter is probably an anagenetic dwarf derived from the larger form of the preceding period. The change in body size of Xenocyon on Java over time is likely the effect of increased competition within the carnivore guild within the restricted boundaries of the island. Simultaneously with a pronounced body size shift, a dietary shift from large-sized prey to much smaller prey must have taken place in order to meet energetic constraints. The degree of endemism of terrestrial mammals of the earliest Middle Pleistocene horizon of the site Trinil, which has also yielded Homo erectus fossils, indicates that during this period, Java was marginally isolated which allowed for a corridor dispersal to the island with subsequent vicariance. The nature and degree of isolation may have been similar to that of Late Pleistocene Sicily, but of a longer duration, given the higher degree of dwarfism of the stegodon, antilope and canid. The following continentalisation enabled the invasion by mainland terrestrial mammals, as is seen in younger layers at Trinil, and eventually by Homo sapiens and Cuon in the Late Pleistocene / Holocene.

In southern China, the Ailuropoda-Stegodon fauna is a very general one with a relatively stable Ailuropoda and Stegodon fossil assemblage throughout the Pleistocene epoch, and cervid remains are also significant elements of this fauna. The taxonomic study on isolated cervid tooth fossils of the Late Pleistocene Ailuropoda-Stegodon fauna is not very adequate. The numerous cervid tooth fossils recently unearthed from Yangjiawan caves and Fuyan Cave give us a possibility to address this issue. Based on morphologic, odontometric and CT scanning studies, six moschid and cervid species have been recognized, including Moschus sp., Elaphodus cephalophus, Muntiacus muntjak, M. reevesi, Rusa unicolor and Cervus nippon. The cervid faunal compositions of these two sites are basically the same, and most of the species can be distinguished from one another in tooth dimensions, except for M. muntjak and E. cephalophus; but the entoflexus of upper premolars, the metacone ribs of upper molars, the structure of p4, and the metastylid of lower molars can be used as distinguishable characters between M. muntjak and E. cephalophus. The cingulum and the entostyle of upper molars, and the metastylid of lower molars of M. reevesi are weaker than those of M. muntjak. R. unicolor possesses tapered entostyle and thick enamel, while the enamel of C. nippon is relatively thinner and the entostyle is columned. Moschus sp. usually has a Dorcatherium fold in lower molars, which is a diagnostic character for Moschus species.

The Himalayan foreland basin is characterised by three principal sedimentary successions. These are the Subathu Group (Palaeocene-Eocene), the Murree or Dharamsala Group (Oligocene-Miocene) and the Siwalik Group (Miocene-Pleistocene). The basin was linked in the east to the Bay of Bengal through the Bengal Basin and its western part was connected to the Arabian Sea through Rajasthan and Kachchh, Gujarat. The Himalayan foothills have long been known for yielding rich mammalian fossil occurrences and significant localities in the Indian Siwaliks include Kalagarh, Ramnagar, Haritalyangar, Chandigarh and Jammu regions. The Bugti Basin and Zinda Pir area of Sulaiman Range of Pakistan were also part of the foreland basin, as Lower and Middle Siwalik fossils are known from the Vihowa and Litra formations, respectively. The Marwat Formation of Bhittani and Marwat Ranges, Pakistan have also yielded Pinjor-age vertebrate fossils. The Middle and Upper Siwalik-age fossils are also known from the Irrawaddy Formation of Myanmar, where older Pegu beds have also yielded scanty Lower Siwalik-age fossils. This faunal succession in the Irrawaddy Valley may have geologically and biogeographically belonged to the Himalayan foreland basin in the past. Additional mammalian fossils having an affinity with the Lower and Middle Siwaliks are known from the Khari Nadi Formation of Kachchh. The Bokabil Formation of Tripura also yielded scanty Lower to Middle Siwalik fossils. A younger Siwalik late Miocene assemblage is known from Piram Island of Gujarat. These areas are also part of the foreland basin. Most recently, a late Miocene suid species was discovered and reported from near Baripada, Odisha. The paper discusses the respective faunal lists of the various successions mentioned above. Finally, the paper discusses the implications of the foreland faunas. The faunas of Lower and Middle Siwaliks of foreland basin have links with Europe, Turkey, Africa and Central Asia and throw light on the uplift of the Himalaya. The post-Siwalik faunas of central and Peninsular India and Indo-Gangetic Plains have also yielded remnant Siwalik taxa, including Stegodon insignis, Elephas hysudricus and Potamochoerus theobaldi. The Kurnool Cave Fauna and Recent Fauna of India have also generic resemblance with the foreland faunas.

It has long been thought that environmental perturbations were the key driving force behind the succession of three distinct mammal faunas in the Pleistocene Ailuropoda-Stegodon faunal complex (sensu lato) of South China: the lower Pleistocene Gigantopithecus-Sinomastodon fauna, the middle Pleistocene Ailuropoda-Stegodon fauna (sensu stricto) and the upper Pleistocene Homo-Elephas fauna. Here, we apply three-dimensional dental microwear texture analysis (DMTA) to three characteristic fossil proboscideans from these mammal faunas to provide preliminary tests for hypotheses of trophic ecology. Despite a few methodological caveats, this study demonstrates the potential of DMTA for understanding the diets of fossil proboscideans. The texture of microwear in Sinomastodon and Stegodon are more reflective of browsing, whereas that of Elephas is suggestive of mixed feeding. The results suggest a more complex process of Pleistocene faunal turnovers in South China. Rather than a unidirectional trend of climate-driven environmental deterioration, biotic factors might have played a more substantial role than previously thought.

Handaxes, the characteristic tool of the Acheulean industrial complex, are predominantly made of stone. Handaxes made of bone are much less common. Only a few have been reported, from sites in Africa, Europe and western Asia. In this study we report a bone handaxe from Chongqing, southwest China, which represents the first bone handaxe ever discovered in China or any other part of East Asia. Typologically, it is somewhat inconsistent with the classic Acheulean handaxe morphology and is therefore classified as a proto-handaxe in this study. The bone from which the handaxe was made, which is now fossilized, has been dated to ∼170 ka based on a U-series technique. The handaxe was manufactured from the mandible of an individual of the stegodontid Stegodon orientalis , a typical member of the Middle Pleistocene Ailuropoda - Stegodon fauna ( sensu stricto ) of southern China. This artifact represents the earliest evidence for a tradition of bone handaxe manufacture in East Asia, and provides important evidence regarding the lifestyle, technology, and environmental surroundings of the humans that occupied the upstream region of the Yangtze River during the Middle Pleistocene.

As one of the most important Quaternary mammalian faunas in southern China, the Gigantopithecus-Sinomastodon fauna has received much attention. The large-primate fossil teeth newly collected from Juyuan karst cave in Boyue Mountain, Guangxi Zhuang Autonomous Region of southern China, can be identified as Gigantopithecus blacki . The morphology and size of these G. blacki teeth differ from those of the middle Pleistocene Gigantopithecus teeth which hold evidently larger dental dimensions and more complex crenulations from Hejiang Cave also in Guangxi and Tham Khuyen Cave in Vietnam. However, the G. blacki teeth from Juyuan Cave are relatively similar to those from Early Pleistocene Mohui Cave and Liucheng Gigantopithecus Cave both in Guangxi, which suggests that the three cave sites have similar age. The Juyuan fauna associated with Gigantopithecus blacki , consisting of 45 mammalian species (such as Sinomastodon yangziensis , Ailuropoda wulingshanensis , Stegodon huananensis , and Rhinoceros fusuiensis ), is a typical Early Pleistocene Gigantopithecus-Sinomastodon fauna of southern China. The Juyuan fauna is mostly similar to Mohui fauna, also implying their contemporaneity. Paleomagnetic analyses demonstrate that the fossil-bearing sediments in Juyuan Cave are dominated by normal polarity. Combining the faunal analysis and magnetostratigraphic evidence, the Juyuan sediments can be best correlated with the Olduvai normal subchron, giving an estimated age of 1.8 Ma.

Gigantopithecus blacki was once a common primate during Early Pleistocene, distributed throughout southern China, but then it gradually declined with time and finally became extinct probably during Middle Pleistocene. Black Cave in Guangxi (the Zhuang Nationality Autonomous Region) was the first site yielding G. blacki teeth in-situ in 1956. It also produced numerous mammalian fossils belonging to the typical Middle Pleistocene Stegodon–Ailuropoda fauna from sediments overlying the G. blacki level. The present study carried out coupled ESR/U-series dating study of five Bovidae teeth from this palaeontological layer and obtained a weighted mean age of 383 ± 20 ka for this Stegodon–Ailuropoda fauna. This age estimate on fossil teeth is constrained by two 230 Th/U ages of 404 ± 24 ka and 382 ± 9 ka obtained on speleothems formed below and above the level of the Stegodon–Ailuropoda fauna. The Black Cave Stegodon–Ailuropoda fauna can be therefore correlated with MIS 11, and the remains of G. blacki should be older than 400 ka.

Sulawesi is an island located in the Wallacean region of Indonesia. Geologically its lying midway between the Asian (Sunda) and Greater Australian (Sahul) continents . As a part of Wallacea islands, Sulawesi is an island that shows complexity either in biology or geology perspective. Though the distinctive quaternary vertebrate faunas has been described from Sulawesi, historical pattern of biogeography still poorly understood due to the lack of the fossil specimens. This paper describes a maxilla fragment with molar root teeth M 1 from an archaic proboscidae called Stegodon that found in the conglomeratic sandstone layer, at Cangkange Area, 4 km to the east of Cabenge Archeological site of South Sulawesi, Indonesia. Based on the comparation measuring data between this specimen with the Stegodon sompoensis and the Stegodon trigonocephalus it can be concluded that this Stegodon maxilla fragment is belong to the Stegodon sompoensis, a dwarf Stegodon from Sulawesi Island. The specimen is a surface collected sample. Based on the attached matrix on the maxilla fragment, this specimen interpreted to be derived from subunit A of Beru Member, Walanae Formation. This Stegodon sompoensis is likely to be lived near the coastal-lagoon around 2,5 million years ago or Late Pliocene to Early Pleistocene. This estimated specimen age is based on the vertebrate fauna biostratigraphy of South Sulawesi. ABSTRAK Pulau Sulawesi di Indonesia terletak di daerah Wallacea. Secara geologi pulau ini berada di antara Asia (paparan Sunda) dan Australia (paparan Sahul). Sebagai bagian dari kepulauan Wallacea, Pulau Sulawesi merupakan pulau yang memiliki kompleksitas baik dari segi biologi maupun geologinya. Meskipun fauna-fauna vertebrata kuarter Sulawesi sudah dideskripsi, tetapi sejarah dan pola biogeografi di pulau ini masih sangat kurang dikarenakan sedikitnya fosil-fosil yang ditemukan. Tulisan ini mendeskripsikan fragmen maxilla dari gajah purba jenis Stegodon dengan akar gigi molar M 1 yang ditemukan di perlapisan batupasir konglomeratan, di daerah Cangkange, sekitar 4 km ke arah timur dari situs arkeologi Cabenge, Sulawesi Selatan, Indonesia. Berdasarkan perbandingan data pengukuran spesimen ini dengan Stegodon sompoensis dan Stegodon trigonocephalus maka disimpulkan bahwa fragmen maksila Stegodon ini berasal dari Stegodon sompoensis , jenis Stegodon kerdil dari Pulau Sulawesi. Spesimen ini merupakan temuan permukaan, tetapi berdasarkan matriks sedimen yang masih menempel di maxilla, spesimen ini diinterpretasikan berasal dari Anggota Beru subunit A. Stegodon sompoensis ini diperkirakan dahulu hidup di lingkungan lagoon dekat pantai pada sekitar 2,5 juta tahun yang lalu atau Pliosen Akhir sampai Pleistosen Awal. Penentuan umur ini didasarkan pada boistratigrafi fauna vertebrata Sulawesi Selatan.

Matar is a locality near Ngandong located on the east bank of the Solo River. Archaeological fieldwork in this locality revealed the existence of several ancient alluvial terraces. Fossils and lithic artifacts found embedded on these terraces were in poorly consolidated conglomerate comprising sand and gravels. Faunal remains from Matar consisted of several extinct taxa such as Stegodon trigonocephalus, Bubalus paleokarabau, Bibos paleosondaicus, and Hexaprotodon sivalensis. These fauna represented typical dry and open-woodland environment in the Middle and Upper Pleistocene in Java. In addition, lithic artifacts were found in the same layer and consisted of flake-tools and a small number of crude core-tools. Flake tools from Matar were quite similar to the description of flake-tools found in Ngandong (see Movius Jr., 1948; Van Heekeren, 1972). However, the raw material used was not limited to chalcedony, but included chert and jasper. Our study has revealed that Matar and its surrounding environment is a promising locality, although it has been forgotten for more than two decades since G.J. Bartstra visited this locality for the first time in 1986.

International audience; The different ecologies, times of extinction and 'last stand' of both Elephas and Stegodon in Southeast Asia is covering almost two million years. For Middle Pleistocene, both taxa belong to the complex Ailuropoda-Stegodon. This regional complex is considered to have a chronological significance and further palaeoecological, palaeobiogeographical or biochronological studies are using this assemblage as a benchmark. Nevertheless, such studies do not provide sufficient information regarding site formation and duration to be consistent enough to do so at an appropriate resolution (MIS timescale). Focussing attention on the occurrence of Proboscideans, a critical review of the robustness of the geological, taphonomical and chronological data of Middle Pleistocene faunal assemblages suggests to undertake a deep reappraisal of this "biochronological benchmark". Recomandations were provided in the 1980s not to use mixtures of faunal assemblages and progress in geochronology for two decades are available. Nevertheless, reviewing Proboscideans suggests the necessity of a severe revision and leads to be very sceptical on the use of the Ailuropoda-Stegodon complex as an ecological marker. Extended to other taxa such a review casts serious doubts on former and current paleoecological modellings and studies.

Aim We investigated the hypothesis that insular body size of fossil elephants is directly related to isolation and surface area of the focal islands. Location Palaeo-islands worldwide. Methods We assembled data on the geographical characteristics (area and isolation) of islands and body size evolution of palaeo-insular species for 22 insular species of fossil elephants across 17 islands. Results Our results support the generality of the island rule in the sense that all but one of the elephants experienced dwarfism on islands. The smallest islands generally harbour the smallest elephants. We found no support for the hypothesis that body size of elephants declines with island isolation. Body size is weakly and positively correlated with island area for proboscideans as a whole, but more strongly correlated for Stegodontidae when considered separately. Average body size decrease is much higher when competitors are present. Main conclusions Body size in insular elephants is not significantly correlated with the isolation of an island. Surface area, however, is a significant predictor of body size. The correlation is positive but relatively weak; c. 23% of the variation is explained by surface area. Body size variation seems most strongly influenced by ecological interactions with competitors, possibly followed by time in isolation. Elephants exhibited far more extreme cases of dwarfism than extant insular mammals, which is consistent with the substantially more extended period of deep geological time that the selective pressures could act on these insular populations.

Understanding global patterns of late Quaternary megafaunal extinction is impeded by geographic variation in data quality and quantity. The magnitude, timing and drivers of megafaunal extinctions remain poorly understood for India, a region with a strong history of palaeontological research. We review available data for all putative extinct Indian megafaunal taxa with direct or indirect reported evidence of potential survival into the global “megafaunal extinction window” (from ∼50,000 years onwards). Assessment of late Quaternary megafaunal species richness is confused by multiple levels of taxonomic uncertainty, including the relationship of named Late Pleistocene taxa to extant taxa, and nomenclatural confusion over correct species names. There is sufficient evidence to recognise up to four genuine global megafaunal species-level extinctions in India during the Late Pleistocene: two proboscideans (Palaeoloxodon namadicus, Stegodon sp.), a hippopotamus (Hexaprotodon sp.), and possibly a horse (Equus namadicus). A fifth extinct megafaunal species, the Indian aurochs Bos [primigenius] namadicus, definitely persisted into the Holocene. Other Indian late Quaternary megafaunal species (Rhinoceros spp., Bubalus palaeindicus, Sus “palaeindicus”, Crocodylus palaeindicus) are likely to be synonyms of extant species. Reconstructing regional extinction dynamics is further impeded by chronological uncertainty; however, attempts to obtain new dates for vertebrate samples from six late Quaternary sites in five regions were unsuccessful. Accurate understanding of the dynamics of megafaunal extinctions in India will require robust taxonomic, chronological and palaeoecological data, and we encourage further investigation of the region’s rich late Quaternary record.

Over the last 50,000 years, a majority of megafauna went extinct globally. Only two regions, Africa, and tropical Asia retain megafaunal communities today. Their persistence in Africa has been attributed to co-evolution with hominins, but faunal dynamics are largely understudied in South Asia. Focusing on the Indian Subcontinent and using a novel dataset compiled from the published literature, we report an extremely size biased, but low magnitude extinction during the terminal Pleistocene and early Holocene—approximately 30,000 years after the arrival of Homo sapiens in the region. We document four mammalian extinctions—Palaeoloxodon namadicus, Stegodon namadicus, Hexaprotodon sp., and Equus namadicus; the extirpation of ostriches (Struthio camelus); and a pseudo-extinction of Indian aurochs (Bos namadicus). The per-capita extinction rate is comparable to eastern and southern Africa, but much lower than elsewhere. Given the long history of hominins in the Indian Subcontinent, and similar extinction patterns to those seen in Africa, we provide the first independent test of the co-evolution hypothesis, and further, suggest that robust population networks and climatic refugia may have allowed for the persistence of megafauna in this region.

To extend the range of materials that can be used to provide chronological constraint in archaeological and palaeontological settings, we explore the use of cave pearls that form in shallow pools. Here, we present U-series dating results of cave pearls found in cave sediments with mammal fossils collected from systematic excavations at the Diaozhongyan Cave (“DZY”), located in Guangxi Zhuang Autonomous Region, southern China. Similar to soda straw stalactites, cave pearls can be transported after formation and subsequently incorporated into sediments. Thus, the U-series ages of the outermost layers of cave pearls are expected to be older than their host sediments and associated archaeological and paleontological materials. In conjunction with U-series dating of mammal teeth, which can provide a reliable minimum age, U-series ages of cave pearls can provide a maximum age limit of the deposits. Thus, dating of the mammal teeth and cave pearls from DZY reasonably constrains the first occurrence of the DZY Ailuropoda-Stegodon fauna to a period between 205.6 ± 1.4 ka and 231.0 ± 12.8 ka. These age constraints agree well with previous estimates for the time range of the late Middle Pleistocene Ailuropoda-Stegodon fauna in southern China. We propose that greater attention should be paid to cave pearls, which are commonly found in cave deposits that were laid down at least in part due to fluvial activity, and can be used for U-series dating to better constrain the age of the associated cave deposits.

The dental evolution of Mammuthus during the Quaternary represents the ultimate level of adaptation to open habitats among all proboscideans. This is the end result of the increased trend in grazing ability in Elephantidae. The combination of mesiodistally flat molar plates and proal (from back to front) jaw movement allowed elephants to increase their grazing effectiveness to an extent that gomphotheres failed to achieve. This does not necessarily mean that this feature evolved as an adaptation to grazing. The acquisition of proal jaw movement in Elephantidae is not well delineated because of the poor fossil record of the earliest elephants. However, it seems that proal jaw movement can be developed independent of the adaptation to grazing. Stegolophodon from the Early Miocene of Japan (ca. 17.3 Ma) suggest that proal jaw movement was fully developed in stegolophodonts by 17.3 Ma. This date roughly coincides with the onset of the Miocene climatic optimum, but forested habitats remained in Southeast Asia and Japan. Thus, at that time, there was no climatic change that selected stegolophodonts to adapt to grazing. Stegodonts remained brachydont throughout their history. Stable carbon isotope data suggest that stegodonts had not become grazers before the latest Pliocene, and this may explain the differences between Elephantidae and Stegodontidae in molar height increase. On the other hand, the browsing habitat of most stegodonts fails to explain the increased number of loph(id)s in stegodonts subsequent to the latest Miocene.

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 south1, and by about 50 thousand years ago, modern humans (Homo sapiens) had crossed to Sahul2,3. On the basis of position, oceanic currents and biogeographical context, Sulawesi probably played a pivotal part in these dispersals4. 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.

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.

The Indonesian islands have long been recognized for their rich and unique avifaunas, but little is known regarding how past processes and events have shaped current avian distributions. Here we compare the modern non-passerine avifauna of Flores with the Late Pleistocene non-passerine fossil assemblage from the cave site of Liang Bua to assess whether the Late Pleistocene assemblage differs from the modern avifauna. Randomized permutation tests failed to detect a statistically significant difference in body size distributions, but a significant difference in dietary guild was found, as the modern fauna lacks scavengers. The emerging pattern of avian extinctions on Flores is characterized by a low proportion of extinct species, a loss of large-bodied species, and apparently minor effects on avian community structure. This is in contrast to other oceanic islands, which experienced dramatic changes in avifauna after the arrival of modern humans. Flores' close proximity to other islands and landmasses likely allowed for population connectivity that buffered populations from extinction. Widespread species may also have been able to recolonize if local extirpations took place. The extinction of the large-bodied avian scavengers Leptoptilos robustus and Trigonoceps sp. on Flores is consistent with the pattern of human-caused extinctions on other oceanic islands. However, the loss of these two large scavenging species may be linked to the extinction of the pygmy proboscidean ( Stegodon florensis insularis ). Such a dependence of avian species on mammalian megafauna, leading to extinction by trophic cascade, is characteristic of continental Late Pleistocene extinctions.

Late Miocene climate change particularly monsoon intensification brought about by tectonic upheavals changed significantly the regional vegetation scenario of the Indian Subcontinent. Siwalik Middle Miocene closed forests gave way to Late Miocene open and seasonal forests, which in turn were gradually replaced by latest Miocene and Early Pliocene wooded grasslands. Here, new stable isotope and microwear data of a Late Miocene rhizomyid rodent, and dental microwear analyses of selected Late Miocene, and Plio-Pleistocene large and small herbivorous mammals from India is presented. These findings are then integrated with the earlier palaeodiet and palaeohabitat data of Siwalik mammals, in order to get an overall picture of climate vis a vis dietary and habitat change between ∼14 and ∼1 Ma. The results indicate that the Late Miocene rhizomyid Rhizomyides sivalensis was primarily a C3 grazer, but might have also consumed leaves, tubers, roots, and seeds. The Miocene murine rodents Antemus chinjiensis and Karnimata cf. K. intermedia most likely ate fruits and insects. Among the Plio-Pleistocene murines Cremnomys and Mus cf. Mus flynni were mixed feeders of seeds, grass and insects. Bandicota was a seasonal grazer and mixed feeder, whereas Golunda was primarily a grazer with some intake of seeds and insects. A late Miocene rhinocerotid from Haritalyangar was a C3 grazer, whereas its Plio-Pleistocene relative was a C4 grazer. The hippo Hexaprotodon was a C4 grazer. Plio-Pleistocene giraffid Sivatherium giganteum was a C4 grazer, whereas its close relative Bramatherium was a mixed feeder with its main diet being C4 grass. Camelus was also a mixed feeder with a dominant C4 grass diet. Damalops was a mixed feeder with C4 grass being its principal diet, whereas Hemibos was a C4 grazer. Plio-Pleistocene elephantids Stegodon and Elephas were C3 browsers and C4 grazers, respectively. The Miocene arboreal frugivore/browser primates (sivapithecenes and sivaladapids) that lived in tropical forests were replaced by immigrant terrestrial grazers (baboons) in the Plio-Pleistocene, when the landscape was dominated by savannah type grasslands. On similar lines, Late Miocene monsoon intensification was responsible for the disappearance of several lineages of frugivore/browser suids, tragulids and bovids, which then were later replaced by omnivores/mixed feeders/grazer immigrants. Lineages of rhinos, equids, giraffids, anthracotheres and murids that had a C3 diet in the Late Miocene started adapting to a C4 diet. However, among the proboscideans the Miocene dinotheres (C3 browsers) were replaced by the Plio-Pleistocene elephantids, which were both browsers and grazers.