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Sundaland, comprising the low-lying Sunda Shelf, the major islands of Borneo, Sumatra, and Java, as well as many smaller surrounding islands, formed a contiguous landmass through much of the Pliocene and Pleistocene. Subsequent late-Pleistocene rising sea levels have often been invoked as major zoogeographic barriers to gene flow and associated phenotypic and taxonomic divergence of Sundaic mammals. However, previous examinations of mammalian biogeography have painted a complex picture, suggesting that mammalian diversification in the region cannot easily be explained by a single overarching biogeographical phenomenon. Instead, it may be that the significant climatic changes and their concomitant impacts on local environments were the more significant control. In this context, we study adult skull morphology of a widely distributed primate in insular and continental Southeast Asia, the long-tailed macaque (Macaca fascicularis), using a 3D geometric morphometric (GM) approach. We used principal component analysis and linear discriminant function analysis to analyze the associations between morphological divergence and island biogeography, taxonomy, and phylogeny. Reduced rank regressions (RRR) were employed to investigate associations with climate of the present and last interglacials and the last glacial maximum (LGM). There was very poor morphological divergence and discrimination among specimens from different biogeographical zones. There was also large overlap between M. fascicularis subspecies in skull size and shape, with M. f. fascicularis occupying the largest range of size and shape variation, consistent with its wide geographical distribution. Nonetheless, sundaic M. f. fascicularis and M. f. philippinensis, restricted to the Philippines, were successfully discriminated on size and shape, supporting the latter as a valid subspecies. There was moderate morphological differentiation associated with the deepest split in mitochondrial DNA lineages (M. f. aureus versus the rest) in skull size and shape. Contrary to biogeography, we detected strong associations between skull morphology and climate. Skull form (size and shape combined) was better explained by climate closer to present conditions and correlated most strongly with the current interglacial, even though past climatic conditions do show correlations with cranial morphology. We found no strong evidence that cranial and mandibular variation in M. fascicularis consistently tracks insularity caused by Quaternary sea level change. Instead, the morphological overlap evident in our sample indicates connections between subspecies and is consistent with the history of Sundaland as a single landmass for large portions of the Quaternary.

We report on results from surveys undertaken in Sumatra during 2018 and 2019. The surveys had three objectives: (1) to examine, sample, and record the extensive Quaternary fossil deposits from caves in West Sumatra; (2) determine the potential for fluvial deposits in Riau and Jambi provinces; and (3) relocate the fossil proboscidean remains reported from Bangka Island. Our surveys produced several significant results. We mapped three important Padang Highland caves, Ngalau Lida Ajer, Ngalau Sampit, and Ngalau Gupin, locating and sampling the main fossil deposits in each, as well as recording additional caves in the region. Our surveys of the fluvial systems in central-west Sumatra did not reveal any vertebrate Pleistocene deposits but did yield Mio-Pliocene trace fossils. Finally, we relocated elephant fossils from Bangka, but no in situ vertebrate remains could be found. These finds add important new data to the geological history of Sumatra.

Orangutan (Pongo spp.) fossils have been found throughout much of Pleistocene Southeast Asia. Today, Pongo is restricted to three living species on the islands of Borneo and Sumatra. They are predominately frugivores, and during fruit masting events their diets can consist entirely of fruit. However, food procurement strategies between masting events differ between islands and thus species. Sumatran orangutans (Pongo abelii) are largely able to fall back on non-masting fruit species, such as figs, such that they can maintain higher quality diets compared to their Bornean counterparts (Pongo pygmaeus), who ingest more lower quality and tougher foods between masting events. However, diets of fossil orangutans remain largely unknown. We investigated the diets of Sumatran orangutans from the Pleistocene deposits of the Padang Highlands, western Sumatra, using dental microwear texture analysis. We tested whether the diets of the fossil species were similar to those of Bornean orangutans, all great apes generally, and macaques that occupy similar habitats to modern orangutans, in order to gain insights into palaeoenvironments present in western Sumatra. We found that fossil orangutans consumed foods less tough than modern Bornean Pongo pygmaeus, and thus presumably more fleshy fruits (absent of significant seed mastication). We found no differences in the amount of hard object feeding between modern and fossil orangutans. Taken together, we suggest that Pleistocene orangutans from western Sumatra had similar diets to Pongo abelii, indicating that masting events were not a significant contributing factor in their intake of fruit. Finally, we suggest anisotropy values in orangutan dental microwear may be an important marker for fruit masting events in the Pleistocene.

A spate of recent discoveries has highlighted the complex and multidimensional nature of movements of hominins, and in particular Homo sapiens, from Africa into Asia and beyond (e.g. Clarkson et al., 2017; Aubert et al., 2018; Groucutt et al., 2018; Ni et al., 2021). The long-accepted idea of a single, late wave of migration of modern humans from Africa to Australia is being replaced by an understanding that there were several dispersal events, of varying levels of penetration and success, contributing to the full global expansion of humans (Rabett, 2018; Lee and Hudock, 2021). The Asia-Pacific region is key to understanding the timing, nature, and environmental and cultural context of the first and last of these migrations (e.g. Stewart et al., 2020; Zhang et al., 2018; Louys and Roberts, 2020; Roberts et al., 2020). While hominins moving out of Africa first encountered Asia, the Pacific, at the other end of the world, records the very last step of these movements. By examining the archaeological, palaeontological, and environmental records of these regions we can begin to develop an understanding of how both local and global factors have influenced the course and trajectory of human evolution outside of Africa.

We employ high-throughput thermal-neutron tomographic imaging to visualise internal diagnostic features of dense fossiliferous breccia from three Pleistocene cave localities in Sumatra, Indonesia. We demonstrate that these seemingly homogeneous breccias are an excellent source of data to aid in determining taphonomic and depositional histories of complex depositional sites such as tropical caves. X-ray Computed Tomographic (CT) imaging is gaining importance amongst palaeontologists as a non-destructive approach to studying fossil remains. Traditional methods of fossil preparation risk damage to the specimen and may destroy contextual evidence in the surrounding matrix. CT imaging can reveal the internal composition and structure of fossils contained within consolidated sediment/rock matrices prior to any destructive mechanical or chemical preparation. Neutron computed tomography (NCT) provides an alternative contrast to X-rays, and in some circumstances, is capable of discerning denser matrices impenetrable to or yielding no contrast with CT imaging. High-throughput neutron imaging reduces neutron fluence during scanning which means there is less residual neutron-induced radioactivation in geological samples; allowing for earlier subsequent analyses. However, this approach remains unutilised in palaeontology, archaeology or geological surveys. Results suggest that the primary agents in the formation of the breccias and concentration of incorporated vertebrate remains are several rapid depositional phases of water and sediment gravity flow. This study highlights the potential for future analyses of breccia deposits in palaeontological studies in caves around the world.

Though hundreds of caves are known across Mongolia, few have been subject to systematic, interdisciplinary archaeological surveys and excavations to understand Late Pleistocene and Holocene environments. Previous cave excavations in Mongolia have demonstrated their potential for preservation of archaeological and biological material, including Palaeolithic assemblages and Holocene archaeology, particularly burials, with associated organic finds. In other cases, cave surveys found that stratigraphic deposits and archaeological materials are absent. The large number of caves makes the Mongolian Altai Mountain Range a potentially attractive region for human occupation in the Pleistocene and Holocene. Here we present the results of an interdisciplinary survey of caves in four carbonate areas across the Gobi-Altai Mountains. We report 24 new caves, some of which contain archaeological material recovered through survey and test excavations. Most caves presented limited sedimentation, and some were likely too small for human habitation. Six caves showed evidence of palaeontological remains, most likely from the Late Holocene and recent periods. The most notable anthropogenic findings included petroglyphs at Gazar Agui 1 & 13. Gazar Agui 1 also contained lithics and a bronze fragment. Tsakhiryn Agui 1 contained 31 wooden fragments that include an unused fire drilling tool kit and items commonly found in association with medieval burials. We observed that the caves remain in contemporary use for religious and economic purposes, such as the construction of shrines, mining and animal corralling. Water samples from the caves, and nearby rivers, lakes, and springs were analysed for their isotopic compositions (δ18O, δD, δ17O, 17Oexcess, d-excess) and the data, combined with backward trajectory modelling revealed that the Gobi-Altai region receives moisture mainly from western sources. These results form a baseline for future archaeological, palaeoclimate and palaeoecological studies about regional seasonality and land use.

Human ribs are thought to be less affected by mechanical strain at the microscopic level than limb bones, implying that rib remodelling better reflects bone physiological homeostasis. Here, we test the hypothesis that rib tissue will be well vascularized and thus enhance susceptibility to metabolic influence. An intra-skeletal comparison of bone vascular canal density was conducted using a macropod animal model adapted to bipedal habitual hopping. The right humerus, ulna, radius, femur, tibia, fibula, a mid-thoracic and upper-thoracic rib of an eastern grey kangaroo (Macropus giganteus) were sectioned at the midshaft, from which histological sections were prepared. Bone vascularity from a maximum of 12 mm2 of sub-periosteal parallel-fibred and lamellar bone was recorded, resulting in a total of 2047 counted vessels. Vascular canal density data were corrected by cortical width, maximum length, and midshaft circumference robusticity indices computed for each bone. The fibula consistently had the highest vascular canal density, even when corrected for maximum length, cortical width and midshaft circumference robusticities. This was followed by the mid- and upper-thoracic ribs. Vascularity differences between bones were relatively consistent whether vascular canal density was controlled for by cortical width or midshaft circumference robusticities. Vascular canal density and robusticity indices were also positively and negatively correlated (p

Since the mid-1840s a diverse fossil vertebrate assemblage, referred to as the Chinchilla Local Fauna, has been collected from the Pliocene deposits of the Chinchilla Sand on the western Darling Downs of South-East Queensland. In large part because of this long history and the numerous collectors who have worked fossil deposits in the area, much ambiguity regarding site and locality names and their specific coordinates exists. Here, we review the vertebrate fossil collection records in the Queensland Museum Fossil, Donor, Collector and Locality Registers, correspondence, and field notes in an effort to pinpoint the location of each named locality and site and develop a digital map which highlights the historical collecting sites at one significant locality in the Chinchilla area. To ensure that a systematic framework for all future collecting from the main collecting area (Chinchilla Rifle Range) is maintained, we recommend the use of consistent nomenclature for sites so that spatial information of the highest possible quality is captured into the future. We recommend future collections include detailed recordings of stratigraphic contexts as well as GPS coordinates.

The continent of Australia is currently warming approximately thirty-five percent faster than the rest of the globe, with the warmest year on record coinciding with the most extreme bush fires. While effects of ongoing climate change are apparent over the past century, Australia has experienced pronounced aridification since the Miocene/Pliocene, transitioning from tropical forests to more open habitats. To better contextualize on-going climate change, we assess the ecology and paleobiology of mammalian faunas in Australia from the Pliocene to the Present in the Darling Downs region of Queensland Australia. Via the analysis of stable isotopes from tooth enamel and dental microwear texture analysis of the chewing surfaces of teeth, we clarify the ecology and paleobiology of medium to large marsupials from the Pliocene Chinchilla Sands and Pleistocene Eastern Darling Downs faunas. By comparing these ancient marsupial mammal communities to extant marsupial mammals that inhabit these regions today, we further demonstrate that the most dramatic changes between past ecosystems are clearly between those of the Present and the Plio-Pleistocene—indicated that the Darling Downs region of today is disparate as compared to the past. Most notably, Macropus giganteus consumes vegetation that is ~5.6‰ higher in δ13C values today than during the past, indicating feeding in a significantly more open landscape. The Pliocene and Pleistocene of the Darling Downs are instead dominated by mixed-feeding and browsing taxa, with several taxa exhibiting diets disparate from modern analogues (e.g., an abundance of C4 browsers). Collectively, these deep-time temporal comparisons are a clear example of how ecological communities observed today do not represent the full range of ecological niches occupied in the past and highlight the dramatic climate-departures experienced today.

The Nusa Tenggara island chain consists of an archipelago that runs roughly east-west in eastern Indonesia. As part of Wallacea, it has never been connected to any continental landmass, and has been subject to a variety of biological invasions that have populated the islands. Here, we examine the craniometric and molecular genetic records of several species of Rattus sensu lato in the island chain. We use the predictions of expanding population edge phenotypic selection in an effort to understand the movement of Rattus rattus and Rattus exulans through the archipelago. We also examine the mitochondrial haplotype networks of R. argentiventer, R. exulans, and the R. rattus Complex (RrC) and microsatellite allele frequency clustering patterns for the RrC, to examine relationships within and between Nusa Tenggara populations, and those of Asia and the Pacific where relevant for each taxon. In the RrC LIV and RrC LII haplotype networks, 20 haplotypes with seven from Nusa Tenggara were observed for RrC LIV, and 100 haplotypes with seven from Nusa Tenggara observed for RrC LII. The top performing RrC craniometric model had a negative association between size and distance from the easternmost point of the samples from Nusa Tenggara, consistent with increasing size moving west to east. The cytochrome b network for the R. exulans sequences comprised 14 haplotypes, with three observed from mainland Southeast Asia, one shared with Nusa Tenggara and regions further east, and another haplotype observed in Nusa Tenggara and in the Pacific. The R. exulans craniometric model selection produced four equally well performing models, with no migration scenario preferred. Finally, the haplotype network of R. argentiventer comprised 10 haplotypes, with six observed in Nusa Tenggara, including a relatively early cluster from the east of the archipelago. Our results are compatible with a polyphasic and polydirectional invasion of Nusa Tenggara by Rattus, likely beginning with RrC from the west to the east, an expansion of R. exulans from Flores, seemingly in no preferred overall direction, and finally the invasion of R. argentiventer from the east to the west. We find some support for the Dong Son drum maritime exchange network contributing to the distribution of the latter species.

The archaeological record of Wallacea remains exceptionally fragmentary. This is especially the case for late Holocene human occupation of the region when lifestyle and culture in marginal island environments is relatively unknown. Here we report on the archaeology of Jareng Bori rockshelter, a Metal-Age site spanning c. 1800 cal. BP up to the late historic period and situated on the eastern coast of Pantar Island in the Lesser Sunda Islands of eastern Indonesia. We use osteoarchaeological (human and vertebrate remains), invertebrate zooarchaeological (crustacean and molluscan remains), technological (lithics, shell, and pottery) and chemical sourcing (obsidian and metal) datasets to discuss networking, migration, and human subsistence strategies during this recent period of history. While some communities were no doubt living in open village settlements where they were producing pottery, the data indicate that aspects of maritime life-ways continued much as in earlier Pleistocene settlements, with people using rockshelters like Jareng Bori to pursue a range of subsistence activities focused on the shoreline. Shellfishing of rocky and reef intertidal species and fishing for mostly small herbivorous and omnivorous fishes was practised, while domestic animals only appear in the late historic period. Wider regional cultural interactions and networking are epitomized by obsidian exchange, dental modification practices, and pottery decorations, while lithic analyses indicates continuity of stone tool technology up until recent times.

Some of the earliest evidence for the presence of modern humans in rainforests has come from the fossil deposits of Lida Ajer in Sumatra. Two human teeth from this cave were estimated to be 73–63 thousand years old, which is significantly older than some estimates of modern human migration out of Africa based on genetic data. The deposits were interpreted as being associated with a rainforest environment based largely on the presence of abundant orangutan fossils. As well as the main fossil-bearing chamber, fossil-bearing passages are present below a sinkhole, although the relationship between the different fossil deposits has only been tenuously established. Here, we provide significant new sedimentological, geochronological and palaeoecological data aimed at reconstructing the speleological and environmental history of the cave and the clastic and fossil deposits therein. Our data suggest that the Lida Ajer fossils were deposited during Marine Isotope Stage 4, with fossils from the lower passages older than the main fossil chamber. Our use of stable carbon and oxygen isotope analyses of mammalian tooth enamel demonstrates that early humans probably occupied a closed-canopy forest very similar to those present in the region today, although the fossil orangutans may have occupied a slightly different niche. 1. Background 2. Material and methods (a) Speleology and sedimentary infill of the cave (b) Geochronology (c) Stable isotope analysis 3. Results (a) Speleology and sedimentary infill of the cave (b) Geochronology (i) Age and growth rate of the stalagmite (ii) Solution uranium-series analyses of dental tissues (iii) Combined uranium-series and electron spin resonance dating of fossil tooth enamel (iv) Luminescence dating (c) Stable isotope analysis 4. Discussion (a) Speleological history (b) Palaeoenvironmental history 5. Conclusion

Dietary variation within species has important ecological and evolutionary implications. While theoreticians have debated the consequences of trait variance (including dietary specialization), empirical studies have yet to examine intraspecific dietary variability across the globe and through time. Here, we use new and published serial sampled δ13Cenamelvalues of herbivorous mammals from the Miocene to the present (318 individuals summarized, 4134 samples) to examine how dietary strategy (i.e. browser, mixed-feeder, grazer) affects individual isotopic variation. We find that almost all herbivores, regardless of dietary strategy, are composed of individual specialists. For example,Cormohipparion emsliei(Equidae) from the Pliocene of Florida (approx. 5 Ma) exhibits a δ13Cenamelrange of 13.4‰, but all individuals sampled have δ13Cenamelranges of less than or equal to 2‰ (mean = 1.1‰). Most notably, this pattern holds globally and through time, with almost all herbivorous mammal individuals exhibiting narrow δ13Cenamelranges (less than or equal to 3‰), demonstrating that individuals are specialized and less representative of their overall species' dietary breadth. Individual specialization probably reduces intraspecific competition, increases carrying capacities, and may have stabilizing effects on species and communities over time. Individual specialization among species with both narrow and broad dietary niches is common over space and time—a phenomenon not previously well recognized or documented empirically.

Analyses of Pleistocene fossil proboscideans have long been used as indirect evidence for climactic and environmental shifts in the Sunda shelf of Southeast Asia. Reconstructing the biological effects of rainforest expansion at the Last Glacial Maximum on elephants can be enhanced by a better understanding of fossil proboscidean palaeobiology. We studied fragmented post-cranial fossil remains of an Asian elephant (Elephas maximus) from Pulau Bangka, an island to the east of Sumatra, hypothesised to be within the Late Pleistocene Sundaland savannah corridor. Bone histology of the humerus, rib, and vertebrae from the Bangka fossil were examined and compared with modern conspecifics to reconstruct remodelling. We measured secondary osteon population density, osteon area, diameter, and infill ratios in Haversian bone. Intra-skeletally, we found that the histology of the largely weight-bearing humerus indicated slower remodelling than that of the ribs and vertebrae, which are less biomechanically constrained. Inter-skeletally, the fossil rib histology showed relatively smaller osteons and Haversian canals when compared to the modern samples. Differences in lifestyles, including range-expansion, may have influenced micro-morphometric differences in elephant rib histology. Our results contribute indirect evidence of the effects of climactic variability in the Sunda palaeoenvironment on Pleistocene fauna.

We describe the partial cranium and skeleton of a new diprotodontian marsupial from the late Oligocene (~26–25 Ma) Namba Formation of South Australia. This is one of the oldest Australian marsupial fossils known from an associated skeleton and it reveals previously unsuspected morphological diversity within Vombatiformes, the clade that includes wombats (Vombatidae), koalas (Phascolarctidae) and several extinct families. Several aspects of the skull and teeth of the new taxon, which we refer to a new family, are intermediate between members of the fossil family Wynyardiidae and wombats. Its postcranial skeleton exhibits features associated with scratch-digging, but it is unlikely to have been a true burrower. Body mass estimates based on postcranial dimensions range between 143 and 171 kg, suggesting that it was ~5 times larger than living wombats. Phylogenetic analysis based on 79 craniodental and 20 postcranial characters places the new taxon as sister to vombatids, with which it forms the superfamily Vombatoidea as defined here. It suggests that the highly derived vombatids evolved from wynyardiid-like ancestors, and that scratch-digging adaptations evolved in vombatoids prior to the appearance of the ever-growing (hypselodont) molars that are a characteristic feature of all post-Miocene vombatids. Ancestral state reconstructions on our preferred phylogeny suggest that bunolophodont molars are plesiomorphic for vombatiforms, with full lophodonty (characteristic of diprotodontoids) evolving from a selenodont morphology that was retained by phascolarctids and ilariids, and wynyardiids and vombatoids retaining an intermediate selenolophodont condition. There appear to have been at least six independent acquisitions of very large (>100 kg) body size within Vombatiformes, several having already occurred by the late Oligocene.

Explanations for the Upper Pleistocene extinction of megafauna from Sahul (Australia and New Guinea) remain unresolved. Extinction hypotheses have advanced climate or human-driven scenarios, in spite of over three quarters of Sahul lacking reliable biogeographic or chronologic data. Here we present new megafauna from north-eastern Australia that suffered extinction sometime after 40,100 (±1700) years ago. Megafauna fossils preserved alongside leaves, seeds, pollen and insects, indicate a sclerophyllous forest with heathy understorey that was home to aquatic and terrestrial carnivorous reptiles and megaherbivores, including the world’s largest kangaroo. Megafauna species diversity is greater compared to southern sites of similar age, which is contrary to expectations if extinctions followed proposed migration routes for people across Sahul. Our results do not support rapid or synchronous human-mediated continental-wide extinction, or the proposed timing of peak extinction events. Instead, megafauna extinctions coincide with regionally staggered spatio-temporal deterioration in hydroclimate coupled with sustained environmental change., The causes of the Upper Pleistocene megafauna extinction in Australia and New Guinea are debated, but fossil data are lacking for much of this region. Here, Hocknull and colleagues report a new, diverse megafauna assemblage from north-eastern Australia that persisted until ~40,000 years ago.

Over the past decade, a growing interest has developed on the archaeology, palaeontology, and palaeoenvironments of the Arabian Peninsula. It is now clear that hominins repeatedly dispersed into Arabia, notably during pluvial interglacial periods when much of the peninsula was characterised by a semiarid grassland environment. During the intervening glacial phases, however, grasslands were replaced with arid and hyperarid deserts. These millennial-scale climatic fluctuations have subjected bones and fossils to a dramatic suite of environmental conditions, affecting their fossilisation and preservation. Yet, as relatively few palaeontological assemblages have been reported from the Pleistocene of Arabia, our understanding of the preservational pathways that skeletal elements can take in these types of environments is lacking. Here, we report the first widespread taxonomic and taphonomic assessment of Arabian fossil deposits. Novel fossil fauna are described and overall the fauna are consistent with a well-watered semiarid grassland environment. Likewise, the taphonomic results suggest that bones were deposited under more humid conditions than present in the region today. However, fossils often exhibit significant attrition, obscuring and fragmenting most finds. These are likely tied to wind abrasion, insolation, and salt weathering following fossilisation and exhumation, processes particularly prevalent in desert environments. Introduction Previous investigations Methods - Surveys - Systematic palaeontology - Taphonomic analysis Results - Systematic Palaeontology - Taphonomic analysis Discussion - Species representation and palaeoenvironmental implications - Taphonomic analysis Conclusions

Skeletal growth rates reconstructed from bone histology in extinct insular hippopotamids, elephants, bovids and sauropods have been used to infer dwarfism as a response to island conditions. Limited published records of osteocyte lacunae densities (Ot.Dn), a proxy for living osteocyte proliferation, have suggested a slower rate of bone metabolism in giant mammals. Here, we test whether insularity might have affected bone metabolism in a series of small to giant murine rodents from Timor. Ten adult femora were selected from a fossil assemblage dated to the Late Quaternary (~5000–18 000 years old). Femur morphometric data were used in computing phylogenetically informed body mass regressions, although the phylogenetic signal was very low (Pagel’s λ = 0.03). Estimates of body weight calculated from these femora ranged from 75 to 1188 g. Osteocyte lacunae densities from histological sections of the midshaft femur were evaluated against bone size and estimated body weight. Statistically significant (P

Ngalau Gupin is a broad karstic cave system in the Padang Highlands of western Sumatra, Indonesia. Abundant fossils, consisting of mostly isolated teeth from small-to large-sized animals, were recovered from breccias cemented on the cave walls and unconsolidated sediments on the cave floor. Two loci on the walls and floors of Ngalau Gupin, named NG-A and NG-B respectively, are studied. We determine that NG-B most likely formed as a result of the erosion and redeposition of material from NG-A. The collection reveals a rich, diverse Pleistocene faunal assemblage (Proboscidea, Primates, Rodentia, Artiodactyla, Perissodactyla, Carnivora) largely analogous to extant fauna in the modern rainforests of Sumatra. The hippopotamid Hexaprotodon represents the only globally extinct taxon in deposits from Sumatra and the first record of this animal from the island. This deposit is dated using combined U-series/ESR dating analyses of several teeth that yield a finite age of between ~160 and ~115 ka, depending on the modalities of the dose rate evaluation. These ages constrain the fossil assemblage, not the breccia deposits. Since the direct U-series estimate obtained on the Hexaprotodon specimen is a minimum age, it is therefore not incompatible with the U-series/ESR results. Thus, the true age of the Hexaprotodon can possibly be between 160 and 115 ka, like the other fossil remains at Ngalau Gupin. These results suggest that the faunal assemblage at Ngalau Gupin correlates with late MIS 6 or early MIS 5. Ngalau Gupin likely reflects the formation of a fossil assemblage with two primary taphonomic pathways: a prime-aged dominated macrofauna component initially produced by carnivores but subsequently accumulated by porcupines and transported to the cave, and a microfauna component likely accumulated by small carnivores. Decalcification of the cemented deposit has further resulted in loss of fossil and other sedimentary material. This site adds important new chronologically constrained fossil mammal data for the Pleistocene record of Sumatra, an island relatively poorly investigated for Southeast Asia.

The study of faunal remains from archaeological sites is often complicated by the presence of large numbers of highly fragmented, morphologically unidentifiable bones. In Australia, this is the combined result of harsh preservation conditions and frequent scavenging by marsupial carnivores. The collagen fingerprinting method known as zooarchaeology by mass spectrometry (ZooMS) offers a means to address these challenges and improve identification rates of fragmented bones. Here, we present novel ZooMS peptide markers for 24 extant marsupial and monotreme species that allow for genus-level distinctions between these species. We demonstrate the utility of these new peptide markers by using them to taxonomically identify bone fragments from a nineteenth-century colonial-era pearlshell fishery at Bandicoot Bay, Barrow Island. The suite of peptide biomarkers presented in this study, which focus on a range of ecologically and culturally important species, have the potential to significantly amplify the zooarchaeological and paleontological record of Australia. 1. Introduction 2. Material and methods 2.1. Materials 2.1.1. Modern reference specimens 2.1.2. Archaeological specimens 2.2. Collagen extraction 2.3. Matrix-assisted laser desorption/ionization–tandem time of flight mass spectrometry 2.4. Liquid chromatography with tandem mass spectrometry 2.5. Identification and confirmation of biomarkers 3. Results 3.1. Novel ZooMS peptide markers 3.2. Marsupial versus monotreme ZooMS markers 3.3. Marsupial ZooMS markers 3.4. Using ZooMS to identify macropods 3.5. Collagen fingerprinting of archaeological specimens 4. Discussion 4.1. ZooMS insights at Bandicoot Bay 4.2. Comparison to published markers 4.3. Challenges and future prospects

Despite its harsh and arid conditions, the Gobi Desert has played an important role in shaping Holocene populations, including the transition from hunting to herding lifeways. Here we present three newly documented rock art sites in the Gobi-Altai Mountains of south-central Mongolia, a cave (Gazar Agui 1), a rock shelter (Gazar Agui 13) and an open-air landscape site overlooking a palaeolake (Unegt Uul). In addition, we re-examine the preservation of the rock art cave site of Saalit Agui some 20 years after its original documentation, using digital technology not available at that time. Comparisons of rock art at Gazar Agui 1 and Saalit Agui with previously documented rock art in Mongolia suggest links with Mesolithic and Neolithic anthropomorphic iconography. Unegt Uul and Gazar Agui 13 show Early Bronze Age to Iron Age symbols, suggesting two distinct periods of production, by hunter-gatherers during the Early Holocene and by later hunter-pastoralists during the Late Holocene. Our findings suggest that wet periods in mountainous basins of the Gobi-Altai were likely key to early human habitation, with pastoralism dominating during arid periods. Our observations further indicate that preservation of rock art sites in the region is currently under threat due to human activity and climate change.

Significance We provide global assessment of the possible link between Pleistocene hominin arrival and island extinction. The existing records on islands around the world do not support a significant and detrimental impact on island biotas following island colonization prior to the Holocene. This suggests that models using island extinctions as evidence in support of anthropogenic megafaunal overhunting, or as extensions of continental-level extinctions, need to be reconsidered., The arrival of modern humans into previously unoccupied island ecosystems is closely linked to widespread extinction, and a key reason cited for Pleistocene megafauna extinction is anthropogenic overhunting. A common assumption based on late Holocene records is that humans always negatively impact insular biotas, which requires an extrapolation of recent human behavior and technology into the archaeological past. Hominins have been on islands since at least the early Pleistocene and Homo sapiens for at least 50 thousand y (ka). Over such lengthy intervals it is scarcely surprising that significant evolutionary, behavioral, and cultural changes occurred. However, the deep-time link between human arrival and island extinctions has never been explored globally. Here, we examine archaeological and paleontological records of all Pleistocene islands with a documented hominin presence to examine whether humans have always been destructive agents. We show that extinctions at a global level cannot be associated with Pleistocene hominin arrival based on current data and are difficult to disentangle from records of environmental change. It is not until the Holocene that large-scale changes in technology, dispersal, demography, and human behavior visibly affect island ecosystems. The extinction acceleration we are currently experiencing is thus not inherent but rather part of a more recent cultural complex.

In recent years, the Arabian Peninsula has emerged as a key region for elucidating hominin and faunal evolution and dispersals between Africa and Eurasia. Central to this research is the middle Pleistocene site of Ti's al Ghadah (TAG) which has yielded a diverse and abundant fossil faunal assemblage and the earliest chronometrically dated evidence for hominins in this part of the world. Here, we present the first detailed taphonomic study of the large Unit 5 fossil assemblage from the site. We aim to assess which actor/s were responsible for the accumulation of the assemblage and evaluate evidence that might be consistent with the accumulation of fauna by hominins. We also describe, for the first time, fossils and lithic artefacts from stratigraphic horizons not previously considered, providing taphonomic insights into their accumulation. The taphonomic work shows that the Unit 5 faunal assemblage was accumulated by ambush predators, likely large felids and hominins, in a lake side environment, and that carcasses were subsequently scavenged by more durophagus carnivores such as hyenas and canids. Less can be reliably said regarding the newly described fossil assemblages given their poor preservation and significant wind abrasion, but large carnivores again appear to have played a role, and hominins probably played a role in the accumulation of at least one of these. This study provides the first detail insights into the interplay between hominins, carnivores, and herbivores in Arabia, and suggests that watering holes have been a focus on the Arabian landscape for resources since the middle Pleistocene.

Rat cortical bone does not typically undergo secondary (Haversian) remodeling. Haversian organization of rat bone has been mainly observed in experimental settings following biomechanical or dietary manipulation. Here, we report an observation of cortical secondary osteons within a histological femur cross-section from an extinct (late Quaternary) form of Timorese giant rat (Murinae gen. et sp. indet). The medio-lateral midshaft diameter of its femur, used as a measure of bone size, is 6.15 mm and indicates a heavier than normal skeletal frame. We compare this sample to bone histology in a small rat's midshaft femur of 2.33 mm diameter. A complete lack of Haversian bone remodeling characteristics is noted for the smaller sample, which is dominated by radial vascular canals. The giant rat shows clear secondary osteons and diffuse vascularity mainly composed of tightly packed longitudinal canals across its cortex. It appears that rat cortical bone can undergo bone remodeling, and is organized in a highly vascularized manner, in insular giant cases. Our findings from Timor align with results reported in experimental rat model skeletal biology literature and other insular fossil rat material. Where macroanatomical examination is limited, histological observations on fossil rat limb bones have the potential to aid reconstructions of life history and skeletal growth aspects in these rodents. Anat Rec, 302:1934-1940, 2019. © 2019 American Association for Anatomy.

Timor has yielded the earliest evidence for modern humans in Wallacea, but despite its long history of modern human occupation, there is little evidence for human-induced Late Pleistocene extinctions. Here, we report on Late Pleistocene and Holocene bird remains from Jerimalai and Matja Kuru 1, sites that have yielded extensive archaeological sequences dating back to >40 ka. Avian remains are present throughout the sequence, and quails (Phasianidae), buttonquails (Turnicidae) and pigeons (Columbidae) are the most abundant groups. Taphonomic analyses suggest that the majority of bird remains, with the exception of large-bodied pigeons, were accumulated by avian predators, likely the Barn owl Tyto sp. All species represent extant taxa that are still present on Timor today, with the exception of a crane, Grus sp., from the Late Pleistocene of Jerimalai, and a large buttonquail, Turnix sp., from Matja Kuru 1. The crane likely represents an extirpated population of cranes, which were much more widespread throughout the Indonesian archipelago during the Quaternary. The large buttonquail is present at Matja Kuru 1 alongside the extant T. maculosus until at least 1372–1300 cal BP. These two species represent the first records of avian extinctions on Timor. However, a causal relationship between the extinction of these two taxa and human impact cannot be demonstrated at this point. acceptedVersion

The study of faunal remains from Kalehkoob, an archeological site in southern Khorasan, Sarayan, represents the first investigation of prehistoric vertebrates in the northern part of the Lut desert, in the east Iranian plateau. A vertebrate assemblage containing >200 specimens was found at various depths (155 cm to 595 cm) of an excavation trench. A combination of morphological and morphometric methods led to the identification of Artiodactyla, namely Bovidae (Ovis, Gazella, and Bos) and Perissodactyla, namely Equidae (Equus asinus and Equus caballus). Combined zooarcheological and sedimentological analyses points to three phases and two sub phases of depositional. Transition between the second phase and the first and third phases suggests that changes in faunal composition were potentially a response to different environmental conditions during each phase, such as a reduction of river water volume and humidity. Radiocarbon dating of bones from two depths, 155 cm (scapula) and 462 cm (horse tooth) from phases 1 and 2, respectively, provides ages of 3227–3060 cal BP and 5914–5749 cal BP for the deposit. Given the high chemical quality of both samples of scapula and tooth, δ13C and δ15N analyses were also conducted, suggesting the presence of arid environmental conditions.