Multiple hominin dispersals into Southwest Asia over the past 400,000 years.

Pleistocene hominin dispersals out of, and back into, Africa necessarily involved traversing the diverse and often challenging environments of Southwest Asia1-4. Archaeological and palaeontological records from the Levantine woodland zone document major biological and cultural shifts, such as alternating occupations by Homo sapiens and Neanderthals. However, Late Quaternary cultural, biological and environmental records from the vast arid zone that constitutes most of Southwest Asia remain scarce, limiting regional-scale insights into changes in hominin demography and behaviour1,2,5. Here we report a series of dated palaeolake sequences, associated with stone tool assemblages and vertebrate fossils, from the Khall Amayshan 4 and Jubbah basins in the Nefud Desert. These findings, including the oldest dated hominin occupations in Arabia, reveal at least five hominin expansions into the Arabian interior, coinciding with brief 'green' windows of reduced aridity approximately 400, 300, 200, 130-75 and 55 thousand years ago. Each occupation phase is characterized by a distinct form of material culture, indicating colonization by diverse hominin groups, and a lack of long-term Southwest Asian population continuity. Within a general pattern of African and Eurasian hominin groups being separated by Pleistocene Saharo-Arabian aridity, our findings reveal the tempo and character of climatically modulated windows for dispersal and admixture.

Environmental drivers of megafauna and hominin extinction in Southeast Asia.

Southeast Asia has emerged as an important region for understanding hominin and mammalian migrations and extinctions. High-profile discoveries have shown that Southeast Asia has been home to at least five members of the genus Homo1-3. Considerable turnover in Pleistocene megafauna has previously been linked with these hominins or with climate change4, although the region is often left out of discussions of megafauna extinctions. In the traditional hominin evolutionary core of Africa, attempts to establish the environmental context of hominin evolution and its association with faunal changes have long been informed by stable isotope methodologies5,6. However, such studies have largely been neglected in Southeast Asia. Here we present a large-scale dataset of stable isotope data for Southeast Asian mammals that spans the Quaternary period. Our results demonstrate that the forests of the Early Pleistocene had given way to savannahs by the Middle Pleistocene, which led to the spread of grazers and extinction of browsers-although geochronological limitations mean that not all samples can be resolved to glacial or interglacial periods. Savannahs retreated by the Late Pleistocene and had completely disappeared by the Holocene epoch, when they were replaced by highly stratified closed-canopy rainforest. This resulted in the ascendency of rainforest-adapted species as well as Homo sapiens-which has a unique adaptive plasticity among hominins-at the expense of savannah and woodland specialists, including Homo erectus. At present, megafauna are restricted to rainforests and are severely threatened by anthropogenic deforestation.

No evidence for widespread island extinctions after Pleistocene hominin arrival.

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.

Global long-term stability of individual dietary specialization in herbivorous mammals.

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 δ13Cenamel values 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 δ13Cenamel range of 13.4‰, but all individuals sampled have δ13Cenamel ranges 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 δ13Cenamel ranges (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.

Micro Methods for Megafauna: Novel Approaches to Late Quaternary Extinctions and Their Contributions to Faunal Conservation in the Anthropocene.

Drivers of Late Quaternary megafaunal extinctions are relevant to modern conservation policy in a world of growing human population density, climate change, and faunal decline. Traditional debates tend toward global solutions, blaming either dramatic climate change or dispersals of Homo sapiens to new regions. Inherent limitations to archaeological and paleontological data sets often require reliance on scant, poorly resolved lines of evidence. However, recent developments in scientific technologies allow for more local, context-specific approaches. In the present article, we highlight how developments in five such methodologies (radiocarbon approaches, stable isotope analysis, ancient DNA, ancient proteomics, microscopy) have helped drive detailed analysis of specific megafaunal species, their particular ecological settings, and responses to new competitors or predators, climate change, and other external phenomena. The detailed case studies of faunal community composition, extinction chronologies, and demographic trends enabled by these methods examine megafaunal extinctions at scales appropriate for practical understanding of threats against particular species in their habitats today.

Somewhere beyond the sea: Human cranial remains from the Lesser Sunda Islands (Alor Island, Indonesia) provide insights on Late Pleistocene peopling of Island Southeast Asia.

The migration of anatomically modern humans (AMH) from Africa to every inhabitable continent included their dispersal through Island Southeast Asia (ISEA) to Australia. Significantly, this involved overwater dispersal through the Lesser Sunda Islands between Sunda (continental Southeast Asia) and Sahul (Australia and New Guinea). However, the timing and direction of this movement is still debated. Here, we report on human skeletal material recovered from excavations at two rockshelters, known locally as Tron Bon Lei, on Alor Island, Indonesia. The remains, dated to the Late Pleistocene, are the first anatomically modern human remains recovered in Wallacea dated to this period and are associated with cultural material demonstrating intentional burial. The human remains from Tron Bon Lei represent a population osteometrically distinct from Late Pleistocene Sunda and Sahul AMH. Instead, morphometrically, they appear more similar to Holocene populations in the Lesser Sundas. Thus, they may represent the remains of a population originally from Sunda whose Lesser Sunda Island descendants survived into the Holocene.

Least-cost pathway models indicate northern human dispersal from Sunda to Sahul.

Archaeological records from Australia provide the earliest, indirect evidence for maritime crossings by early modern humans, as the islands to the north-west of the continent (Wallacea) have never been connected to the mainland. Suggested in 1977 by Joseph B. Birdsell, the two main routes from Sunda (mainland Southeast Asia) to Sahul (Australia-New Guinea), still in debate today, are a northern route through Sulawesi with a landing in New Guinea, or a southern route through Bali, Timor and thence landing in northern Australia. Here we construct least-cost pathway models of human dispersal from Sunda to Sahul at 65 ka and 70 ka by extending previous out-of-Africa least-cost models through the digitization of these routes. We recover overwhelming support for a northern route into Sahul, with a landing location on present-day Misool Island. Minimal support is also recovered for the southern route at 70 ka, with a possible crossing to Sahul from eastern Timor. Review of archaeological records on the Wallacean islands crossed by our northern route indicate a dearth of archaeological research in this region. Meanwhile, the comparatively better studied southern islands still lack any archaeological dates comparable to those known for initial occupation in Sunda and Sahul. Based on our model results we suggest Misool Island as the initial landing site for early modern humans on Sahul and recommend a future focus on archaeological fieldwork in the northern Wallacean islands.

Seasonal migration of marsupial megafauna in Pleistocene Sahul (Australia-New Guinea).

Seasonal two-way migration is an ecological phenomenon observed in a wide range of large-bodied placental mammals, but is conspicuously absent in all modern marsupials. Most extant marsupials are typically smaller in body size in comparison to their migratory placental cousins, possibly limiting their potential to undertake long-distance seasonal migrations. But what about earlier, now-extinct giant marsupial megafauna? Here we present new geochemical analyses which show that the largest of the extinct marsupial herbivores, the enormous wombat-like Diprotodon optatum, undertook seasonal, two-way latitudinal migration in eastern Sahul (Pleistocene Australia-New Guinea). Our data infer that this giant marsupial had the potential to perform round-trip journeys of as much as 200 km annually, which is reminiscent of modern East African mammal migrations. These findings provide, to our knowledge, the first evidence for repetitive seasonal migration in any metatherian (including marsupials), living or extinct, and point to an ecological phenomenon absent from the continent since the Late Pleistocene.

Exploring morphological generality in the Old World monkey postcranium using an ecomorphological framework.

Nearly all primates are ecologically dependent on trees, but they are nonetheless found in an enormous range of habitats, from highly xeric environments to dense rainforest. Most primates have a relatively 'generalised' skeleton, enabling locomotor flexibility and facilitating other crucial functions, such as manual foraging and grooming. This paper explores the associations between habitat, locomotion and morphology in the forelimbs of cercopithecids (Old World monkeys), contextualising their skeletal ecomorphological patterns with those of other mammals, and complementing functional morphological analyses with phylogenetic comparative techniques. The ecomorphological signals present in the generalised primate postcranium, and how an ancestral arboreal 'bauplan' might be modified to incorporate terrestriality or exploit distinct arboreal substrates, are investigated. Analysis of ecomorphological variation in guenons indicates that terrestrial Chlorocebus species retain core elements of a general guenon form, with modifications for terrestriality that vary by species. Adaptation to different modes of arboreality has also occurred in Cercopithecus. The considerable morphological similarity in the guenons sampled emphasises the importance of generality in the primate postcranium - much forelimb variation appears to have emerged stochastically, with a smaller number of traits having a strong functional signal. Analysis of a broader sample of cercopithecids and comparison with felids, suids and bovids indicates that although the cercopithecid humerus has functional morphological signals that enable specimens to be assigned with a reasonable degree of certainty to habitat groups, there is considerable overlap in the specimens assigned to each habitat group. This probably reflects ecological dependence on trees, even in predominantly terrestrial species, as well as the multiple functions of the forelimb and, in some cases, wide geographic distributions that promote intraspecific variation. The use of phylogenetic correction reduced the discriminatory power of the models, indicating that, like allometry, phylogeny contains important ecomorphological information, and should not necessarily be factored out of analyses.

Climate change frames debate over the extinction of megafauna in Sahul (Pleistocene Australia-New Guinea).

Around 88 large vertebrate taxa disappeared from Sahul sometime during the Pleistocene, with the majority of losses (54 taxa) clearly taking place within the last 400,000 years. The largest was the 2.8-ton browsing Diprotodon optatum, whereas the ∼100- to 130-kg marsupial lion, Thylacoleo carnifex, the world's most specialized mammalian carnivore, and Varanus priscus, the largest lizard known, were formidable predators. Explanations for these extinctions have centered on climatic change or human activities. Here, we review the evidence and arguments for both. Human involvement in the disappearance of some species remains possible but unproven. Mounting evidence points to the loss of most species before the peopling of Sahul (circa 50-45 ka) and a significant role for climate change in the disappearance of the continent's megafauna.

Analytical framework for reconstructing heterogeneous environmental variables from mammal community structure.

We test the performance of two models that use mammalian communities to reconstruct multivariate palaeoenvironments. While both models exploit the correlation between mammal communities (defined in terms of functional groups) and arboreal heterogeneity, the first uses a multiple multivariate regression of community structure and arboreal heterogeneity, while the second uses a linear regression of the principal components of each ecospace. The success of these methods means the palaeoenvironment of a particular locality can be reconstructed in terms of the proportions of heavy, moderate, light, and absent tree canopy cover. The linear regression is less biased, and more precisely and accurately reconstructs heavy tree canopy cover than the multiple multivariate model. However, the multiple multivariate model performs better than the linear regression for all other canopy cover categories. Both models consistently perform better than randomly generated reconstructions. We apply both models to the palaeocommunity of the Upper Laetolil Beds, Tanzania. Our reconstructions indicate that there was very little heavy tree cover at this site (likely less than 10%), with the palaeo-landscape instead comprising a mixture of light and absent tree cover. These reconstructions help resolve the previous conflicting palaeoecological reconstructions made for this site.

Bovid ecomorphology and hominin paleoenvironments of the Shungura Formation, lower Omo River Valley, Ethiopia.

The Shungura Formation in the lower Omo River Valley, southern Ethiopia, has yielded an important paleontological and archeological record from the Pliocene and Pleistocene of eastern Africa. Fossils are common throughout the sequence and provide evidence of paleoenvironments and environmental change through time. This study developed discriminant function ecomorphology models that linked astragalus morphology to broadly defined habitat categories (open, light cover, heavy cover, forest, and wetlands) using modern bovids of known ecology. These models used seven variables suitable for use on fragmentary fossils and had overall classification success rates of >82%. Four hundred and one fossils were analyzed from Shungura Formation members B through G (3.4-1.9 million years ago). Analysis by member documented the full range of ecomorph categories, demonstrating that a wide range of habitats existed along the axis of the paleo-Omo River. Heavy cover ecomorphs, reflecting habitats such as woodland and heavy bushland, were the most common in the fossil sample. The trend of increasing open cover habitats from Members C through F suggested by other paleoenvironmental proxies was documented by the increase in open habitat ecomorphs during this interval. However, finer grained analysis demonstrated considerable variability in ecomorph frequencies over time, suggesting that substantial short-term variability is masked when grouping samples by member. The hominin genera Australopithecus, Homo, and Paranthropus are associated with a range of ecomorphs, indicating that all three genera were living in temporally variable and heterogeneous landscapes. Australopithecus finds were predominantly associated with lower frequencies of open habitat ecomorphs, and high frequencies of heavy cover ecomorphs, perhaps indicating a more woodland focus for this genus.

The microstratigraphy and depositional environments of Lida Ajer and Ngalau Gupin, two fossil-bearing tropical limestone caves of west Sumatra.

Lida Ajer and Ngalau Gupin are karstic caves situated in the Padang Highlands, western Sumatra, Indonesia. Lida Ajer is best known for yielding fossil evidence that places the arrival of Homo sapiens in Southeast Asia during Marine Isotope Stage 4, one of the earliest records for the region. Ngalau Gupin recently produced the first record of hippopotamid Hexaprotodon on the island, representing the only globally extinct taxon in Pleistocene deposits from Sumatra. Microstratigraphic (micromorphological) analyses were applied to unconsolidated fossil-bearing cave sediments from these two sites. We use micromorphology as part of a micro-contextualised taphonomic approach to identify the diagenetic processes affecting fossils and sediments within these caves, through phases of their depositional history. The fossil-bearing sediments in Lida Ajer have been subjected to a suite of natural sedimentation processes ranging from water action to carnivore occupation, which would indicate the fossils underwent significant reworking prior to lithification of the deposit. The results demonstrate that the base of the unconsolidated fossil-bearing sediments in Ngalau Gupin were derived from the interior of the cave, where the matrix was partially phosphatized as a result of guano-driven diagenesis. These observations can be used to test hypotheses about the integrity of incorporated vertebrate remains and to aid in local palaeoenvironmental reconstructions. The methods employed in this research have not previously been applied to cave sediments from sites in the Padang Highlands and provide key new insights into the palaeontological and natural history of the western region of Sumatra.

Expanded geographic distribution and dietary strategies of the earliest Oldowan hominins and Paranthropus.

The oldest Oldowan tool sites, from around 2.6 million years ago, have previously been confined to Ethiopia's Afar Triangle. We describe sites at Nyayanga, Kenya, dated to 3.032 to 2.581 million years ago and expand this distribution by over 1300 kilometers. Furthermore, we found two hippopotamid butchery sites associated with mosaic vegetation and a C4 grazer-dominated fauna. Tool flaking proficiency was comparable with that of younger Oldowan assemblages, but pounding activities were more common. Tool use-wear and bone damage indicate plant and animal tissue processing. Paranthropus sp. teeth, the first from southwestern Kenya, possessed carbon isotopic values indicative of a diet rich in C4 foods. We argue that the earliest Oldowan was more widespread than previously known, used to process diverse foods including megafauna, and associated with Paranthropus from its onset.

Stable isotopes provide independent support for the use of mesowear variables for inferring diets in African antelopes.

We examine the relationship between mesowear variables and carbon and nitrogen isotopes in 16 species of African antelope (Mammalia: Bovidae). We show significant differences in carbon and nitrogen isotope values between individuals exhibiting sharp versus round cusps, and high versus low occlusal relief. We show significant correlations between mesowear variables and both carbon and nitrogen isotopes. We find significant correlations between mesowear score and nitrogen, but not carbon isotopes. Finally, we find no significant correlations between hypsodonty index and either isotope examined. Our results provide strong support for the use of mesowear variables in palaeodietary reconstructions of antelopes. Our results further suggest that for the antelopes examined here, mesowear signals are a direct result of diet, while hyposodonty may be the result of phylogenetic legacy.

Speleological and environmental history of Lida Ajer cave, western Sumatra.

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. This article is part of the theme issue 'Tropical forests in the deep human past'.

Intra-skeletal vascular density in a bipedal hopping macropod with implications for analyses of rib histology.

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 < 0.05). Results confirm that the ribs are well vascularized, which facilitates bone metabolic processes such as remodelling, but the fibula also appears to be a well vascularized bone. Future research investigating human bone metabolism will benefit from examining thoracic rib or fibula samples.