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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.

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 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.

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.

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

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.

Gigantism is a common phenomenon observed in murids from islands, particularly the Indonesian islands belonging to the Lesser Sunda chain. Rats in this island group are often considerably larger than their continental sister taxa and have been reported from the islands of Timor, Sumba, and Flores. Here, we describe the first record of a giant rat from the island of Alor, at the northeastern end of the Lesser Sundas. Alormys aplini gen. et sp. nov. is described from isolated molars and a partial lower jaw from excavations in the twilight zone of Makpan Cave, a lava tube cave situated on the southwestern margin of the island. Speleological and taphonomic information suggests that these remains were deposited as a result of owl activities. Alormys aplini is characterized by its large size relative to members of Rattus and a relatively unspecialized, moderately hypsodont dentition. Among the larger rodents of the Lesser Sunda chain, Alormys most closely resembles Milimonggamys and Papagomys. Carbon and oxygen isotope analysis indicates that it consumed C3 plants almost exclusively. Its extinction in the late Holocene appears coeval with the disappearance of Timor’s and Sumba’s giant rats and may be related to increased aridity and deforestation on these islands. Keywords: Gigantism, Island rule, Muridae, Nusa Tenggara

Traditional fish vertebrae identification relies on the availability of comprehensive reference collections that include every element from the neural spine for each taxon. In regions with great taxonomic diversity, such as the Indo-Pacific, the identification of fish vertebrae to species is difficult. This results in taxonomic lists with many skeletal elements identified only to family. However family level identifications often tell us little about the environmental preferences of the fish and thus, by inference, human fishing practices. Here we apply geometric morphometrics (GM) to examine shape variations within vertebrae in modern specimens of a variety of pelagic and reef species to determine if this method can be used to reliably inform on habitat preferences. Digitized vertebral elements of reef (Acanthuridae, Balistidae, Labridae, Lethrinidae, Lutjanidae and Serranidae) and pelagic/open water (Scombridae and Carangidae) families were scored using 2D landmarks. These were subjected to Generalized Procrustes Analysis and discriminatory multivariate analyses (Linear Discriminant Analysis and Discriminant Function Analysis) in order to assess whether shape can be used to differentiate habitats. Our results suggest that geometric morphometrics do allow the differentiation of habitat in vertebrae and provide an alternative method for the classification of archaeological fish assemblages. These analyses were applied to a sample of archaeological fish remains from a site in Alor Island (Nusa Tenggara Timur, Indonesia) and compared with the results of an earlier traditional comparative icthyoarchaeological analysis. We found that the main component of the Pleistocene marine human diet comprised reef species, with the sporadic addition of open water species, supporting the pattern recorded with traditional analyses. This methodology could be widely applied to archaeological fish material from across the Indo-Pacific allowing a greater number of bones in assemblages to contribute to insights into human exploitation of coastal habitats and fishing techniques over time.

Chinchilla in the Western Downs Region of Queensland is home to the Chinchilla Rifle Range, a palaeontological site that has produced a significant well-preserved Pliocene vertebrate assemblage. Here, we describe and discuss the histology of a crocodile tooth recovered from the ca 3.5-million-year-old Chinchilla Sand deposit in the Rifle Range. The tooth is from the posterior jaw and likely belongs to a species of Paludirex. We discuss the tooth micro-morphology in relation to what is known about tooth histology in extant and extinct crocodylians with brevirostine and platyrostral skull morphology. We hypothesised that there should be several similarities in the tooth micro-structures between related extinct and extant taxa. We found that the Chinchilla Sand fossil tooth is characterised by thin enamel that is likely prismless but shows incremental striations (also seen in dentine), similar to other crocodylians. This short study highlights the importance of microscopic techniques applied to fossil material. With further fossil evidence emerging from Chinchilla, and application of three-dimensional microscopy techniques to understand the nature of Paludirex enamel prisms, a better understanding of reptile palaeobiology can be developed for Queensland and Australia.

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. Stable isotope data for Southeast Asian mammals across the Quaternary period shed light on environmental change from the Early Pleistocene to the Holocene epoch, contextualizing hominin evolution and megafauna extinction in the region.

The significance of the role of birds in hominin evolution in Island Southeast Asia is not clear. Few avian vertebrate deposits have been recovered from archaeological or fossil sites in the region, and their association with either hominin or natural deposition in caves and rock shelters complicates their usefulness in hominin behavioural and palaeoecological reconstructions. In this paper, we assess the taphonomic history of the Pleistocene avian vertebrate remains recovered from Laili Cave, Timor-Leste, dated to between ca. 44.6 to 11.2 ka and in association with abundant lithic material. We use avian taxonomic composition, skeletal element abundance, and bone surface modification data to determine the agent of avian skeletal deposition. Our analyses indicate that the small grassland and woodland birds (quail, buttonquail, song birds), which dominate the assemblage, were deposited by avian predators (probably barn owls) throughout the sequence. Humans possibly hunted the small quantity of larger birds (imperial pigeon, duck). The bird remains suggest that grasslands, woodland savannahs, wetlands, and forest environments were present locally during the Pleistocene.

The thousands of islands east of Huxley’s Line have never formed a single land mass or been connected to Sunda or Sahul. The earliest records of hominins in this area are stone tools recovered from Pleistocene deposits on Flores and Sulawesi. Subsistence by these hominins as well as the later subsistence patterns exhibited by Homo floresiensis suggest that exploitation of marine resources was, at best, rare and opportunistic. Likewise, the fragmentary hominin remains recovered from Late Pleistocene deposits from Callao Cave in the Philippines exploited large game at the expense of marine resources. In contrast, the earliest zooarchaeological records of modern humans are dominated by marine fish and shellfish and include the earliest evidence of pelagic fishing using shell tools, implying complex fish-capture technology. Pleistocene lithic assemblages on these islands are unspecialized, indicating reduction of predominantly locally available stone to produce flakes and irregularly retouched pieces. By the ...

Caves have been an important source of vertebrate fossils for much of Southeast Asia, particularly for the Quaternary. Despite this importance, the mechanisms by which vertebrate remains accumulate and preserve in Southeast Asian caves has never been systematically reviewed or examined. Here, we present the results of three years of cave surveys in Indonesia and Timor-Leste, describing cave systems and their attendant vertebrate accumulations in diverse geological, biogeographical, and environmental settings. While each cave system is unique, we find that the accumulation and preservation of vertebrate remains are highly dependent on local geology and environment. These factors notwithstanding, we find the dominant factor responsible for faunal deposition is the presence or absence of biological accumulating agents, a factor directly dictated by biogeographical history. In small, isolated, volcanic islands, the only significant accumulation occurs in archaeological settings, thereby limiting our understanding of the palaeontology of those islands prior to human arrival. In karstic landscapes on both oceanic and continental islands, our understanding of the long-term preservation of vertebrates is still in its infancy. The formation processes of vertebrate-bearing breccias, their taphonomic histories, and the criteria used to determine whether these represent syngenetic or multiple deposits remain critically understudied. The latter in particular has important implications for arguments on how breccia deposits from the region should be analysed and interpreted when reconstructing palaeoenvironments.

Genetic evidence for anatomically modern humans (AMH) out of Africa before 75 thousand years ago (ka)1 and in island southeast Asia (ISEA) before 60 ka (93–61 ka)2 predates accepted archaeological records of occupation in the region3. Claims that AMH arrived in ISEA before 60 ka (ref. 4) have been supported only by equivocal5 or non-skeletal evidence6. AMH evidence from this period is rare and lacks robust chronologies owing to a lack of direct dating applications7, poor preservation and/or excavation strategies8 and questionable taxonomic identifications9. Lida Ajer is a Sumatran Pleistocene cave with a rich rainforest fauna associated with fossil human teeth7,10. The importance of the site is unclear owing to unsupported taxonomic identification of these fossils and uncertainties regarding the age of the deposit, therefore it is rarely considered in models of human dispersal. Here we reinvestigate Lida Ajer to identify the teeth confidently and establish a robust chronology using an integrated dating approach. Using enamel–dentine junction morphology, enamel thickness and comparative morphology, we show that the teeth are unequivocally AMH. Luminescence and uranium-series techniques applied to bone-bearing sediments and speleothems, and coupled uranium-series and electron spin resonance dating of mammalian teeth, place modern humans in Sumatra between 73 and 63 ka. This age is consistent with biostratigraphic estimations7, palaeoclimate and sea-level reconstructions, and genetic evidence for a pre-60 ka arrival of AMH into ISEA2. Lida Ajer represents, to our knowledge, the earliest evidence of rainforest occupation by AMH, and underscores the importance of reassessing the timing and environmental context of the dispersal of modern humans out of Africa.

SUMMARYExtinctions have altered island ecosystems throughout the late Quaternary. Here, we review the main historic drivers of extinctions on islands, patterns in extinction chronologies between islands, and the potential for restoring ecosystems through reintroducing extirpated species. While some extinctions have been caused by climatic and environmental change, most have been caused by anthropogenic impacts. We propose a general model to describe patterns in these anthropogenic island extinctions. Hunting, habitat loss and the introduction of invasive predators accompanied prehistoric settlement and caused declines of endemic island species. Later settlement by European colonists brought further land development, a different suite of predators and new drivers, leading to more extinctions. Extinctions alter ecological networks, causing ripple effects for islands through the loss of ecosystem processes, functions and interactions between species. Reintroduction of extirpated species can help restore ecosystem function and processes, and can be guided by palaeoecology. However, reintroduction projects must also consider the cultural, social and economic needs of humans now inhabiting the islands and ensure resilience against future environmental and climate change.

The Pig-footed Bandicoot, Chaeropus ecaudatus, an extinct arid-adapted bandicoot, was named in 1838 based on a specimen without a tail from the Murray River in New South Wales. Two additional species were later named, C. castanotis and C. occidentalis, which have since been synonymised with C. ecaudatus. Taxonomic research on the genus is rather difficult because of the limited material available for study. Aside from the types of C. castanotis and C. occidentalis housed at the Natural History Museum in London, and the type of C. ecaudatus at the Australian Museum in Sydney, there are fewer than 30 other modern specimens in other collections scattered around the world. Examining skeletal and dental characters for several specimens, and using a combination of traditional morphology, morphometrics, palaeontology and molecular phylogenetics, we have identified two distinct species, C. ecaudatus and C. yirratji sp. nov., with C. ecaudatus having two distinct subspecies, C. e. ecaudatus and C. e. occidentalis. We use palaeontological data to reconstruct the pre-European distribution of the two species, and review the ecological information known about these extinct taxa.

Despite its largely hyper-arid and inhospitable climate today, the Arabian Peninsula is emerging as an important area for investigating Pleistocene hominin dispersals. Recently, a member of our own species was found in northern Arabia dating to ca. 90 ka, while stone tools and fossil finds have hinted at an earlier, middle Pleistocene, hominin presence. However, there remain few direct insights into Pleistocene environments, and associated hominin adaptations, that accompanied the movement of populations into this region. Here, we apply stable carbon and oxygen isotope analysis to fossil mammal tooth enamel (n = 21) from the middle Pleistocene locality of Ti’s al Ghadah in Saudi Arabia associated with newly discovered stone tools and probable cutmarks. The results demonstrate productive grasslands in the interior of the Arabian Peninsula ca. 300–500 ka, as well as aridity levels similar to those found in open savannah settings in eastern Africa today. The association between this palaeoenvironmental information and the earliest traces for hominin activity in this part of the world lead us to argue that middle Pleistocene hominin dispersals into the interior of the Arabian Peninsula required no major novel adaptation. Stable carbon and oxygen isotope analysis of mammal teeth associated with stone tools and cut-marked bone dated to between 300,000 and 500,000 years ago reveals that, at the time of the earliest-known hominin presence, the Arabian peninsula was home to productive grasslands similar to modern-day African savannahs.

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.

The Sangihe-Talaud Archipelago represents a group of 77 remote islands located between the Philippines and North Sulawesi, in the northern sector of Wallacea. The Talaud Islands have a rich and significant archaeological record going back to the Pleistocene and are instrumental in understanding Pleistocene colonisation of small islands and later models of Austronesian language dispersal. Here we report on vertebrate material excavated from Leang Mande'et, a late Holocene rockshelter on Karakelang, the main island in the Talaud group. The site represents a periodically occupied shelter used for gardening. Fauna recovered predominately comprises murid elements, with at least four taxa (Rattus rattus, Rattus exulans and two Melomys species) found. The rodents show clear signs of being deposited by raptors rather than humans, unlike the small number of fish remains also recovered. DNA sequences from several Rattus rattus specimens are referrable to Rattus rattus complex lineage IV, a lineage with a southern Indochinese origins and broad modern representation through Indonesia and the Philippines. The lack of any Rattus rattus complex lineage II from Leang Mande’et suggests that the first agricultural inhabitants of this island did not come from Taiwan or northern Indochina, but rather that they either originated from the south or that, once established in the Talauds, opened up significant trade networks to the south and in the process acquired a significant pest.

The common wombat (Vombatus ursinus) is equipped with a set of physiological and morphological adaptations suited to a fossorial lifestyle. These allow wombats to engage in efficient scratch-digging and maintaining a low basal metabolic rate while living underground. While bone microstructure has been described for several subterranean animals, wombat bone histology has received very little attention to date. Here, we present preliminary insights into bone histology in modern adult V. ursinus (Mt Fairy, New South Wales) and Pleistocene fossil Vombatus sp. (Bakers Swamp, New South Wales) midshaft humeri. The modern sample was well preserved, allowing us to identify varying bone tissue types (woven, parallel-fibred, lamellar). The sample showed vascularity composed of primary and secondary osteons, and simple longitudinal and radial vessels. We also observed evidence for Haversian remodelling (i.e. localised replacement of pre-existing bone) and coarse compact cancellous bone within the inner cortex of the diaphysis. The fossil histology was poorly preserved, but likely showed bone matrix organisation similar to the modern specimen. We use these preliminary data to discuss hypotheses for wombat forelimb biomechanical and physiological microscopic adaptation to a burrow environment. We encourage future intraskeletal examination of microstructure in wombat populations to better inform their ecological adaptations and behaviour in palaeontological contexts.

International audience; We report archaeological findings from a significant new cave site on Alor Island, Indonesia, with an in situ basal date of 40,208-38,454 cal BP. Twenty thousand years older than the earliest Pleistocene site previously known from this island, Makpan retains dense midden deposits of marine shell, fish bone, urchin and crab remains, but few terrestrial species; demonstrating that protein requirements over this time were met almost exclusively from the sea. The dates for initial occupation at Makpan indicate that once Homo sapiens moved into southern Wallacea, settlement of the larger islands in the archipelago occurred rapidly. However, the Makpan sequence also suggests that the use of the cave following initial human arrival was sporadic prior to the terminal Pleistocene about 14,000 years ago, when occupation became intensive, culminating in the formation of a midden. Like the coastal sites on the larger neighbouring island of Timor, the Makpan assemblage shows that maritime technology in the Pleistocene was highly developed in this region. The Makpan assemblage also contains a range of distinctive personal ornaments made on Nautilus shell, which are shared with sites located on Timor and Kisar supporting connectivity between islands from at least the terminal Pleistocene. Makpan's early inhabitants responded to sea-level change by altering the way they used both the site and local resources. Marine food exploitation shows an initial emphasis on sea-urchins, followed by a subsistence switch to molluscs, barnacles, and fish in the dense middle part of the sequence, with crabs well represented in the later occupation. This new record provides further insights into early modern human movements and patterns of occupation between the islands of eastern Nusa Tenggara from ca. 40 ka

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.

Fragmentary long bone material from fossil Carnivora is rarely considered to support palaeoenvironmental reconstructions. Here, we use morphometry of the radius in extant carnivorans of the dog family (Canidae) to reconstruct the palaeobiology of extinct canids from Olduvai Gorge, Tanzania (Bed I and II) and Koobi Fora, Kenya. We use radius morphometrics to predict adaptation to prey size and introduce a new method for quantifying canid habitat adaptations based on the geographic distributions of the extant species sampled. Linear Discriminant Function Analyses (DFA) and cluster neighbour-joining algorithms are employed to investigate radial morphometrics as described by 29 linear measurements. Results of our analyses suggest that a phylogenetic signal is present in radial morphometrics, even if it does not allow us to accurately discriminate among genera. A binary prey size categorisation of “small-medium” versus “large” prey can be more accurately predicted than a habitat categorisation scheme (Open, Mixed, Closed). The East African fossil specimens examined show morphometric affinities with the golden jackal (Canis aureus) and coyote (Canis latrans) and are likely attributable to the genus Canis. Fragmentary fossil specimens from Olduvai Gorge are predicted as habitat generalists (Open for Bed I and Mixed for Bed II) adapted for hunting small-medium prey, whereas the specimen from Koobi Fora was predicted as inhabiting mixed habitats and adapted for killing large prey. This study supports the inclusion of fossil Canidae in palaeoecological analyses attempting to clarify the palaeoenvironment of early hominin fossil sites.

An obvious but key prerequisite to testing hypotheses concerning the role of humans in the extinction of late Quaternary ‘megafauna’ is demonstrating that humans and the extinct taxa overlapped, both temporally and spatially. In many regions, a paucity of reliably dated fossil occurrences of megafauna makes it challenging, if not impossible, to test many of the leading extinction hypotheses. The giant monitor lizards of Australia are a case in point. Despite commonly being argued to have suffered extinction at the hands of the first human colonisers (who arrived by 50 ka), it has never been reliably demonstrated that giant monitors and humans temporally overlapped in Australia. Here we present the results of an integrated U–Th and 14C dating study of a late Pleistocene fossil deposit that has yielded the youngest dated remains of giant monitor lizards in Australia. The site, Colosseum Chamber, is a cave deposit in the Mt Etna region, central eastern Australia. Sixteen new dates were generated and demonstrate that the bulk of the material in the deposit accumulated since ca. 50 ka. The new monitor fossil is, minimally, 30 ky younger than the previous youngest reliably dated record for giant lizards in Australia and for the first time, demonstrates that on a continental scale, humans and giant lizards overlapped in time. The new record brings the existing geochronological dataset for Australian giant monitor lizards to seven dated occurrences. With such sparse data, we are hesitant to argue that our new date represents the time of their extinction from the continent. Rather, we suspect that future fossil collecting will yield new samples both older and younger than 50 ka. Nevertheless, we unequivocally demonstrate that humans and giant monitor lizards overlapped temporally in Australia, and thus, humans can only now be considered potential drivers for their extinction.

The species-level systematics of the marsupial family Phalangeridae, particularly Phalanger, are poorly understood, due partly to the family’s wide distribution across Australia, New Guinea, eastern Indonesia, and surrounding islands. In order to refine the species-level systematics of Phalangeridae, and improve our understanding of their evolution, we generated 36 mitochondrial ND2 DNA sequences from multiple species and sample localities. We combined our new data with available sequences and produced the most comprehensive molecular phylogeny for Phalangeridae to date. Our analyses (1) strongly support the monophyly of the three phalangerid subfamilies (Trichosurinae, Ailuropinae, Phalangerinae); (2) reveal the need to re-examine all specimens currently identified as ‘Phalanger orientalis’; and (3) suggest the elevation of the Solomon Island P. orientalis subspecies to species level (P. breviceps Thomas, 1888). In addition, samples of P. orientalis from Timor formed a clade, consistent with an introduction by humans from a single source population. However, further research on east Indonesian P. orientalis populations will be required to test this hypothesis, resolve inconsistencies in divergence time estimates, and locate the source population and taxonomic status of the Timor P. orientalis.

Fossils from caves in the Manning Karst Region, New South Wales, Australia have long been known, but until now have never been assessed for their palaeontological significance. Here, we report on late Quaternary faunal records from eight caves in the region. Extinct Pleistocene megafaunal taxa are recognised in two systems and include giant echidnas (Tachyglossidae gen. et sp. indet.), devils (Sarcophilus laniarius), koalas (Phascolarctos stirtoni), marsupial ‘lions’ (Thylacoleo carnifex), and kangaroos (Macropus giganteus titan). Some caves contain skeletal remains of introduced exotics such as sheep and dogs, but also provide a rich record of small-bodied native species including Eastern Bettongs (Bettongia gaimardi), Eastern Chestnut Mice (Pseudomys gracilicaudatus), and White-footed Rabbit Rats (Conilurus albipes). These endemics are either locally extirpated or have suffered total extinction in the historic period. Their skeletal and dental remains were recorded as unmineralised surface specimens in the caves, indicating that they are recent in age. Extant populations have never been recorded locally, thus, their probable loss from the region in historic times had gone unnoticed in the absence of palaeo-evidence. Our findings suggest that the supposed habitat tolerances of such species have been substantially underestimated. It is highly likely that modern populations have suffered niche contraction since the time of European colonisation of the continent. The local extirpations of several species of digging mammal has likely led to decreased functionality of the current ecosystem.

Alarm over the prospects for survival of species in a rapidly changing world has encouraged discussion of translocation conservation strategies that move beyond the focus of 'at-risk' species. These approaches consider larger spatial and temporal scales than customary, with the aim of recreating functioning ecosystems through a combination of large-scale ecological restoration and species introductions. The term 'rewilding' has come to apply to this large-scale ecosystem restoration program. While reintroductions of species within their historical ranges have become standard conservation tools, introductions within known paleontological ranges-but outside historical ranges-are more controversial, as is the use of taxon substitutions for extinct species. Here, we consider possible conservation translocations for nine large-bodied taxa in tropical Asia-Pacific. We consider the entire spectrum of conservation translocation strategies as defined by the IUCN in addition to rewilding. The taxa considered are spread across diverse taxonomic and ecological spectra and all are listed as 'endangered' or 'critically endangered' by the IUCN in our region of study. They all have a written and fossil record that is sufficient to assess past changes in range, as well as ecological and environmental preferences, and the reasons for their decline, and they have all suffered massive range restrictions since the late Pleistocene. General principles, problems, and benefits of translocation strategies are reviewed as case studies. These allowed us to develop a conservation translocation matrix, with taxa scored for risk, benefit, and feasibility. Comparisons between taxa across this matrix indicated that orangutans, tapirs, Tasmanian devils, and perhaps tortoises are the most viable taxa for translocations. However, overall the case studies revealed a need for more data and research for all taxa, and their ecological and environmental needs. Rewilding the Asian-Pacific tropics remains a controversial conservation strategy, and would be difficult in what is largely a highly fragmented area geographically.

The Pliocene fossil record of Australia has revealed the oldest known occurrences of modern peramelemorphian (bandicoot and bilbies) genera such as Perameles, cf. Peroryctes, and Chaeropus. Recent phylogenetic analyses based on morphology have questioned the previously accepted understanding about generic relationships of some of these Pliocene taxa. These doubts limit our ability to develop independent divergence models based on molecular data because they depend on fossil records in order to calibrate minimum rates of change. Hence, there is a need to critically review the Pliocene fossil record. To this end, we have examined Pliocene specimens of peramelemorphians in museum collections across Australia and performed a comprehensive phylogenetic analysis based on dental and cranial morphology to review and accordingly revise subfamilial taxonomy. As part of this revision, we describe here two new species, one from the Chinchilla Local Fauna (LF) in Queensland (Perameles wilkinsonorum, sp. nov.) and one from both the Big Sink LF in the Wellington Caves area and the Bow LF in New South Wales (Silvicultor karae, gen. et sp. nov.). We reassign the two ‘Peroryctes’ species from Hamilton LF, Victoria, to a new genus (Silvicultor). We summarize the distribution of peramelemorphians during the Pliocene and show how climate change appears to have shaped their subsequent Quaternary distribution. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP Citation for this article: Travouillon, K. J., J. Louys, G. J. Price, M. Archer, S. J. Hand, and J. Muirhead. 2017. A review of the Pliocene bandicoots of Australia, and descriptions of new genus and species. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2017.1360894.

Understanding the evolutionary history of the herbivore niche within African bovids has traditionally relied on examining anatomical adaptations to diet, particularly those related to digestive strategy. More recently, mesowear and stable isotope analyses have been used to great effect to reconstruct dietary preferences. We use these dietary proxies to construct a morphology-free dietary ecospace and examine the topology of the phylogenetic relationships of African bovids mapped onto this ecospace. The reconstructed dietary ecospace provides evidence for four distinct dietary classes: species with C3- or C4-dominated diets that produce low or high occlusal relief, likely related to diets high or low in abrasives, respectively. We detected no evidence for a discrete mixed feeder category; the species often categorized as such represent the end members of groups of species with either C3- or C4- dominated diets. Our analysis reveals high variability within the C4 grazing ecospace, and phylogenetic evidence indicates at least two pathways to grazing, likely related to the abrasive qualities of ingested food, which may be determined by the moisture content or the height of consumed grasses. These different pathways probably contribute to the high diversity of African grazers, both today and in the fossil record. C3 browsers (non-frugivores) also display a high degree of variation, but there are no species associated with highly abrasive diets and there is evidence for only a single evolutionary pathway. We find evidence for only one evolutionary route towards frugivory, which includes species with diets that produce both high and low occlusal reliefs. The cause of abrasive wear in frugivores may be related to grit and/or the hard parts of fruits, but this requires further examination.

Mesowear analysis is a form of dental wear analysis used to infer the diets of herbivorous mammal species. It makes use of percentage indices of blunt, round and sharp cusp shape and high occlusal relief to classify the diet of species into one of three categories: browser, grazer or mixed feeder. Previously, this form of analysis has been limited to placental mammals, restricting the use of such analyses in Australia where the dominant herbivorous mammalian fauna consist of marsupials. In order to address this limitation, mesowear variables of extant marsupials were examined to determine whether their diets can accurately be predicted using mesowear analyses. Discriminant Function Analysis of mesowear variables and analysis of variance (ANOVA) of univariate mesowear scores for marsupial species demonstrate that mesowear analysis can be used to classify marsupial diets. A dataset of 24 typical marsupial species considered to be representative of the three dietary categories with respect to mesowear was generated and significantly increased cross-validated classification levels from 74.4% to 100% for the second molar. Mesowear analysis for marsupial species is most effective for the second molar with high predictive power also being evident for the first and third molars. When mesowear analysis was applied to six Plio-Pleistocene macropods (Marsupialia: Macropodidae) from the Darling Downs region, southeast Queensland, all species were classified as mixed feeders with the exception of Protemnodon roechus which was classified as a grazer. This study demonstrates the effectiveness of mesowear analysis as a dietary proxy for herbivorous marsupial species.

Much of Southeast Asia's large terrestrial carnivores appeared, evolved and disappeared from the region for reasons that remain poorly understood. Two of the most significant extinctions are represented by the charismatic Pleistocene megacarnivores Pachycrocuta and Pliocrocuta. Southeast Asia hosts the last populations of these species globally. Their persistence in southern China until the late Pleistocene suggests their extinction was not tied to that of the machairodont cats, which like the rest of the world became extinct sometime in the early Pleistocene in this region. Instead the disappearance of the hyenids is probably related to climate change and deteriorating environmental conditions. There is some evidence that the wolf and domesticated dog first appeared in Southeast Asia, although confirmation of this awaits more detailed fossil records. There does not appear to be a large carnivore guild turnover of the same scale or time as recorded in Europe and Africa, although an extinction event in the late Pleistocene is provisionally recorded. Environmental changes and fluctuating sea levels have had a unique impact on the region's large carnivore guild. Several large carnivores from Java show unique genetic and morphological variations, and this could potentially be related to the connection between Java and the Indochinese mainland sometime during the middle Pleistocene. The effects of islands on the large carnivores are complicated and at times contradictory. Nevertheless, periods of isolation of large carnivores on Java, Sumatra and Borneo from the continent had impacts on both extinctions and speciations, with at least one well documented endemic large carnivore evolving in Sundaland (Sunda clouded leopard). Hunting and deforestation ongoing since the mid- to late Holocene means that many extant members of the large carnivore guild are at high risk of extinction.

The finding of archaeological evidence predating 1 Ma and a small hominin species (Homo floresiensis) on Flores, Indonesia, has stimulated much research on its origins and ancestry. Here we take a different approach and examine two key questions – 1) how did the ancestors of H. floresiensis reach Flores and 2) what are the prospects and difficulties of estimating the likelihood of hominin persistence for over 1 million years on a small island? With regard to the first question, on the basis of the biogeography we conclude that the mammalian, avian, and reptilian fauna on Flores arrived from a number of sources including Java, Sulawesi and Sahul. Many of the terrestrial taxa were able to float or swim (e.g. stegodons, giant tortoises and the Komodo dragon), while the rodents and hominins probably accidentally rafted from Sulawesi, following the prevailing currents. The precise route by which hominins arrived on Flores cannot at present be determined, although a route from South Asia through Indochina, Sulawesi and hence Flores is tentatively supported on the basis of zoogeography. With regards to the second question, we find the archaeological record equivocal. A basic energetics model shows that a greater number of small-bodied hominins could persist on Flores than larger-bodied hominins (whether H. floresiensis is a dwarfed species or a descendent of an early small-bodied ancestor is immaterial here), which may in part explain their apparent long-term success. Yet the frequent tsunamis and volcanic eruptions in the region would certainly have affected all the taxa on the island, and at least one turnover event is recorded, when Stegodon sondaari became extinct. The question of the likelihood of persistence may be unanswerable until we know much more about the biology of H. floresiensis.

We document morphological variation (both geographical and sexual) in the dentition of the extant koala, Phascolarctos cinereus, in order to facilitate discrimination of species boundaries in extinct phascolarctids. Considerable variation is evident in dental structures previously used to diagnose several phascolarctid fossil species. Consistent patterns of morphological variation are not evident between sexes or geographic regions, with variation as great between samples as within them. Metric variation is evident between the sexes in upper molar dimensions with Victorian (southern) males significantly larger than Victorian females, although this is not reflected in lower molar dimensions or in the Queensland (northern) sample. Male koalas from southern populations generally display significantly larger molars than their northern counterparts; however this trend is not evident in female upper molar dimensions. In both males and females, some, but not all, lower molar dimensions are larger in southern pop...

Mammalian carnivores are rarely incorporated in paleoenvironmental reconstructions, largely because of their rarity within the fossil record. However, multivariate statistical modeling can be successfully used to quantify specific anatomical features as environmental predictors. Here we explore morphological variability of the humerus in a closely related group of predators (Felidae) to investigate the relationship between morphometric descriptors and habitat categories. We analyze linear measurements of the humerus in three different morphometric combinations (log-transformed, size-free, and ratio), and explore four distinct ways of categorizing habitat adaptations. Open, Mixed, and Closed categories are defined according to criteria based on traditional descriptions of species, distributions, and biome occupancy. Extensive exploratory work is presented using linear discriminant analyses and several fossils are included to provide paleoecological reconstructions.We found no significant differences in the predictive power of distinct morphometric descriptors or habitat criteria, although sample splitting into small and large cat guilds greatly improves the stability of the models. Significant insights emerge for three long-canine cats:Smilodon populator,Paramachairodus orientalis, andDinofelissp. from Olduvai Gorge (East Africa).S. populatorandP. orientalisare both predicted to have been closed-habitat adapted taxa. The false “sabertooth”Dinofelissp. from Olduvai Gorge is predicted to be adapted to mixed habitat. The application of felid humerus ecomorphology to the carnivoran record of Olduvai Gorge shows that the older stratigraphic levels (Bed I, 1.99–1.79 Ma) included a broader range of environments than Beds II or V, where there is an abundance of cats adapted to open environments.

Stegodons are a commonly recovered extinct proboscidean (elephants and allies) from the Pleistocene record of Southeast Asian oceanic islands. Estimates on when stegodons arrived on individual islands and the timings of their extinctions are poorly constrained due to few reported direct geochronological analyses of their remains. Here we report on uranium-series dating of a stegodon tusk recovered from the Ainaro Gravels of Timor. The six dates obtained indicate the local presence of stegodons in Timor at or before 130 ka, significantly pre-dating the earliest evidence of humans on the island. On the basis of current data, we find no evidence for significant environmental changes or the presence of modern humans in the region during that time. Thus, we do not consider either of these factors to have contributed significantly to their extinction. In the absence of these, we propose that their extinction was possibly the result of long-term demographic and genetic declines associated with an isolated island population.

Morphological convergence amongst species inhabiting similar environments but having different evolutionary histories is a concept central to evolutionary biology. Cases of divergent evolution, where there is morphological divergence between closely related species exploiting different environments, are less well studied. Here we show divergent evolution in the morphology of the proximal phalanges of several closely related African antelope species inhabiting different environments. This morphological divergence was consistently observed in both a neutral morphospace and an externally ordinated morphospace. Divergence, but not convergence, was also observed when size and shape were considered independently. Finally, convergent evolution of the morphology of the proximal phalanges was observed, but only in the externally ordinated morphospace. Size shows less correlation with phylogeny than does shape. Therefore, we suggest that divergence in size will occur more readily when a species encounters new environmental conditions than divergence in shape. These findings are compatible with observations of rapid dwarfing on islands (Foster’s rule).

Analysis of dental mesowear is a useful tool to examine dietary strategies in ungulates, and provides a palaeoenvironmental proxy at archaeological and palaeontological sites. We examine the distribution of traditional qualitative mesowear variables amongst 33 species of extant African antelope. We also examine the relationship of our dataset with the original dataset of Fortelius and Solounias (2000) . We find that confining mesowear analyses to antelope species alone reduces their discriminating power relative to that found in analyses using a broader range of taxa. Variability in mesowear scores along the toothrow suggest that different teeth do not wear equivalently over an individual’s life-time, so it is important to confine comparisons to individual tooth positions for palaeodietary classifications. Mesowear scores from the third upper molar were found to have the greatest discriminating power. Frugivores displayed unique mesowear signatures, suggesting that this trophic guild should be taken into consideration separately in future mesowear analyses. These results have implications which can improve the application and the discriminatory power of mesowear analyses for the determination of the palaeodiets of extinct taxa, and, by extension, their palaeoenvironments.

Aim To quantify how mammal community structure relates to heterogeneity of vegetation for palaeoecological reconstructions, and to test whether historical or environmental factors are more important in structuring communities. Location Sixty-three natural protected areas in Asia, Africa and South and Central America. Methods We defined faunal communities by allocating species to ecological guilds and calculating proportional representation within each guild. Vegetation heterogeneity for each natural protected area was calculated from satellite images. The relationship between these ecospaces was calculated using canonical correlations analysis, redundancy analysis and principal components analysis. We expected that large, herbivorous mammals would be most strongly correlated with open areas. Convergence was tested by independently eliminating the effects of geography and vegetation heterogeneity on the structure of the mammal communities. We expected that vegetation would more strongly structure communities than geographical position. Results We show that the guild structure of communities across habitats is significantly correlated with vegetation heterogeneity. The highest correlation was between small, scansorial-arboreal secondary consumers and heavy tree cover. The first convergence analysis shows American communities distinguished from Asian and African communities; these latter communities show a remarkable convergence in structure. Historical factors only affected the continent whose mammals had experienced a long period of isolation. The second convergence analysis shows that almost all biomes have the same or very similar community structure regardless of continent. Main conclusions Communities from the same environments in different continents showed remarkable convergence. Communities from the same continents only converged when those continents shared a recent geological and biological history. These results suggest that historical and environmental factors are operating over different timescales. This study confirms that environmental reconstructions made on the basis of whole communities will accurately reflect the environment that the community lived in. However, reconstructions made for fossil sites in deep time need to take historical factors into consideration. Small, arboreal and scansorial secondary consumers show the strongest correlation with vegetation, correlating with continuous tree canopy cover. This relationship allows simple reconstructions of the amount of tree cover occurring in a landscape from the proportion of species from the community falling in this ecological guild.

Partial crania of two fossil species of koala (family Phascolarctidae) from Oligo-Miocene deposits in the Riversleigh World Heritage Area, one referable to Litokoala kutjamarpensis and another to Nimiokoala greystanesi, are described. Comparison with the extant koala Phascolarctos cinereus and other diprotodontian marsupials reveals a high degree of similarity in basicranial morphology between fossil and modern phascolarctids but substantial differences in the architecture of the masticatory system. Key specialisations present in Phascolarctos but absent in both Litokoala and Nimiokoala include forward displacement of the palate, enlargement of the occlusal surface of the molar teeth, thickening of the maxillae above the toothrow with resultant lowering of the occlusal plane of the cheekteeth relative to the glenoid fossa, and a decrease in the size of the pterygoid fossae. These extreme aspects of the cranial morphology of Phacolarctos probably reflect its dependence on eucalypt leaves, a nutrie...

Louys, J., Black, K., Archer, M., Hand, S.J. & Godthelp, H., June, 2007. Descriptions of koala fossils from the Miocene of Riversleigh, northwestern Queensland and implications for Litokoala (Marsupialia, Phascolarctidae). Alcheringa 31, 99‐110. ISSN 0311-5518. Litokoala is revised on the basis of new material from the early to middle Miocene deposits of Riversleigh. A new species of Litokoala is described from the System B Outasite Local Fauna. Litokoala garyjohnstoni sp. nov. possesses a unique combination of both plesiomorphic and apomorphic dental features, making its phylogenetic position within Litokoala difficult to interpret. In some aspects of dental morphology, L. garyjohnstoni is convergent on the early Miocene phascolarctid Nimiokoala greystanesi Black & Archer, 1997. The dentition of a partial skull of Litokoala from Jims Carousel Site, System C is also described. On the basis of this dentition, it is suggested that the morphological variation between L. kutjamarpensis Stirton, 1967 and L. ka...

The extinction of large-bodied taxa from the Pleistocene in Southeast Asia is examined. Although the chronological resolution of these extinctions is poor, and number of excavations in the region relatively few, broad characteristics of these extinctions can be described. Many taxa which became extinct appear to have been endemic to regions within Southeast Asia, while some taxa which experienced extinction or severe range reduction occurred in several regions. Members of the latter group include proboscideans (Stegodon and Palaeloxodon), the pygmy hippopotamus (Hexaprotodon), the orangutan (Pongo), hyenas (Crocuta and Hyaena), the giant panda (Ailuropoda), tapirs (Tapirus and Megatapirus), rhinoceroses (Rhinoceros), and the giant Asian ape Gigantopithecus. The loss of these species cannot be assigned to a single cause. Rather their disappearance is likely tied to both climatic and human agents. Unlike other regions which experienced megafauna extinctions, eustatic changes in sea level in Southeast Asia seems to have been an important factor.

Current lists of species-level representation in faunas from 80 Cenozoic fossil localities at the Riversleigh World Heritage Area have been compiled by review of recorded occurrences of taxa obtained from both published and unpublished sources. More than 290 species-level taxa are represented, comprising mammals, amphibians, reptiles, birds, fishes, molluscs and crustaceans. The data are presented for the purpose of ongoing palaeoecological and biochronological studies.

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.

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.

The Chinchilla Local Fauna is a diverse assemblage of both terrestrial and aquatic Pliocene vertebrates from the fluviatile Chinchilla Sand deposits of southeastern Queensland, Australia. It represents one of Australia's few but exceptionally rich Pliocene vertebrate localities, and as such is an important source of paleoecological data concerning Pliocene environmental changes and its effects on ecosystems. Prior inferences about the paleoenvironment of this locality made on the basis of qualitative observations have ranged from grassland to open woodland to wetland. Examination of the carbon and oxygen isotopes in the tooth enamel of marsupials from this site represents a quantitative method for inferring the paleoenvironments and paleoecology of the fossil fauna. Results from Chinchilla show that Protemnodon sp. indet. consumed both C3 and C4 photosynthesis plant types (mean δ(13)C = -14.5±2.0‰), and therefore probably occupied a mixed vegetation environment. Macropus sp. indet. from Chinchilla also consumed a mixed diet of both C3 and C4 plants, with more of a tendency for C4 plant consumption (mean δ(13)C = -10.3±2.3‰). Interestingly, their isotopic dietary signature is more consistent with tropical and temperate kangaroo communities than the sub-tropical communities found around Chinchilla today. Other genera sampled in this study include the extinct kangaroo Troposodon sp. indet. and the fossil diprotodontid Euryzygoma dunense each of which appear to have occupied distinct dietary niches. This study suggests that southeastern Queensland hosted a mosaic of tropical forests, wetlands and grasslands during the Pliocene and was much less arid than previously thought.