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Synopsis Among extant great apes, orangutans are considered the most sexually dimorphic in body size. However, the expression of sexual dimorphism in orangutans is more complex than simply males being larger than females. At sexual maturity, some male orangutans develop cheek pads (flanges), while other males remain unflanged even after becoming reproductively capable. Sometimes flange development is delayed in otherwise sexually mature males for a few years. In other cases, flange development is delayed for many years or decades, with some males even spending their entire lifespan as unflanged adults. Thus, unflanged males of various chronological ages can be mistakenly identified as “subadults.” Unflanged adult males are typically described as “female-sized,” but this may simply reflect the fact that unflanged male body size has only ever been measured in peri-pubescent individuals. In this study, we measured the skeletons of 111 wild adult orangutans (Pongo spp.), including 20 unflanged males, 45 flanged males, and 46 females, resulting in the largest skeletal sample of unflanged males yet studied. We assessed long bone lengths (as a proxy for stature) for all 111 individuals and recorded weights-at-death, femoral head diameters, bi-iliac breadths, and long bone cross-sectional areas (CSA) (as proxies for mass) for 27 of these individuals, including seven flanged males, three adult confirmed-unflanged males, and three young adult likely-unflanged males. ANOVA and Kruskal–Wallis tests with Tukey and Dunn post-hoc pairwise comparisons, respectively, showed that body sizes for young adult unflanged males are similar to those of the adult females in the sample (all P ≥ 0.09 except bi-iliac breadth), whereas body sizes for adult unflanged males ranged between those of adult flanged males and adult females for several measurements (all P

At Liang Bua, the type site of Homo floresiensis on the Indonesian island of Flores, the stone artefact assemblages are dominated by two raw materials, qualitatively classified as chert and silicified tuff in previous studies. Field observations describe both stone types as locally abundant and of good flaking quality, but no systematic analysis has yet been carried out to characterise their nature. In this study, we conducted the first geological, mechanical, and quantitative assessment of these two raw materials using a suite of analytical approaches. Our results show that the two stone types are mineralogically alike in composition and derive from fossiliferous limestone that had undergone diagenetic silica replacement, but they clearly differ from one another geochemically. Therefore, the ‘chert’ and ‘silicified tuff’ categories used in previous studies are more aptly described as silica-dominated (i.e., SiO2-dominated) nodular chert and iron-rich (i.e., Fe2O3-rich) nodular chert, respectively. We discuss the implications of our results on the shift in raw material utilisation patterns at Liang Bua that occurred after ~ 46 ka and coincided with the arrival of Homo sapiens at the site.

Liang Bua (Flores, Indonesia) has yielded remains of a faunal community that included small-bodied and small-brained hominins, dwarf proboscideans, Komodo dragons, vultures and giant marabou storks (

Mountain gorillas are particularly inbred compared to other gorillas and even the most inbred human populations. As mountain gorilla skeletal material accumulated during the 1970s, researchers noted their pronounced facial asymmetry and hypothesized that it reflects a population-wide chewing side preference. However, asymmetry has also been linked to environmental and genetic stress in experimental models. Here, we examine facial asymmetry in 114 crania from three

Mountain gorillas are particularly inbred compared to other gorillas and even the most inbred human populations. As mountain gorilla skeletal material accumulated during the 1970s, researchers noted their pronounced facial asymmetry and hypothesized that it reflects a population-wide chewing side preference. However, asymmetry has also been linked to environmental and genetic stress in experimental models. Here, we examine facial asymmetry in 114 crania from three Gorilla subspecies using 3D geometric morphometrics. We measure fluctuating asymmetry (FA), defined as random deviations from perfect symmetry, and population-specific patterns of directional asymmetry (DA). Mountain gorillas, with a current population size of about 1000 individuals, have the highest degree of facial FA (explaining 17% of total facial shape variation), followed by Grauer gorillas (9%) and western lowland gorillas (6%), despite the latter experiencing the greatest ecological and dietary variability. DA, while significant in all three taxa, explains relatively less shape variation than FA does. Facial asymmetry correlates neither with tooth wear asymmetry nor increases with age in a mountain gorilla subsample, undermining the hypothesis that facial asymmetry is driven by chewing side preference. An examination of temporal trends shows that stress-induced developmental instability has increased over the last 100 years in these endangered apes. European Union [798117]; Ohio State University President's Postdoctoral Scholars Program, National Geographic Society [8486-08]; National Science Foundation [BCS 0852866, 0964944, 1520221, 1753651]; Leakey Foundation; Smithsonian Collections Care and Preservation Fund; 2010 Max Planck Research Award; French Government; SYNTHESYS Project - European Community Research Infrastructure Action under the FP7 'Capacities' Program Published version This project has received funding from the European Union's Horizon 2020 research and innovation program under the Marie Sklodowska-Curie grant agreement no. 798117, The Ohio State University President's Postdoctoral Scholars Program, National Geographic Society (8486-08), National Science Foundation (BCS 0852866, 0964944, 1520221, 1753651), The Leakey Foundation, the Smithsonian Collections Care and Preservation Fund, and the 2010 Max Planck Research Award to Timothy G. Bromage endowed by the German Federal Ministry of Education and Research to the Max Planck Society and the Alexander von Humboldt Foundation. This study received support from the French Government in the framework of the Investments for the Future Programme IdEx Universite de Bordeaux/GPR Human Past, and from the SYNTHESYS Project http://www.synthesys.info, which is financed by European Community Research Infrastructure Action under the FP7 'Capacities' Program.

Domesticated chickens (Gallus gallus domesticus) are a dominant part of the global human diet. Although the early domestication history of this species remains disputed, Mainland Southeast Asia (MSEA) is assumed to have been the initial domestication center. The eastward spread of chickens into Island Southeast Asia (ISEA) and the Pacific is typically attributed to human-mediated dispersals. Chicken remains are relatively common at Pacific Neolithic sites but are extremely rare in the archaeological records of MSEA and ISEA. Therefore, the exact routes and timing of the human-mediated spread of chickens from their native range in MSEA into the Pacific remain questions of interest. Here, we present the earliest evidence of Gallus on the Indonesian island of Flores at Liang Bua. This site has yielded an extensive stratigraphic sequence that spans from ~190,000 calendar years (ka) ago until the present and includes dense accumulations of faunal remains. Twelve bones from the cave's Holocene deposits have been identified as Gallus. The oldest remains, a right and left coracoid, were each directly dated to ~2,250 calibrated radiocarbon years before present (ka cal. BP), whereas the youngest Gallus elements are ~0.3 ka old. Although wild Green Junglefowl (Gallus varius) and Red Junglefowl (Gallus gallus) are found on Flores today, the absence of either of these species in deposits at Liang Bua older than ~2.5 ka as well as the size and shape of the oldest coracoids suggests that these remains likely represent domesticated G. gallus. This is the first evidence for domesticated chickens in the Neolithic of Flores and the first directly dated Gallus remains in Wallacea. The absence of chickens in the fossil record of ISEA suggests that Red Junglefowl (and perhaps Green Junglefowl also) reached Wallacea via human-mediated dispersal(s) at least ~2.25 ka cal. BP. publishedVersion

Liang Bua (Flores, Indonesia) has yielded remains of a faunal community that included small-bodied and small-brained hominins, dwarf proboscideans, Komodo dragons, vultures and giant marabou storks ( Leptoptilos robustus ). Previous research suggested that L. robustus evolved from a smaller L eptoptilos dubius -like Middle Pleistocene ancestor and may have been flightless. However, analyses of this species' considerably expanded hypodigm ( n = 43, MNI = 5), which includes 21 newly discovered bones described here for the first time, reveals that the wing bones of L. robustus were well-developed and this species was almost certainly capable of active flight. Moreover, L. robustus bones are broadly similar to Leptoptilos falconeri remains from sites in Africa and Eurasia, and its overall size range is comparable to fossils attributed to L. falconeri and similar specimens, as well as those of Leptoptilos lüi (China) and Leptoptilos titan (Java). This suggests that a Pleistocene dispersal of L. falconeri into Island Southeast Asia may have given rise to populations of giant marabou storks in this region. As L. robustus and L. titan are the most recent known representatives of these once plentiful giant marabou storks, Island Southeast Asia likely acted as a refugium for the last surviving members of this lineage.

Organic biomarker and lithic use-wear analyses of archaeological implements manufactured and/or used by hominins in the past offers a means of assessing how prehistoric peoples utilised natural resources. Currently, most studies focus on one of these techniques, rather than using both in sequence. This study aims to assess the potential of combining both methods to analyse stone artefacts, using a set of 69 stones excavated from the cave site of Liang Bua (Flores, Indonesia). Prior to chemical analysis, an initial inspection of the artefacts revealed potential use-wear traces but no visible residues. Gas chromatography mass spectrometry (GC-MS) analysis, including the targeting of 86 lipids, terpenes, terpenoids, alkanes and their analogues, found compounds with plant or animal origin on 27 of the 69 stones. The artefacts were subsequently cleaned, and use-wear analysis identified traces of use on 43 artefacts. Use-wear analysis confirmed traces of use on 23 of the 27 artefacts with potential use-residues that were determined by GC-MS. The GC-MS results were broadly consistent with the functional classes identified in the later use-wear analysis. This inclusive approach for stone artefact analysis strengthens the identifications made through multiple lines of enquiry. There remain conflicts and uncertainties in specific cases, suggesting the need for further refinement and analyses of the relationships between use-wear and residues.

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

Altres ajuts: CERCA Programme/Generalitat de Catalunya Current approaches to quantify phalangeal curvature assume that the long axis of the bone's diaphysis approximates the shape of a portion of a circle (included angle method) or a parabola (second-degree polynomial method). Here we developed, tested, and employed an alternative geometric morphometrics-based (GM) approach to quantify diaphysis shape of proximal phalanges in humans, apes and monkeys with diverse locomotor behaviors. One hundred landmarks of the central longitudinal axis were extracted from 3D surface models and analyzed using 2DGM methods, including generalized Procrustes analyses. Principal components analyses were performed and PC1 scores (>80% of variation) represented the dorsopalmar shape of the bone's central longitudinal axis and separated taxa consistently and in accord with known locomotor behavioral profiles. The most suspensory taxa, including orangutans, hylobatids and spider monkeys, had significantly lower PC1 scores reflecting the greatest amounts of phalangeal curvature. In contrast, bipedal humans and the quadrupedal cercopithecoid monkeys sampled (baboons, proboscis monkeys) exhibited significantly higher PC1 scores reflecting flatter phalanges. African ape (gorillas, chimpanzees and bonobos) phalanges fell between these two extremes and were not significantly different from each other. PC1 scores were significantly correlated with both included angle and the a coefficient of a second-degree polynomial calculated from the same landmark dataset, but had a significantly higher correlation with included angles. Our alternative approach for quantifying diaphysis shape of proximal phalanges to investigate dorsopalmar curvature is replicable and does not assume a priori either a circle or parabola model of shape, making it an attractive alternative compared with existing methodologies.

Homo naledi fossils from the Rising Star cave system provide important insights into the diversity of hand morphology within the genus Homo. Notably, the pollical (thumb) metacarpal (Mc1) displays an unusual suite of characteristics including a median longitudinal crest, a narrow proximal base, and broad flaring intrinsic muscle flanges. The present study evaluates the affinities of H. naledi Mc1 morphology via 3D geometric morphometric analysis of shaft shape using a broader comparative sample (n = 337) of fossil hominins, recent humans, apes, and cercopithecoid monkeys than in prior work. Results confirm that the H. naledi Mc1 is distinctive from most other hominins in being narrow at the proximal end but surmounted by flaring muscle flanges distally. Only StW 418 (Australopithecus cf. africanus) is similar in these aspects of shape. The gracile proximal shaft is most similar to cercopithecoids, Pan, Pongo, Australopithecus afarensis, and Australopithecus sediba, suggesting that H. naledi retains the condition primitive for the genus Homo. In contrast, Neandertal Mc1s are characterized by wide proximal bases and shafts, pinched midshafts, and broad distal flanges, while those of recent humans generally have straight shafts, less robust muscle flanges, and wide proximal shafts/bases. Although uncertainties remain regarding character polarity, the morphology of the H. naledi thumb might be interpreted as a retained intermediate state in a transformation series between the overall gracility of the shaft and the robust shafts of later hominins. Such a model suggests that the addition of broad medial and lateral muscle flanges to a primitively slender shaft was the first modification in transforming the Mc1 into the overall more robust structure exhibited by other Homo taxa including Neandertals and recent Homo sapiens in whose shared lineage the bases and proximal shafts became expanded, possibly as an adaptation to the repeated recruitment of powerful intrinsic pollical muscles.

Wallace's Line demarcates a biogeographical boundary between the Indomalaya and Australasian ecoregions. Most placental mammalian genera, for example, occur to the west of this line, whereas most marsupial genera occur to the east. However, macaque monkeys are unusual because they naturally occur on both western and eastern sides. To further explore this anomalous distribution, we analyzed 222 mitochondrial genomes from ∼20 macaque species, including new genomes from 60 specimens. These comprise a population sampling of most Sulawesi macaques, Macaca fascicularis (long-tailed macaques) specimens that were collected by Alfred R. Wallace and specimens that were recovered during archaeological excavations at Liang Bua, a cave on the Indonesian island of Flores. In M. fascicularis, three mitochondrial lineages span the southernmost portion of Wallace's Line between Bali and Lombok, and divergences within these lineages are contemporaneous with, and possibly mediated by, past dispersals of modern human populations. Near the central portion of Wallace's Line between Borneo and Sulawesi, a more ancient dispersal of macaques from mainland Asia to Sulawesi preceded modern human colonization, which was followed by rapid dispersal of matrilines and was subsequently influenced by recent interspecies hybridization. In contrast to previous studies, we find no strong signal of recombination in most macaque mitochondrial genomes. These findings further characterize macaque evolution before and after modern human dispersal throughout Southeast Asia and point to possible effects on biodiversity of ancient human cultural diasporas.

This chapter describes 12 tarsal bones from Sterkfontein, and compares them to those of extant hominoids and other fossil hominins, adding new analyses for some of the bones which have been studied previously. In general, the Sterkfontein calcanei and tali more closely resemble those attributed to Australopithecus sediba from nearby Malapa than to Australopithecus afarensis tarsals from Hadar, Ethiopia. Overall, the bones are similar to those of humans, but African ape-like characters indicating selection on adaptive use of arboreal substrates are persistent throughout the Sterkfontein tarsals, although more prevalent on some bones than on others. Planes of the navicular and first metatarsal facets on the medial cuneiform are relatively parallel, suggesting the hallux was not notably divergent from the metatarsus. However, the anteroposteriorly short intermediate and lateral cuneiforms suggest that the anterior tarsal row had not yet elongated as in modern humans, adversely affecting the foot’s ability to serve as a rigid lever. A large groove for the flexor hallucis longus muscle and large navicular tubercle may be related to strong grasping and climbing. Overall, the Sterkfontein tarsal elements show mixture of modern human-like and African ape-like features.

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

One hundred dental casts of modern Pima Amerindian children, 50 male and 50 female, wereexamined for the presence and expression of thirteen deciduous nonmetric traits. The effects of sexual dimorphism,asymmetry, and inter-trait association on trait presence were examined to evaluate their utility in populationdistance studies. No statistically significant differences between the sexes were observed. The majority of examinedvariants displayed a strong trend toward bilateral expression and no statistically significant differences betweenantimeres occurred. These data support the hypothesis that strong genetic components coupled with negligibleenvironmental influences are involved in deciduous trait presence. Five statistically significant associations betweenvariants were detected. Four of these involved a combination of incisor and canine shoveling within and betweenjaws. This indicates that their combined use in biological distance studies violates the mathematical assumption ofindependence. The lack of significant sexual dimorphism and asymmetry in the deciduous discrete traits examinedherein supports their use in population distance analyses if precautions are taken to use non-associated traits.

A sensitive and selective gas chromatography-tandem mass spectrometry (GC–MS/MS) method for the detection and quantification of terpenes, monoterpenoids and alkanes was developed and validated, to complement an existing analytical workflow set up for organic biomarker analysis of stone artefacts. This method was applied to seven stone artefacts—six of which contained potential use-residues based on a previous study using non-volatile low molecular weight lipid biomarkers—recovered from Liang Bua, an archaeological cave site on the Indonesian island of Flores. Following localised solvent extractions of the artefacts, aliquots of the solvent extracts were directly assayed using the optimised method. Identification of an analyte was considered positive when three criteria were met: (1) the retention time was the same as observed for a reference standard; (2) the three selected multiple reaction monitoring transitions for a reference standard compound were observed for the analyte; and (3) the qualitative ions (relative to the quantitative ion) were present in the expected ratios consistent with the reference standard. Alkane chemical profiles indicated that the presence of plant residues cannot be excluded, but of particular interest was the detection of camphor on one of the artefacts. Camphor-containing plants are found throughout Asia, including Indonesia, and are known historically to have been used for medicinal and culinary purposes. The likelihood of resource processing was high for three of the artefacts, and medium for the remaining four artefacts, based on the specificity, quantity and combination of the analytes identified.

Excavations in southeast Asia have unearthed a previously unreported hominin species named Homo luzonensis. The discovery has implications for ideas about early hominin evolution and dispersal from Africa. The discovery of a species of hominin called Homo luzonensis.

Edge-glossed flakes, usually recognised as stone tools with macroscopically visible polish from use, have been documented in parts of Southeast Asia from contexts spanning the past 12,000 years and possibly more than 30,000 years. It has been proposed that the edges of these flakes were polished by cutting, splitting or otherwise processing siliceous and fibrous plants for matting, basketry and bindings, and possibly by the reaping of cereals such as rice. Previous studies of use-wear on edge-glossed flakes in Southeast Asia have suggested several tool functions, including the cutting, splitting and scraping of plants with few or no phytoliths (e.g., Pandanaceae) and highly siliceous plants, with high phytolith content, such as Arecaceae palms (e.g., rattan) and Poaceae grasses (e.g., rice, bamboo and cattail reeds). Our study of the use-wear on 16 edge-glossed flakes from layers spanning the past 18,000 years at Liang Bua, a limestone cave on the Indonesian island of Flores, Indonesia, indicates processing of various plant taxa, including Poaceae grasses, Arecaceae palm and wood. The edge-glossed flakes from Liang Bua have features of micropolish that indicate they were probably linked with fibrecraft. Eleven of the 16 Liang Bua artefacts with edge-gloss are between about 18,580 and 11,750 years old, from artefact assemblages associated with modern humans; they currently represent the earliest evidence of these behaviours on Flores. No edge-glossed flakes have yet been identified in the 190,000–50,000-year-old artefact assemblages associated with Homo floresiensis.

Analyses of ancient micro-residues and usewear preserved on stone artefacts can potentially provide detailed information about how prehistoric humans used the artefacts to process materials such as food, pigments and/or adhesives. However, ancient micro-residues are likely degraded, and there are multiple potential sources of contamination, such as contact with sediments, groundwater, recent handling, storage materials or laboratory conditions, any of which can inhibit reliable identification of micro-residues and other traces of prehistoric use. In this pilot study, five stone tools from the archaeological site of Liang Bua (Flores, Indonesia) were used to evaluate the viability of Raman spectroscopy to identity ancient micro-residues preserved on stone artefact surfaces that are due specifically to prehistoric use as opposed to some form of ancient or modern source of contamination. Inorganic and organic deposits that occur commonly in the cave environment, including iron oxide, manganese oxide and biofilms, were identified in both the sediment and on the artefacts. Protein and saturated fatty acid micro-residues were identified on edges of all artefacts and may partially originate from modern handling. Proteins, plant fibres and other micro-residues associated with calcium nitrate are possibly archaeologically significant. Detection of plant fibres and starch grains may indicate either modern contamination or prehistoric contact with plant material that was transferred incidentally or during tool manufacture and/or tool use. These results demonstrate the viability of Raman microscopy to screen, at an early stage of archaeological residue analysis, for modern contaminants and micro-residues related to tool manufacture and/or tool use. This approach serves as a base for planning strategies and analytical protocols for future work that targets larger samples of artefacts, integrates Raman microscopy with GC–MS/LC–MS and includes more comprehensive studies of usewear. Copyright © 2017 John Wiley & Sons, Ltd.

Liang Bua, a karstic cave located on the island of Flores in eastern Indonesia, is best known for yielding the holotype of the diminutive hominin Homo floresiensis from Late Pleistocene sediments. Modern human remains have also been recovered from the Holocene deposits, and abundant archaeological and faunal remains occur throughout the sequence. The cave, the catchment in which it is located and the gross aggradational phases of the sediment sequence have all been subject to a great deal of scientific scrutiny since the discovery of the holotype of H. floresiensis in 2003. A recent program of geoarchaeological research has extended analyses of the site’s deposits to the microstratigraphic (micromorphological) level. The stratigraphic sequence in the cave is well defined but complex, comprising interstratified sediments of diverse lithologies and polygenetic origins, including volcanic tephras, fine-grained colluvium, coarse autogenic limestone gravels, speleothems and anthropogenic sediments, such as combustion features. The sedimentological and chemical heterogeneity suggest that processes of site formation and diagenesis varied markedly through time, both laterally and vertically. We present initial results from samples collected in 2014 from an excavation area near the rear of the cave, which yielded radiocarbon ages from charcoal that fill an important temporal gap in the chrono-stratigraphic sequence of previously excavated areas of the site. The results indicate marked changes in site environment and hominin activity during the Late Pleistocene, relating primarily to the degree to which the cave was connected to the hydrogeological system and to the varying intensities of use of the cave by hominins. Importantly, we identify anthropogenic signs of fire-use at the site between 41 and 24 thousand years ago, most likely related to the presence of modern humans.

Source determination of use-related residues on prehistoric stone tools is especially challenging, due to issues related to preservation, contamination and the contribution of residues from multiple sources. To increase confidence in this process, an analytical workflow was developed to include: (1) a sampling strategy that retains spatial information of the recovered residues and enables monitoring of environmental contamination; and (2) a sensitive and selective gas chromatography-tandem mass spectrometry (GC-MS/MS) procedure to quantify non-volatile low molecular weight lipids on stone artefacts. This workflow was applied to 14 stone artefacts excavated from deposits at Liang Bua, a limestone cave on the Indonesian island of Flores. These artefacts range in age between ∼14 000 and 1000 years old, and were preliminarily classified as either potentially showing traces of use (n = 7) or not (n = 7) using low magnification microscopy. Residues were sampled by direct solvent extraction off the surface of the artefacts. The aliquots were spiked with internal standards and derivatised. The trimethylsilyl derivatives of 40 saturated fatty acids, sterols, di- and tri-terpenoids and their analogues were quantified using optimised multiple reaction monitoring (MRM) transitions. Six of the potentially used artefacts contained sterols, phytosterols and terpenoids, either individually or in combination, whereas none of these compounds was commonly found on the seven artefacts preliminarily classified as unused. This suggests that these six artefacts may have been used as implements to process resources, and provides scope for further investigation. This workflow can also be adapted for the analysis of other archaeological objects.

The primate foot functions as a grasping organ. As such, its bones, soft tissues, and joints evolved to maximize power and stability in a variety of grasping configurations. Humans are the obvious exception to this primate pattern, with feet that evolved to support the unique biomechanical demands of bipedal locomotion. Of key functional importance to bipedalism is the morphology of the joints at the forefoot, known as the metatarsophalangeal joints (MTPJs), but a comprehensive analysis of hominin MTPJ morphology is currently lacking. Here we present the results of a multivariate shape and Bayesian phylogenetic comparative analyses of metatarsals (MTs) from a broad selection of anthropoid primates (including fossil apes and stem catarrhines) and most of the early hominin pedal fossil record, including the oldest hominin for which good pedal remains exist, Ardipithecus ramidus. Results corroborate the importance of specific bony morphologies such as dorsal MT head expansion and “doming” to the evolution of terrestrial bipedalism in hominins. Further, our evolutionary models reveal that the MT1 of Ar. ramidus shifts away from the reconstructed optimum of our last common ancestor with apes, but not necessarily in the direction of modern humans. However, the lateral rays of Ar. ramidus are transformed in a more human-like direction, suggesting that they were the digits first recruited by hominins into the primary role of terrestrial propulsion. This pattern of evolutionary change is seen consistently throughout the evolution of the foot, highlighting the mosaic nature of pedal evolution and the emergence of a derived, modern hallux relatively late in human evolution.

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

The Rising Star cave system has produced abundant fossil hominin remains within the Dinaledi Chamber, representing a minimum of 15 individuals attributed to Homo naledi. Further exploration led to the discovery of hominin material, now comprising 131 hominin specimens, within a second chamber, the Lesedi Chamber. The Lesedi Chamber is far separated from the Dinaledi Chamber within the Rising Star cave system, and represents a second depositional context for hominin remains. In each of three collection areas within the Lesedi Chamber, diagnostic skeletal material allows a clear attribution to H. naledi. Both adult and immature material is present. The hominin remains represent at least three individuals based upon duplication of elements, but more individuals are likely present based upon the spatial context. The most significant specimen is the near-complete cranium of a large individual, designated LES1, with an endocranial volume of approximately 610 ml and associated postcranial remains. The Lesedi Chamber skeletal sample extends our knowledge of the morphology and variation of H. naledi, and evidence of H. naledi from both recovery localities shows a consistent pattern of differentiation from other hominin species. DOI: http://dx.doi.org/10.7554/eLife.24232.001, eLife digest Species of ancient humans and the extinct relatives of our ancestors are typically described from a limited number of fossils. However, this was not the case with Homo naledi. More than 1500 fossils representing at least 15 individuals of this species were unearthed from the Rising Star cave system in South Africa between 2013 and 2014. Found deep underground in the Dinaledi Chamber, the H. naledi fossils are the largest collection of a single species of an ancient human-relative discovered in Africa. After the discovery was reported, a number of questions still remained. These questions included: why were so many fossils from a single species found at the one site, and how did they come to rest so far into the cave system? Possible explanations such as H. naledi living in the cave or being washed in by a flood were considered but ruled out. Instead, the evidence was largely consistent with intact bodies being deliberately disposed of in the cave and then decomposing. Now, Hawks et al. – who include many of the researchers who were involved in the discovery of H. naledi – report that yet more H. naledi fossils have been unearthed from a second chamber in the Rising Star cave system, the Lesedi Chamber. The chamber is 30 meters below the surface and there is no direct route between it and the Dinaledi Chamber. Again, the evidence is most consistent with the bodies arriving intact into the chamber, and there were no signs that the remains had been exposed to the surface environment. Also like the Dinaledi Chamber, no remains of other ancient humans or their relatives were found in the Lesedi Chamber. In total, 133 fossils of H. naledi have been found in this second chamber representing at least three individuals: two adults and a juvenile. However, and as Hawks et al. point out, only a small volume of the chamber has been excavated so far, and so there are likely more fossils still to be found. The fossils in the Lesedi Chamber are similar to those found before but include intact examples of bones, like the collarbone, that were previously known only from fragments. Perhaps the most impressive among the new fossils is a relatively complete skull that is part of a partial skeleton. The skull could have housed a brain that was 9% larger than the maximum estimate calculated from the previous H. naledi fossils. Though these new fossils provide us with yet more information about H. naledi, some questions still remain unanswered – the material from the Lesedi Chamber is undated, for example. However, a related study by Dirks et al. does give an estimate for the age of the fossils from the Dinaledi Chamber, while Berger et al. provide an explanation for why this date might be much younger than was previously predicted. DOI: http://dx.doi.org/10.7554/eLife.24232.002

Western gorillas (Gorilla gorilla) are known to climb significantly more often than eastern gorillas (Gorilla beringei), a behavioral distinction attributable to major differences in their respective habitats (i.e., highland vs. lowland). Genetic evidence suggests that the lineages leading to these taxa began diverging from one another between approximately 1 and 3 million years ago. Thus, gorillas offer a special opportunity to examine the degree to which morphology of recently diverged taxa may be "fine-tuned" to differing ecological requirements. Using three-dimensional (3D) geometric morphometrics, we compared talar morphology in a sample of 87 specimens including western (lowland), mountain (highland), and grauer gorillas (lowland and highland populations). Talar shape was captured with a series of landmarks and semilandmarks superimposed by generalized Procrustes analysis. A between-group principal components analysis of overall talar shape separates gorillas by ecological habitat and by taxon. An analysis of only the trochlea and lateral malleolar facet identifies subtle variations in trochlear shape between western lowland and lowland grauer gorillas, potentially indicative of convergent evolution of arboreal adaptations in the talus. Lastly, talar shape scales differently with centroid size for highland and lowland gorillas, suggesting that ankle morphology may track body-size mediated variation in arboreal behaviors differently depending on ecological setting. Several of the observed shape differences are linked biomechanically to the facilitation of climbing in lowland gorillas and to stability and load-bearing on terrestrial substrates in the highland taxa, providing an important comparative model for studying morphological variation in groups known only from fossils (e.g., early hominins).

Three-dimensional geometric morphometrics (3DGM) is a powerful tool for capturing and visualizing the “pure” shape of complex structures. However, these shape differences are sometimes difficult to interpret from a functional viewpoint, unless specific approaches (mostly based on biomechanical modeling) are employed. Here, we use 3DGM to explore the complex shape variation of the hamate, the disto-ulnar wrist bone, in anthropoid primates. Major trends of shape variation are explored using principal components analysis along with analyses of shape and size covariation. We also evaluate the phylogenetic patterning of hamate shape by plotting an anthropoid phylogenetic tree onto the shape space (i.e., phylomorphospace) and test against complete absence of phylogenetic signal using posterior permutation. Finally, the covariation of hamate shape and locomotor categories is explored by means of 2-block partial least squares (PLS) using shape coordinates and a matrix of data on arboreal locomotor behavior. Our results show that 3DGM is a valuable and versatile tool for characterizing the shape of complex structures such as wrist bones in anthropoids. For the hamate, a significant phylogenetic pattern is found in both hamate shape and size, indicating that closely related taxa are typically the most similar in hamate form. Our allometric analyses show that major differences in hamate shape among taxa are not a direct con

Gorillas occupy a variety of habitats from the west coast to eastern central Africa. These habitats differ considerably in altitude, which has a pronounced effect on forest ecology. Although all gorillas are obligate terrestrial knuckle-walking quadrupeds, those that live in lowland habitats eat fruits and climb more often than do those living in highland habitats. Here we test the hypothesis that gorilla talus morphology falls along a morphocline that tracks locomotor function related to a more inverted or everted foot set. This proposed morphocline predicts that gorillas living in lowland habitats may have a talocrural joint configured to facilitate a more medially oriented foot during climbing, suggesting that they may be more adaptively committed to arboreality than gorillas living in highland habitats. To quantify the relative set of the foot in gorillas, we chose two three-dimensional measurements of the talocrural joint: mediolateral curvature of the trochlea and relative surface area of the lateral malleolus. Our results show that, in comparison to their eastern counterparts, western gorillas have talar features that reflect a more medially directed sole of the foot. This morphology likely facilitates foot placement in a wider range of positions and minimization of shearing stresses across the joint when the foot is loaded on more curved or vertically oriented substrates as occurs during climbing and other arboreal behaviors. In contrast, eastern gorilla talar morphology is consistent with habitual placement of the foot with the sole directed more inferiorly, suggesting more effective loading during plantigrade push-off on terrestrial substrates. Am J Phys Anthropol 153:526–541, 2014. © 2013 Wiley Periodicals, Inc.

Despite discoveries of relatively complete hands from two early hominin species (Ardipithecus ramidus and Australopithecus sediba) and partial hands from another (Australopithecus afarensis), fundamental questions remain about the evolution of human-like hand anatomy and function. These questions are driven by the paucity of hand fossils in the hominin fossil record between 800,000 and 1.8 My old, a time interval well documented for the emergence and subsequent proliferation of Acheulian technology (shaped bifacial stone tools). Modern and Middle to Late Pleistocene humans share a suite of derived features in the thumb, wrist, and radial carpometacarpal joints that is noticeably absent in early hominins. Here we show that one of the most distinctive features of this suite in the Middle Pleistocene to recent human hand, the third metacarpal styloid process, was present ∼1.42 Mya in an East African hominin from Kaitio, West Turkana, Kenya. This fossil thus provides the earliest unambiguous evidence for the evolution of a key shared derived characteristic of modern human and Neandertal hand morphology and suggests that the distinctive complex of radial carpometacarpal joint features in the human hand arose early in the evolution of the genus Homo and probably in Homo erectus sensu lato.

Liang Bua, a limestone cave on the Indonesian island of Flores, has a depositional sequence that spans the last 95,000 years and includes well-preserved faunal remains. Birds are well represented throughout the stratigraphic sequence at Liang Bua. Here, we present the results of the first comprehensive study of avian remains retrieved from Sector XI, a 2 m by 2 m archaeological excavation along the east wall of the cave. A total of 579 specimens were identified as avian, with 244 belonging to at least 26 non-passerine taxa in 13 families. The late Pleistocene assemblage (23 taxa) includes the first recorded occurrence of vultures in Wallacea, as well as kingfishers, snipes, plovers, parrots, pigeons, and swiftlets. Together, these taxa suggest that during this time the surrounding environment was floristically diverse and included several habitat types. Two of these taxa, the giant marabou Leptoptilos robustus and the vulture Trigonoceps sp., are extinct. Eight taxa were identified in the Holocen...

The carpals from the Homo floresiensis type specimen (LB1) lack features that compose the shared, derived complex of the radial side of the wrist in Neandertals and modern humans. This paper comprises a description and three-dimensional morphometric analysis of new carpals from at least one other individual at Liang Bua attributed to H. floresiensis: a right capitate and two hamates. The new capitate is smaller than that of LB1 but is nearly identical in morphology. As with capitates from extant apes, species of Australopithecus, and LB1, the newly described capitate displays a deeply-excavated nonarticular area along its radial aspect, a scaphoid facet that extends into a J-hook articulation on the neck, and a more radially-oriented second metacarpal facet; it also lacks an enlarged palmarly-positioned trapezoid facet. Because there is no accommodation for the derived, palmarly blocky trapezoid that characterizes Homo sapiens and Neandertals, this individual most likely had a plesiomorphically wedge-shaped trapezoid (like LB1). Morphometric analyses confirm the close similarity of the new capitate and that of LB1, and are consistent with previous findings of an overall primitive articular geometry. In general, hamate morphology is more conserved across hominins, and the H. floresiensis specimens fall at the far edge of the range of variation for H. sapiens in a number of metrics. However, the hamate of H. floresiensis is exceptionally small and exhibits a relatively long, stout hamulus lacking the oval-shaped cross-section characteristic of human and Neandertal hamuli (variably present in australopiths). Documentation of a second individual with primitive carpal anatomy from Liang Bua, along with further analysis of trapezoid scaling relative to the capitate in LB1, refutes claims that the wrist of the type specimen represents a modern human with pathology. In total, the carpal anatomy of H. floresiensis supports the hypothesis that the lineage leading to the evolution of this species originated prior to the cladogenetic event that gave rise to modern humans and Neandertals.

Understanding the life history correlates of ontogenetic differences in hominoid brain growth requires information from multiple species. At present, however, data on how brain size changes over the course of development are only available from chimpanzees and modern humans. In this study, we examined brain growth in wild Virunga mountain gorillas using data derived from necropsy reports (N = 34) and endocranial volume (EV) measurements (N = 86). The youngest individual in our sample was a 10-day-old neonatal male with a brain mass of 208 g, representing 42% of the adult male average. Our results demonstrate that Virunga mountain gorillas reach maximum adult-like brain mass by 3-4 years of age; adult-sized EV is reached by the time the first permanent molars emerge. This is in contrast to the pattern observed in chimpanzees, which despite their smaller absolute brain size, reportedly attain adult brain mass approximately 1 year later than Virunga mountain gorillas. Our findings demonstrate that brain growth is completed early in Virunga mountain gorillas compared to other great apes studied thus far, in a manner that appears to be linked with other life history characteristics of this population.

Purpose Trapezial-metacarpal (TM) joint surfaces appear to be shallower in Asian than in white postmortem specimens, and the frequency of TM osteoarthritis seems to be substantially lower in Asian TM joints. This study tested the hypothesis that there are significant differences among human populations in TM joint surface curvature and that populations of Asian descent have less curvature than those of recent European descent. Methods The sample included trapeziums and first metacarpals from skeletons of 80 individuals of recent European and Asian descent and from skeletons of 34 African and 9 Australian aboriginal individuals. We scanned the surfaces using a laser digitizer to generate 3-dimensional models of each articular surface. We calculated dorsovolar, radioulnar, and root mean square curvatures by fitting modeled quadric surfaces to the TM joint surfaces. We tested pairwise comparisons of mean curvatures between populations for statistical significance using a standard resampling method (ie, bootstrapping). We also made pairwise comparisons of mean curvatures between males and females for a combined African and European sample. Results Mean dorsovolar metacarpal curvature was significantly higher in the European sample than in the Asian, African, and Australian samples. Mean root mean square curvature of the trapezial surface was significantly higher in the European sample than in the Asian sample. The European sample had the highest root mean square and dorsovolar trapezial curvatures of all the populations. We found no significant differences between male and female specimens. Conclusions A tendency toward higher mean dorsovolar curvature of both the metacarpal and trapezial surface in the European sample may help to explain the higher frequency of TM osteoarthritis reported in Europeans. Clinical relevance The greater TM curvatures affect basal thumb joint mechanics in thumb opposition and therefore may be a factor in the development of osteoarthritis at this joint in Europeans.

Molecular evidence indicates that the last common ancestor of the genus Pan and the hominin clade existed between 8 and 4 million years ago (Ma). The current fossil record indicates the Pan-Homo last common ancestor existed at least 5 Ma and most likely between 6 and 7 Ma. Together, the molecular and fossil evidence has important consequences for interpreting the evolutionary history of the hand within the tribe Hominini (hominins). Firstly, parsimony supports the hypothesis that the hand of the last common ancestor most likely resembled that of an extant great ape overall (Pan, Gorilla, and Pongo), and that of an African ape in particular. Second, it provides a context for interpreting the derived changes to the hand that have evolved in various hominins. For example, the Australopithecus afarensis hand is likely derived in comparison with that of the Pan–Homo last common ancestor in having shorter fingers relative to thumb length and more proximo-distally oriented joints between its capitate, second metacarpal, and trapezium. This evidence suggests that these derived features evolved prior to the intensification of stone tool-related hominin behaviors beginning around 2.5 Ma. However, a majority of primitive features most likely present in the Pan-Homo last common ancestor are retained in the hands of Australopithecus, Paranthropus/early Homo, and Homo floresiensis. This evidence suggests that further derived changes to the hands of other hominins such as modern humans and Neandertals did not evolve until after 2.5 Ma and possibly even later than 1.5 Ma, which is currently the earliest evidence of Acheulian technology. The derived hands of modern humans and Neandertals may indicate a morphological commitment to tool-related manipulative behaviors beyond that observed in other hominins, including those (e.g. H. floresiensis) which may be descended from earlier tool-making species.

Studies of infant feeding and weaning patterns in past populations that rely on a cross-sectional approach must make the assumption that no infant mortality bias exists. Previous investigations of infant weaning patterns at the Dakhleh Oasis, Egypt, relied on cross-sectional isotope data. In this study, we re-examine this weaning pattern, using a simulated longitudinal approach, which does not require any assumptions regarding potential infant mortality biases. This involves examining the dental isotopic signatures of individuals who survived the weaning process. Stable isotope signatures from juveniles and adults (102 individuals, 297 teeth) were examined to reconstruct the weaning history of those that survived the weaning process. Both deciduous and permanent teeth were sampled. Homogenized enamel and dentin samples were isolated from each tooth and analyzed for delta(13)C(ap) and delta(18)O(ap) from the enamel and delta(15)N(coll) and delta(13)C(coll) from dentin collagen. We investigate differences between in utero versus postbirth, preweaning versus postweaning, and juvenile versus adult stable isotope values as reflected in the dentition. A random permutation procedure was used to test for statistically significant differences in stable isotope values between tooth types. Statistically significant differences were observed in all stable isotopes between permanent and deciduous teeth, and between early and later forming permanent teeth in delta(13)C(ap) and delta(15)N(coll) isotopes. These results indicate dietary change between in utero and postbirth, and changes occurring during the weaning period. These results provide a more comprehensive picture of infant weaning practices at Kellis and provide further support that complete weaning occurred by 3 years of age.

(2007). The comparative method and the inference of venom-delivery systems in fossil mammals. Journal of Vertebrate Paleontology: Vol. 27, No. 2, pp. 541-546.

Homo rudolfensis Samples from Olduvai Gorge, East Lake Turkana, and Lake Malawi were included in this study. The East Lake Turkana fossils available prior to 2010 were examined first-hand, while for the Olduvai and Lake Malawi fossils and KNM-ER 60000, 62000, and 62003 we relied on original observations on fossils and casts as well as published reports (Schrenk et al., 1993; Blumenschine et al., 2003; Leakey et al., 2012). We include the following fossils in the hypodigm of H. rudolfensis: KNM-ER 819, KNM-ER 1470, KNM-ER 1482, KNM-ER 1483, KNM-ER 1590, KNM-ER 1801, KNM-ER 1802, KNM-ER 3732, KNM-ER 3891, KNM-ER 60000, KNM-ER 62000, KNM-ER 62003, OH 65, and UR 501. We do recognize that KNM-ER 60000 and KNM-ER 1802 present some conflicting anatomy that some authors have argued precludes them as conspecific specimens (Leakey et al., 2012); by considering both, we aim to be conservative as they encompass more variation within H. rudolfensis., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Australopithecus africanus The samples attributed to Au. africanus from Taung, Sterkfontein and Makapansgat were employed. Original specimens were examined first-hand by the authors., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Paranthropus boisei Samples from the Omo Shungura sequence, East Lake Turkana, Olduvai Gorge and Konso were included in this study. Original specimens from Olduvai Gorge and East Lake Turkana were examined first-hand, while casts and published reports (Tobias, 1967; Suwa et al., 1996, 1997; Do ḿınguez- Rodrigo et al., 2013) were used to study the Omo and Konso materials. Our postcranial considerations of P. boisei are very limited and we did not rely upon the association of KNM-ER 1500 (Grausz et al., 1988) to derive information about the postcranial skeleton of P. boisei. Paranthropus robustus The samples from Kromdraai, Swartkrans, Sterkfontein, Drimolen, Gondolin, and Coopers were included in this study. First-hand observations of original specimens from all localities were used with the exception of Drimolen fossils, which were compared using published reports (Keyser, 2000; Keyser et al., 2000)., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560

Paranthropus aethiopicus The cranium KNM-WT 17000 was examined first-hand for this study., Published as part of Lee R Berger, John Hawks, Darryl J de Ruiter, Steven E Churchill, Peter Schmid, Lucas K Delezene, Tracy L Kivell, Heather M Garvin, Scott A Williams, Jeremy M DeSilva, Matthew M Skinner, Charles M Musiba, Noel Cameron, Trenton W Holliday, William Harcourt-Smith, Rebecca R Ackermann, Markus Bastir, Barry Bogin, Debra Bolter, Juliet Brophy, Zachary D Cofran, Kimberly A Congdon, Andrew S Deane, Mana Dembo, Michelle Drapeau, Marina C Elliott, , Elen M Feuerriegel, Daniel Garcia-Martinez, David J Green, Alia Gurtov, Joel D Irish, Ashley Kruger, Myra F Laird, Damiano Marchi, Marc R Meyer, Shahed Nalla, Enquye W Negash, Caley M Orr, Davorka Radovcic, Lauren Schroeder, Jill E Scott, Zachary Throckmorton, Matthew W Tocheri, Caroline VanSickle, Christopher S Walker, Pianpian Wei & Bernhard Zipfel, 2015, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa, pp. 1-35 in eLife e 09560 4 on page 26, DOI: 10.7554/eLife.09560