Fossil Hominoids Research Articles

Paleoecology of Pleistocene mammals and paleoclimatic change in South China: Evidence from stable carbon and oxygen isotopes

The role of climate change in the evolution and diversification of hominoids remains a hotly debated issue. Stable isotope analyses of fossil mammals that coexisted with the hominoids can provide insights into hominoid palaeoenvironments and shed light on this debate. Here, we report results of stable carbon and oxygen isotope analyses of tooth enamel samples from a variety of Pleistocene mammals including pandas, deer, elephants, pigs, rhinos, and bovids from two hominoid fossil localities (Yugong Cave and Baxian Cave) in South China. Enamel δ13C values indicate that most of the mammals living in the study area during the late Middle Pleistocene had C3-based diets but a small number of individuals consumed some C4 grasses. This indicates the presence of C4 plants in the region during the late Middle Pleistocene, most likely in patches of open areas in a predominantly forested environment. However, during the early Late Pleistocene, all of the mammals examined had C3-based diets, except one bovid and one panda that may have ingested small amounts of C4 plants. This indicates a dense forested environment with little C4 grasses during the early Late Pleistocene. Like the Early Pleistocene pygmy panda (Ailuropoda microta) from Yanliang Cave, the late Middle Pleistocene Ailuropoda baconi from Yugong Cave and the early Late Pleistocene Ailuropoda melanoleuca from Baxian Cave had higher mean diet-δ13C values than other co-occurring herbivores, indicating they preferred relatively open forest habitats and had more restricted diets compared to other mammals. The reconstructed mean paleo-meteoric water δ18Ow values are lower than the annual average δ18Ow value of modern precipitation in the region, suggesting that the climatic conditions during the times when these Pleistocene mammals were alive were colder and/or wetter than today. In addition, δ18O values of the obligate drinkers (pigs, rhinos, bovids) display an overall decreasing trend, accompanied by increased range of δ18O variations, from the Early Pleistocene to the early Late Pleistocene. This suggests that the regional climate became colder and/or wetter, with increased seasonality, from the Early Pleistocene to the early Late Pleistocene, likely related to intensified glaciation. The change in climate to colder conditions may be responsible for the extinction of the Gigantopithecus in this region.

A new method to quantify mandibular corpus shape in extant great apes and its potential application to the hominoid fossil record.

Mandibular corpus robusticity (corpus breadth/corpus height) is the most commonly utilized descriptor of the mandibular corpus in the great ape and hominin fossil records. As a consequence of its contoured shape, linear metrics used to characterize mandibular robusticity are inadequate to quantify the shape of the mandibular corpus. Here, we present an alternative to the traditional assessment of mandibular shape by analyzing the outline of the mandibular corpus in cross-section using landmarks and semilandmarks. Outlines of the mandibular corpus in cross-section between M1 and M2 were quantified in a sample of hominoids and analyzed using generalized Procrustes analysis, Procrustes ANOVA, CVA, and cluster analysis. Corpus breadth and width were also collected from the same sample and analyzed using regression, ANOVA, and cluster analysis. Analysis of corpus outline shape revealed significant differences in mandibular corpus shape that are independent of size and sex at the genus level across hominoids. Cluster analysis based on the analysis of corpus outline shape results in almost all specimens grouping based on taxonomic affinity (99.28% correct classification). Comparison of these results to results using traditional measures of mandibular robusticity shows that analysis of the outline of the corpus in cross-section discriminate extant great apes more reliably. The strong taxonomic signal revealed by this analysis indicates that quantification of the outline of the mandibular corpus more fully captures mandibular corpus shape and offers the potential for greater power in discriminating among taxa in the hominoid fossil record.

Using machine learning to classify extant apes and interpret the dental morphology of the chimpanzee-human last common ancestor

Author(s): Monson, Tesla A.; Armitage, David W.; Hlusko, Leslea J. | Abstract: Machine learning is a formidable tool for pattern recognition in large datasets. We developed and expanded on these methods, applying machine learning pattern recognition to a problem in paleoanthropology and evolution. For decades, paleontologists have used the chimpanzee as a model for the chimpanzee-human last common ancestor (LCA) because they are our closest living primate relative. Using a large sample of extant and extinct primates, we tested the hypothesis that machine learning methods can accurately classify extant apes based on dental data. We then used this classification tool to observe the affinities between extant apes and Miocene hominoids. We assessed the discrimination accuracy of supervised learning algorithms when tasked with the classification of extant apes (n=175), using three types of data from the postcanine dentition: linear, 2-dimensional, and the morphological output of two genetic patterning mechanisms that are independent of body size: molar module component (MMC) and premolar-molar module (PMM) ratios. We next used the trained algorithms to classify a sample of fossil hominoids (n=95), treated as unknowns. Machine learning classifies extant apes with greater than 92% accuracy with linear and 2-dimensional dental measurements, and greater than 60% accuracy with the MMC and PMM ratios. Miocene hominoids are morphologically most similar in dental size and shape to extant chimpanzees. However, relative dental proportions of Miocene hominoids are more similar to extant gorillas and follow a strong trajectory through evolutionary time. Machine learning is a powerful tool that can discriminate between the dentitions of extant apes with high accuracy and quantitatively compare fossil and extant morphology. Beyond detailing applications of machine learning to vertebrate paleontology, our study highlights the impact of phenotypes of interest and the importance of comparative samples in paleontological studies.

Can Pallars i Llobateres: A new hominoid-bearing locality from the late Miocene of the Vallès-Penedès Basin (NE Iberian Peninsula)

In the Iberian Peninsula, Miocene apes (Hominoidea) are generally rare and mostly restricted to the Vallès-Penedès Basin. Here we report a new hominoid maxillary fragment with M2 from this basin. It was surface-collected in March 2017 from the site of Can Pallars i Llobateres (CPL, Sant Quirze del Vallès), where fossil apes had not been previously recorded. The locality of provenance (CPL-M), which has delivered no further fossil remains, is located very close (ca. 50 m) to previously known CPL outcrops, and not very far (ca. 500 m in NW direction) from the classical hominoid-bearing locality of Can Poncic 1. Here we describe the new fossil and, based on the size and proportions of the M2, justify its taxonomic attribution to Hispanopithecus cf. laietanus, a species previously recorded from several Vallesian sites of the Vallès-Penedès Basin. Based on the associated mammalian fauna from CPL, we also provide a biochronological dating and a paleoenvironmental reconstruction for the site. The associated fauna enables an unambiguous correlation to the Cricetulodon hartenbergeri – Progonomys hispanicus interval local subzone, with an estimated age of 9.98–9.73 Ma (late Vallesian, MN10). Therefore, CPL-M is roughly coeval with the Hispanopithecus laietanus-bearing localities of Can Llobateres 1 and Can Feu 1, and minimally older than those of La Tarumba 1 and Can Llobateres 2. In contrast, CPL-M is younger than the early Vallesian (MN9) localities of Can Poncic 1 (the type locality of Hispanopithecus crusafonti) as well as Polinyà 2 (Gabarró) and Estació Depuradora d’Aigües Residuals–Riu Ripoll 13, where Hispanopithecus sp. is recorded. The associated fauna from CPL indicates a densely forested and humid paleoenvironment with nearby freshwater. This supports the view that Hispanopithecus might have been restricted to dense wetland forests soon before its extinction during the late Vallesian, due to progressive climatic deterioration. Coupled with the existence of other fossiliferous outcrops in the area, this find is most promising for the prospect of discovering additional fossil hominoid remains in the future.

Trabecular and cortical bone structure of the talus and distal tibia in Pan and Homo.

Internal bone structure, both cortical and trabecular bone, remodels in response to loading and may provide important information regarding behavior. The foot is well suited to analysis of internal bone structure because it experiences the initial substrate reaction forces, due to its proximity to the substrate. Moreover, as humans and apes differ in loading of the foot, this region is relevant to questions concerning arboreal locomotion and bipedality in the hominoid fossil record. We apply a whole-bone/epiphysis approach to analyze trabecular and cortical bone in the distal tibia and talus of Pan troglodytes and Homo sapiens. We quantify bone volume fraction (BV/TV), degree of anisotropy (DA), trabecular thickness (Tb.Th), bone surface to volume ratio (BS/BV), and cortical thickness and investigate the distribution of BV/TV and cortical thickness throughout the bone/epiphysis. We find that Pan has a greater BV/TV, a lower BS/BV and thicker cortices than Homo in both the talus and distal tibia. The trabecular structure of the talus is more divergent than the tibia, having thicker, less uniformly aligned trabeculae in Pan compared to Homo. Differences in dorsiflexion at the talocrural joint and in degree of mobility at the talonavicular joint are reflected in the distribution of cortical and trabecular bone. Overall, quantified trabecular parameters represent overall differences in bone strength between the two species, however, DA may be directly related to joint loading. Cortical and trabecular bone distributions correlate with habitual joint positions adopted by each species, and thus have potential for interpreting joint position in fossil hominoids.

Lower Ilium Evolution in Apes and Hominins.

Elucidating the pelvic morphology of the Pan-Homo last common ancestor (LCA) is crucial for understanding ape and human evolution. The pelvis of Ardipithecus ramidus has been the basis of controversial interpretations of the LCA pelvis. In particular, it was proposed that the lower ilium became elongate independently in the orangutan and chimpanzee clades, making these taxa poor analogues for the pelvis of the LCA. This study examines the variation in relative lower ilium height between and within living and fossil hominoid species (and other anthropoids), and models its evolution using available fossil hominoids as calibration points. We find nuanced differences in relative lower ilium height among living hominoids, particularly in regards to gorillas, which do not have elongate lower ilia (because they are likely to represent the plesiomorphic hominoid condition for this trait). We also show that differences in relative lower ilium height among hominoid taxa are not readily explained by differences in size between species. Our maximum likelihood ancestral state reconstructions support inferences that chimpanzees (Pan troglodytes in particular) and orangutans evolved their elongate lower ilia independently. We also find that the predicted lower ilium height of the Pan-Homo LCA is shorter than all great apes except gorillas. This study adds to a growing body of evidence that finds different regions of the body show different evolutionary histories in different hominoids, and underscores that the unique combinations of morphologies of each modern and fossil hominoid species should be considered when reconstructing the mosaic nature of the Pan-Homo LCA. Anat Rec, 300:828-844, 2017. © 2017 Wiley Periodicals, Inc.

The late Miocene hominoid-bearing site in the Maragheh Formation, Northwest Iran

After a long period of inactivity, recent excavations at the late Miocene Maragheh Formation unexpectedly resulted in the discovery of the first fossil hominoid and second Mesopithecus remains from this area. The discovery motivated a new international initiative to conduct research in these rich fossil sites. These studies focused on the fossil hominoid and its locality, aiming to reveal more about the context of this fossil discovery. Detailed stratigraphy, sedimentology and magnetostratigraphy studies were conducted. New samples from volcaniclastic key horizons (pumice beds) in Dareh Gorg, where the hominoid fossil site is located, were dated by radiometric methods. The radiometric age determinations provide a firm tie-point for the geochronology. The polarity pattern in the palaeomagnetically investigated section corroborates the K-Ar results. The preliminary magnetostratigraphic results suggest that the hominoid locality can be correlated to the normal polarity chron C4n.2n (8.108–7.695 Ma), C4n.1n (7.642–7.528 Ma) or C3Br.1n (7.285–7.251 Ma), placing it at intervals corresponding to the mammal units MN11 or possibly early MN12. The study of fossil hominoid indicates broad affinities with a number of contemporaneous taxa from the Balkan-Iranian palaeoprovince, as well as Siwaliks and southeast Asia. A preliminary analysis of the accompanying (in situ) fauna at the hominoid site indicates the highest similarity of this level to Turolian hominoid- and Mesopithecus-bearing localities in Turkey, Greece and Bulgaria. However, some environmental differences are observed among these localities, based on their faunal structure and taxon properties, as well as in the different masticatory adaptations of their hominoids.

Rodent-based age appraisal of the Lower Siwalik Subgroup of Kalaunta, Ramnagar, Jammu, India

Parmar, V., Magotra, R., Norboo, R. & Prasad, G.V.R., July 2016. Rodent-based age appraisal of the Lower Siwalik Subgroup of Kalaunta, Ramnagar, Jammu, India. Alcheringa 41, xx-xx. ISSN 0311-5518.Rocks of the Lower Siwalik Subgroup exposed around Ramnagar town in Jammu, India, have been known for more than 90 years to yield Chinji-equivalent (14.2–11.2 Ma) hominoid fossils. In 1997, the fossil ape Sivapithecus reported near Kalaunta village in the vicinity of Ramnagar was argued to be about 18 Ma old by its discoverers. This had implications for hominoid evolutionary studies, as molecular clock calibrations are commonly based on Sivapithecus first appearance datum (FAD). In a paper published in 2010, the identification of this hominoid tooth was questioned and reassigned to a suid. Recently, based upon the discovery of an indeterminate fossil bovid from the supposedly hominoid-yielding site near Kalaunta, an age estimate between ca 13.9 and 10.8 Ma was provided. As rodent biochronology has proven to be useful in age assignment of upper Cenozoic terrestrial sequences, the age of the fossil-bearing Lower Siwalik Subgroup of Kalaunta is re-assessed based upon newly recovered cricetid taxa. The cricetid rodents from this site comprise Punjabemys downsi and P. mikros. These two species are known to exist in the well-dated Potwar Siwalik Group succession between 14.3 and 12.6 Ma and between 16.3 and 13.0 Ma, respectively. Based on the overlapping temporal ranges of the cricetid taxa and an indeterminate Bovidae (ca 13.9–10.8 Ma) reported previously from this area, an age of 13.9–13.0 Ma is inferred for rocks occurring in the vicinity of Kalaunta. The new finds refute earlier claims of rocks in the area extending back to ca 18 Ma or as young as 10.8 Ma.Varun Parmar [varunparmarvp@gmail.com], Rahul Magotra [rahulmagotra01@gmail.com] and Rigzin Norboo [norboo04@gmail.com], Post Graduate Department of Geology, University of Jammu, Jammu, 180 006, India; Guntupalli Veera Raghavendra Prasad [guntupalli.vrprasad@gmail.com], Department of Geology, Centre for Advanced Studies, University of Delhi, Delhi, 110 007, India.

The first hominoid from the Maragheh Formation, Iran

Miocene hominoid fossils are known from Africa and Eurasia, in the latter ranging widely from western Europe to Anatolia and from South Asia to Southeast/East Asia. Iran is located between the known western and eastern Eurasian hominoid distributions and is potentially important in understanding Miocene hominoid dispersal patterns. Maragheh is a late Miocene fossil locality in northwestern Iran, well known since the nineteenth century for its abundant mammalian fossils. However, until now, the only primate fossils reported from Maragheh or Iran were the Old World monkey Mesopithecus pentelicus. Recent field research at Maragheh has changed this situation by the discovery of the first hominoid fossil from Iran, a maxillary fragment with well-preserved second and third molars. Here, we provide a detailed description of this new specimen, comparing it with other similarly large-sized Eurasian late Miocene hominoids, Ouranopithecus, Ankarapithecus, Sivapithecus, and Indopithecus. Molar morphology of the Maragheh hominoid is similar to that of these Eurasian Miocene genera, with only minor differences in morphology and wear pattern. Based on the presently available materials, we tentatively prefer the interpretation that the Maragheh hominoid may be related more closely to either Ankarapithecus or Sivapithecus rather than to Ouranopithecus, but the fragmentary nature of the fossil makes evaluations difficult. Future discoveries of this Iranian hominoid are needed to determine its phylogenetic position with more certainty.

The stable isotope ecology of Pan in Uganda and beyond.

Stable isotope analysis has long been used to study the dietary ecology of living and fossil primates, and there has been increasing interest in using stable isotopes to study primate habitat use and anthropogenic impacts on non-human primates. Here, we examine the stable carbon and nitrogen isotope compositions of chimpanzees (Pan troglodytes) from seven communities in Uganda across a continuum of habitat structure (closed to more open) and access to anthropogenic resources (no reliance to heavy reliance). In general, the hair δ(13) C, but not δ(15) N, values of these communities vary depending on forest structure and degree of anthropogenic influence. When integrated with previously published hair δ(13) C and δ(15) N values for Pan, it is apparent that modern "savanna" and "forest" Pan form discrete clusters in carbon and nitrogen isotope space, although there are exceptions probably relating to microhabitat specialization. The combined dataset also reveals that Pan δ(13) C values (but not δ(15) N values) are inversely related to rainfall (r(2) = 0.62). We converted Pan hair δ(13) C values to enamel equivalents and made comparisons to the fossil hominoids Sivapithecus sp., Gigantopithecus blacki, Ardipithecus ramidus, and Australopithecus anamensis. The δ(13) C values of the fossil hominins Ar. ramidus and Au. anamensis do not cluster with the δ(13) C values of modern Pan in "forest" habitats, or with fossil hominoids that are believed to have inhabited forests. Am. J. Primatol. 78:1070-1085, 2016. © 2016 Wiley Periodicals, Inc.

Dental development in living and fossil orangutans

Numerous studies have investigated molar development in extant and fossil hominoids, yet relatively little is known about orangutans, the only great ape with an extensive fossil record. This study characterizes aspects of dental development, including cuspal enamel daily secretion rate, long-period line periodicities, cusp-specific molar crown formation times and extension rates, and initiation and completion ages in living and fossil orangutan postcanine teeth. Daily secretion rate and periodicities in living orangutans are similar to previous reports, while crown formation times often exceed published values, although direct comparisons are limited. One wild Bornean individual died at 4.5 years of age with fully erupted first molars (M1s), while a captive individual and a wild Sumatran individual likely erupted their M1s around five or six years of age. These data underscore the need for additional samples of orangutans of known sex, species, and developmental environment to explore potential sources of variation in molar emergence and their relationship to life history variables. Fossil orangutans possess larger crowns than living orangutans, show similarities in periodicities, and have faster daily secretion rate, longer crown formation times, and slower extension rates. Molar crown formation times exceed reported values for other fossil apes, including Gigantopithecus blacki. When compared to African apes, both living and fossil orangutans show greater cuspal enamel thickness values and periodicities, resulting in longer crown formation times and slower extension rates. Several of these variables are similar to modern humans, representing examples of convergent evolution. Molar crown formation does not appear to be equivalent among extant great apes or consistent within living and fossil members of Pongo or Homo.

Isotopic reconstructions of habitat change surrounding the extinction of Oreopithecus, the last European ape.

Oreopithecus bambolii was the last hominoid to survive in Europe. The purpose of this investigation was to reconstruct, through stable isotope analyses, Oreopithecus' habitat, subsistence behavior, and changes in habitat that may have led to its extinction. Carbon and oxygen stable isotopes from inorganic carbonate in tooth enamel from Oreopithecus and its contemporaneous faunas from localities in Tuscany and Sardinia were sampled. Also the fauna from localities in Tuscany shortly after Oreopithecus went extinct were sampled. Results indicated that Oreopithecus, compared with most modern hominoids, inhabited forests that probably had a more open canopy. At Tuscan localities, Oreopithecus yields some of the highest carbon isotope values but some of the lowest oxygen, suggesting a diet that may have included tubers or aquatic vegetation. Relatively higher oxygen values in Sardinia suggested that its diet included arboreal foods as well. Among modern and fossil hominoids, Oreopithecus only resembled chimpanzees living outside of rainforests. It also resembled Ardipithecus in carbon isotope values, suggesting possible similarities in feeding strategies concordant with shared skeletal features between Oreopithecus and early hominins. Isotope values from post-Oreopithecus faunas indicated a shift to more forested conditions, unlike other hominoid extinctions associated with loss of forest. Isotopic reconstructions of Oreopithecus' habitat and changes associated with its extinction indicated that its paleoecology was unique among hominoids. However, these reconstructions also suggested that like other hominoids, Oreopithecus was susceptible to changes in seasonality of precipitation, and it may have used wetlands as a buffer to seasonal regimes. Am J Phys Anthropol 160:254-271, 2016. © 2016 Wiley Periodicals, Inc.

A review of trabecular bone functional adaptation: what have we learned from trabecular analyses in extant hominoids and what can we apply to fossils?

Many of the unresolved debates in palaeoanthropology regarding evolution of particular locomotor or manipulative behaviours are founded in differing opinions about the functional significance of the preserved external fossil morphology. However, the plasticity of internal bone morphology, and particularly trabecular bone, allowing it to respond to mechanical loading during life means that it can reveal greater insight into how a bone or joint was used during an individual's lifetime. Analyses of trabecular bone have been commonplace for several decades in a human clinical context. In contrast, the study of trabecular bone as a method for reconstructing joint position, joint loading and ultimately behaviour in extant and fossil non-human primates is comparatively new. Since the initial 2D studies in the late 1970s and 3D analyses in the 1990 s, the utility of trabecular bone to reconstruct behaviour in primates has grown to incorporate experimental studies, expanded taxonomic samples and skeletal elements, and improved methodologies. However, this work, in conjunction with research on humans and non-primate mammals, has also revealed the substantial complexity inherent in making functional inferences from variation in trabecular architecture. This review addresses the current understanding of trabecular bone functional adaptation, how it has been applied to hominoids, as well as other primates and, ultimately, how this can be used to better interpret fossil hominoid and hominin morphology. Because the fossil record constrains us to interpreting function largely from bony morphology alone, and typically from isolated bones, analyses of trabecular structure, ideally in conjunction with that of cortical structure and external morphology, can offer the best resource for reconstructing behaviour in the past.

Clay mineralogy indicates a mildly warm and humid living environment for the Miocene hominoid from the Zhaotong Basin, Yunnan, China.

Global and regional environmental changes have influenced the evolutionary processes of hominoid primates, particularly during the Miocene. Recently, a new Lufengpithecus cf. lufengensis hominoid fossil with a late Miocene age of ~6.2 Ma was discovered in the Shuitangba (STB) section of the Zhaotong Basin in Yunnan on the southeast margin of the Tibetan Plateau. To understand the relationship between paleoclimate and hominoid evolution, we have studied sedimentary, clay mineralogy and geochemical proxies for the late Miocene STB section (~16 m thick; ca. 6.7–6.0 Ma). Our results show that Lufengpithecus cf. lufengensis lived in a mildly warm and humid climate in a lacustrine or swamp environment. Comparing mid to late Miocene records from hominoid sites in Yunnan, Siwalik in Pakistan, and tropical Africa we find that ecological shifts from forest to grassland in Siwalik are much later than in tropical Africa, consistent with the disappearance of hominoid fossils. However, no significant vegetation changes are found in Yunnan during the late Miocene, which we suggest is the result of uplift of the Tibetan plateau combined with the Asian monsoon geographically and climatically isolating these regions. The resultant warm and humid conditions in southeastern China offered an important refuge for Miocene hominoids.

Magnetic polarity of Masol 1 Locality deposits, Siwalik Frontal Range, northwestern India

The Mio-Pleistocene Siwalik formations have been known worldwide since the 19th century for their fossil hominoids. Numerous paleomagnetic studies have contributed to build the chronological framework of the Siwalik Group subdivided into Lower, Middle and Upper Siwalik Subgroups. Our study concerns the Tatrot Formation (Late Pliocene) of the Upper Siwalik Subgroup located at Masol in the Chandigarh Siwalik Frontal Range (India), and is accessible by the Patiali Rao River. At Masol (district Mohali, Punjab), the erosion of the anticline structure has formed an inlier and exposed paleontological assemblages characterizing the Late Pliocene “Quranwala fossiliferous zone”. Since 2008, the Indo-French research program, “Siwaliks”, has conducted surveys in the Masol inlier and has collected stone tools on the surface of the outcrops among fossilized bones, a few with cut marks. The first cut-marked bone was discovered in 2009 at Masol 1 (M1). The study of the magnetic polarities of some stratigraphic units of M1 revealed that the deposits recorded a normal polarity. According to the paleontology and the previous magnetostratigraphy of the Patiali Rao, it appeared that the deposits of Masol 1 are older than the Gauss-Matuyama reversal, dated to 2.58 Ma.

Morphology and environment in some fossil Hominoids and Pedetids (Mammalia).

Linking the environment to functional anatomy is not an easy exercise, especially when dealing with fossils, which are often fragmentary and represent animals that are extinct. A holistic approach permits us to fill the gaps in reconstructing the evolutionary patterns in fossil groups. Identifying the environment where animals lived can help to interpret some anatomical structures and, vice versa, the functional morphological pattern can help to refine some fossil environments. Two examples focusing on locomotor behaviours in fossil mammals are considered in this paper: the hominoids and the origins of hominid bipedalism and the springing adaptations in fossil rodents (Pedetidae) in connection with different habitats. In the first case, the limits of the chimp-based models and the necessity to take into account detailed environmental reconstructions will be addressed. The famous 'savannah hypothesis' is no longer tenable because the palaeontological data support a more vegetated environment for the origins of bipedal hominids. Data from the environment will be considered. The earliest putative hominid fossils which preserve skeletal remains of the locomotor apparatus show mixed adaptations to terrestrial bipedalism and arboreal activities. The second example focuses on the variation in springing adaptations in Pedetidae in the Lower Miocene of East Africa and Southern Africa. In the East, the sites where Pedetidae were preserved were mainly forested, whereas in the South the region was more open and drier, with extensive grassy patches. In the first case, pedetids were robust and heavy jumpers, whereas in the South they were smaller, their skeleton more gracile and their springing was lighter. During the desertification of the southern part of Africa, the large pedetid species became extinct, but a smaller species developed. In the case of primates, as in the case of rodents, the skeletal morphology was adapted to its environment.

Evolution of the hominoid vertebral column: The long and the short of it.

The postcranial axial skeleton exhibits considerable morphological and functional diversity among living primates. Particularly striking are the derived features in hominoids that distinguish them from most other primates and mammals. In contrast to the primitive catarrhine morphotype, which presumably possessed an external (protruding) tail and emphasized more pronograde trunk posture, all living hominoids are characterized by the absence of an external tail and adaptations to orthograde trunk posture. Moreover, modern humans evolved unique vertebral features that satisfy the demands of balancing an upright torso over the hind limbs during habitual terrestrial bipedalism. Our ability to identify the evolutionary timing and understand the functional and phylogenetic significance of these fundamental changes in postcranial axial skeletal anatomy in the hominoid fossil record is key to reconstructing ancestral hominoid patterns and retracing the evolutionary pathways that led to living apes and modern humans. Here, we provide an overview of what is known about evolution of the hominoid vertebral column, focusing on the currently available anatomical evidence of three major transitions: tail loss and adaptations to orthograde posture and bipedal locomotion.

Virtual anthropology.

Comparative morphology, dealing with the diversity of form and shape, and functional morphology, the study of the relationship between the structure and the function of an organism's parts, are both important subdisciplines in biological research. Virtual anthropology (VA) contributes to comparative morphology by taking advantage of technological innovations, and it also offers new opportunities for functional analyses. It exploits digital technologies and pools experts from different domains such as anthropology, primatology, medicine, paleontology, mathematics, statistics, computer science, and engineering. VA as a technical term was coined in the late 1990s from the perspective of anthropologists with the intent of being mostly applied to biological questions concerning recent and fossil hominoids. More generally, however, there are advanced methods to study shape and size or to manipulate data digitally suitable for application to all kinds of primates, mammals, other vertebrates, and invertebrates or to issues regarding plants, tools, or other objects. In this sense, we could also call the field "virtual morphology." The approach yields permanently available virtual copies of specimens and data that comprehensively quantify geometry, including previously neglected anatomical regions. It applies advanced statistical methods, supports the reconstruction of specimens based on reproducible manipulations, and promotes the acquisition of larger samples by data sharing via electronic archives. Finally, it can help identify new, hidden traits, which is particularly important in paleoanthropology, where the scarcity of material demands extracting information from fragmentary remains. This contribution presents a current view of the six main work steps of VA: digitize, expose, compare, reconstruct, materialize, and share. The VA machinery has also been successfully used in biomechanical studies which simulate the stress and strains appearing in structures. Although methodological issues remain to be solved before results from the two domains can be fully integrated, the various overlaps and cross-fertilizations suggest the widespread appearance of a "virtual functional morphology" in the near future.

New carnivore material from the Middle Miocene of Turkey. Implications on biochronology and palaeoecology

New carnivore remains, as well as some old material from the middle Miocene of Turkey are studied and reviewed in the present article. The middle Miocene mammal faunas of Southeastern Europe and Western Asia are rare and every new finding contributes to their better knowledge. The studied material originates from the Çandır, Paşalar and Inönü localities, which are well known, especially the two first ones because of the presence of hominoid fossil remains. The following species have been recognized in the studied material from each locality (those marked by an asterisk are found for the first time): Çandır: Amphicyon major, “Thalassictis” cf. montadai*, Percrocuta tungurensis*, Percrocuta sp.; Paşalar: Amphicyonidae sp., Ischyrictis sp., Protictitherium intermedium, Protictitherium cingulatum*, Protictitherium cf. crassum*, Styriofelis lorteti; Inönü: Sansanosmilus cf. palmidens*. The carnivoran faunas of the Çandır and Paşalar localities are similar, with eight shared species. They differ in the presence of ailurids in Çandır and the higher abundance of mustelids in Paşalar. The age of the three localities is discussed; the similarity of the Çandır and Paşalar faunas suggests that they have similar ages. The carnivore guild structure of Çandır and Paşalar and its comparison with modern assemblages from known environments suggest a mixed, open woodland-like environment.

ESR, U-series and paleomagnetic dating of Gigantopithecus fauna from Chuifeng Cave, Guangxi, southern China

Several Gigantopithecus faunas associated with taxonomically undetermined hominoid fossils and/or stone artifacts are known from southern China. These faunas are particularly important for the study of the evolution of humans and other mammals in Asia. However, the geochronology of the Gigantopithecus faunas remains uncertain. In order to solve this problem, a program of geochronological studies of Gigantopithecus faunas in Guangxi Province was recently initiated. Chuifeng Cave is the first studied site, which yielded 92 Gigantopithecus blacki teeth associated with numerous other mammalian fossils. We carried out combined ESR/U-series dating of fossil teeth and sediment paleomagnetic studies. Our ESR results suggest that the lower layers at this cave can be dated to 1.92±0.14Ma and the upper layers can be dated to older than 1.38±0.17Ma. Correlation of the recognized magnetozones to the geomagnetic polarity timescale was achieved by combining magnetostratigraphic, biostratigraphic and ESR data. The combined chronologies establish an Olduvai subchron (1.945–1.778Ma) for the lowermost Chuifeng Cave sediments. We also analyzed the enamel δ13C values of the Gigantopithecus faunas. Our results show that southern China was dominated by C3 plants during the early Pleistocene and that the Gigantopithecus faunas lived in a woodland-forest ecosystem.