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Objectives Gigantopithecus blacki, the largest hominoid known, is one of the representative Pleistocene mammals in southern China and northern Southeast Asia. Here we investigate the feeding ecology of G. blacki in its core habitat (Guangxi, Southern China) during the early Early Pleistocene, which was the early period in its evolution. Materials and methods The stable isotopic (C, O) analysis of tooth enamel of the fauna associated with G. blacki (n = 58), including the largest number of G. blacki teeth (n = 12) to date from the Liucheng Gigantopithecus Cave (~2 Ma), Guangxi, China, is undertaken. Results The δ13 C values of Liucheng fauna range from -12.9 to -19.0‰ with an average of -16.1 ± 1.3‰ (n = 58) and the δ18 O values range from -4.3 to -9.6‰ with an average of -6.9 ± 1.2‰ (n = 58). The δ13 C values of G. blacki range from -15.9‰ to -17.0‰ with an average of -16.5 ± 0.4‰ (n = 12), and the δ18 O values vary from -5.9‰ to -7.5‰ with an average of -6.6 ± 0.5‰ (n = 12). Conclusions The isotopic data show Guangxi was characterized by closed C3 forest and humid climate in the early Early Pleistocene. Niche partitioning is found among G. blacki, Sinomastodon, Ailuropoda and Stegodon, the typical megafauna in South China in the early Early Pleistocene. This could be one of the important factors for them to co-exist until the Middle Pleistocene. Smallest isotopic variations of G. blacki are found compared with those of contemporary animals, indicating a conservative foraging ecology i.e., limited foraging area and/or narrow dietary flexibility. Furthermore, the more confined foraging ecology of G. blacki is also seen in comparison with fossil and extant large-bodied primates. However, the unique dietary pattern of G. blacki does not seem to have hindered its survival. The environment in Guangxi during the early Early Pleistocene offered the suitable conditions for G. blacki to become one of the typical species in the faunal assemblages.

It has long been thought that environmental perturbations were the key driving force behind the succession of three distinct mammal faunas in the Pleistocene Ailuropoda-Stegodon faunal complex (sensu lato) of South China: the lower Pleistocene Gigantopithecus-Sinomastodon fauna, the middle Pleistocene Ailuropoda-Stegodon fauna (sensu stricto) and the upper Pleistocene Homo-Elephas fauna. Here, we apply three-dimensional dental microwear texture analysis (DMTA) to three characteristic fossil proboscideans from these mammal faunas to provide preliminary tests for hypotheses of trophic ecology. Despite a few methodological caveats, this study demonstrates the potential of DMTA for understanding the diets of fossil proboscideans. The texture of microwear in Sinomastodon and Stegodon are more reflective of browsing, whereas that of Elephas is suggestive of mixed feeding. The results suggest a more complex process of Pleistocene faunal turnovers in South China. Rather than a unidirectional trend of climate-driven environmental deterioration, biotic factors might have played a more substantial role than previously thought.

As one of the most important Quaternary mammalian faunas in southern China, the Gigantopithecus-Sinomastodon fauna has received much attention. The large-primate fossil teeth newly collected from Juyuan karst cave in Boyue Mountain, Guangxi Zhuang Autonomous Region of southern China, can be identified as Gigantopithecus blacki . The morphology and size of these G. blacki teeth differ from those of the middle Pleistocene Gigantopithecus teeth which hold evidently larger dental dimensions and more complex crenulations from Hejiang Cave also in Guangxi and Tham Khuyen Cave in Vietnam. However, the G. blacki teeth from Juyuan Cave are relatively similar to those from Early Pleistocene Mohui Cave and Liucheng Gigantopithecus Cave both in Guangxi, which suggests that the three cave sites have similar age. The Juyuan fauna associated with Gigantopithecus blacki , consisting of 45 mammalian species (such as Sinomastodon yangziensis , Ailuropoda wulingshanensis , Stegodon huananensis , and Rhinoceros fusuiensis ), is a typical Early Pleistocene Gigantopithecus-Sinomastodon fauna of southern China. The Juyuan fauna is mostly similar to Mohui fauna, also implying their contemporaneity. Paleomagnetic analyses demonstrate that the fossil-bearing sediments in Juyuan Cave are dominated by normal polarity. Combining the faunal analysis and magnetostratigraphic evidence, the Juyuan sediments can be best correlated with the Olduvai normal subchron, giving an estimated age of 1.8 Ma.

The origin of the Old World brevirostrine gomphotheriid taxon Sinomastodon has been debated intensively. The discovery of the oldest known Sinomastodon cranium, reported herein, supports its endemic origin and contradicts the prevalent theory of its North America origin. The new cranium was discovered from the Shuitangba locality, southwestern China, and is dated at about 6.5–6.0 Ma, corresponding to the latest Miocene. The new specimen shows distinct characters from the other species of Sinomastodon and was therefore named Sinomastodon praeintermedius, sp. nov. Newly discovered, isolated Sinomastodon-like teeth from the upper Miocene to the lower Pleistocene of southwestern China and Southeast Asia indicate a long evolution of Sinomastodon endemically. Remains of this species are frequently accompanied by those of stegodontid species. These two groups may have had a similar migration route, invading northern China and Japan during the latest Miocene, and retreating or becoming extinct from the Palearctic realm by the end of the Pliocene. The migrations of proboscideans may have been sparked by major paleoenviromental changes, i.e., the strengthened summer monsoon beginning in the late Miocene (~7–8 Ma) and global cooling due to the expansion of ice sheets from the middle Pliocene to the early Pleistocene. The new finding reveals a close relationship of the early Pliocene fauna of northern China and the latest Miocene fauna of southwestern China, and thus provides novel insight into the origin and components of Pliocene fauna in northern China.

Since its sub-aerial emergence, Java has experienced multiple tectonic, geographic and climatic changes, which affected the megafaunal occupation, adaptation and succession. Six Proboscidean taxa have been found from various localities throughout Java extending back to the Early to Late Pleistocene. The six taxa are: Stegoloxodon indonesicus, Sinomastodon bumiajuensis, pygmy Stegodon sp., Stegodon trigonocephalus, Elephas hysudrindicus and Elephas maximus, in which respective taxa are included in successive faunal stages. The aim of this research was to reconstruct the succession of Proboscidea in Java and the adaptation of each taxon to environmental changes by incorporating stable isotope analysis with the fossil faunal record, geology and stratigraphy. We conducted stable carbon (δ13C) and oxygen (δ18O) isotope analysis of the carbonate phase in the tooth enamel of six proboscidean taxa from numerous localities and ages. Our results suggest that for the earliest terrestrial fauna from Java, represented by Stegoloxodon indonesicus, the feeding ecology was restricted to a closed canopy rainforest during the earliest Pleistocene, while towards the late Early Pleistocene, the successive taxon, Sinomastodon bumiajuensis, was adapted to different or increasingly drier grassy habitats. C4 expansion on the island took place since the later part of the Early Pleistocene (before 1.5 Ma) and continued until the Middle Pleistocene, as suggested by the carbon and oxygen isotope composition of assemblages of proboscidean taxa from Sangiran, Kedung Brubus, Trinil, and western Java, which also predominantly display the expected range of C4-dominant feeders. However, the occurrence of mixed C3/C4 feeder Proboscidea in the Ngandong Fauna suggests that fragmented dense evergreen forests, shrubby or woodland vegetation reappeared towards the end of the Middle Pleistocene. This environmental shift is detected in the later stage of the Middle Pleistocene, as the dietary preference of all analysed samples from individuals from this age shifted back from a C4-dominated into a C3-dominated diet, which suggests a change from a dry to more humid climate conditions.

The Mohui fauna, associated with Gigantopithecus blacki, has been recovered from the Pleistocene karst cave deposit in Bubing Basin, Guangxi, South China. The large-mammalian assemblage derives from 28 species, 15 of which have no historic descendants. Occurrence of some ancient species implies that this fauna is early Early Pleistocene, such as Hystrix magna, Sinomastodon yangziensis, Stegodon huananensis, Ailuropoda microta, Pachycrocuta licenti, Tapirus sanyuanensis, Hespertherium sp., and Dorcabune liuchengense. Comparisons to already-dated early Pleistocene faunas in South China, combined with preliminary paleomagnetic, electron spin resonance (ESR) and U-series analysis of this cave, indicate mammalian fauna age around 1.7 Ma. Comparisons to five faunal assemblages in the same basin show species-level differences in Ailuropoda, Stegodon, Hystrix, Tapirus, and Sus, implying the large-mammal fauna has passed through a slow process of evolution during the Quaternary in East Asia.

Sinomastodon (Proboscidea, Mammalia) is a unique brevirostrine trilophodont gomphotheriid from Eurasia, which mainly occurs in China during the Plio-Pleistocene. Although much is known about adult individuals, the morphology of juveniles of this taxon is still poorly understood due to the scarcity of pertinent fossils. The present contribution represents the first detailed systematic study of the mandible and deciduous dentition of juvenile individuals of Sinomastodon jiangnanensis from the early Early Pleistocene (2.14–2.15 Ma) Renzidong Cave of eastern China. Based on morphological comparisons, the Renzidong juvenile mandible bears, on one hand, the transitional characteristics between the typical long-jawed trilophodont and elephantoid proboscideans, and, on the other hand, more primitive features than adult mandibles of Sinomastodon . This study of juvenile individuals of S. jiangnanensis provides new information to enhance the diagnosis of S. jiangnanensis , and confirms the validity of the species. It also provides evidence to show that the morphological transition of genus Sinomastodon from Neogene to Quaternary is mirrored in adult and juvenile individuals.

Since 2004, five new early Pleistocene cave sites with Gigantopithecus blacki and other fossil mammals have been discovered in the Chongzuo, Zuojiang River area, Guangxi Zhuang Autonomous Region (Guangxi ZAR), South China. These sites include Baikong Cave of Liyu Mountain, Juyuan Cave of Boyue Mountain, Sanhe and Queque Caves of Wuming Mountain, and Wuming Cave of Mulan Mountain. The present study provides a preliminary account of these newly discovered Gigantopithecus faunas, and discusses their taxonomic composition, evolutionary trends and chronology based on recent biostratigraphic and geochronological studies. Combining faunal and paleomagnetic data, the Gigantopithecus faunas from the Zuojiang River area can be divided into three temporal stages during the early Pleistocene. The early stage (2.6–1.8 Ma) is represented by the Baikong and Boyue faunas, which include some Neogene relics (e.g., Sinomastodon, Hesperotherium, Dicoryphochoerus, Dorcabune and Cervavitus) and several species that make their first appearance during the Pleistocene (e.g., Sinicuon dubius, Pachycrocuta licenti, Ailuropoda microta, Sinomastodon jiangnanensis, Stegodon huananensis and Tapirus sanyuanensis). The middle stage (1.8–1.2 Ma) is represented by the Sanhe fauna, which is characterized by the appearance of Ailuropoda wulingshanensis, Cuon antiquus, and Tapirus sinensis. The late stage (1.2–0.8 Ma) is represented by the Queque fauna, which is characterized by a marked decrease in Neogene relics and the appearance of a few species that are more typical of the middle Pleistocene (e.g., Ailuropoda baconi and Stegodon orientalis). Evolutionary trends in the early Pleistocene Gigantopithecus faunas include a gradual increase in dental size in some of the typical taxa, such as G. blacki, Ailuropoda and Tapirus, and the successive turnover of representative species, such as A. microta – A. wulingshanensis – A. baconi. Thus far, Eleven G. blacki sites of early Pleistocene age have been recorded in southern China, and these are primarily concentrated in Guangxi ZAR, where more than 70% of early Pleistocene G. blacki cave sites are known. Given this temporal and geographic distribution it is possible to infer that Guangxi ZAR represents an important early evolutionary and zoogeographic center for G. blacki.

As a valid genus belonging to the family Gomphotheriidae, Sinomastodon is the unique brevirostrine trilophodont gomphotheriid known from the Old World and was primarily indigenous to China from the Early Pliocene to the Pleistocene. The new bunodont molars from the Sanhe karst Cave in Chongzuo, Guangxi Zhuang Autonomous Region (ZAR), South China have been described as Sinomastodon yanziensis of the middle Early Pleistocene, with associated reliable paleomagnetic dating of 1.2–1.6 Ma. This represents the southernmost distribution of Sinomastodon remains found in China. The diagnosis of S. yanziensis is revised here, the evolution of Pleistocene Sinomastodon in China is divided into three temporal stages, and the evolutionary trends on Early Pleistocene Sinomastodon molars are also summarized. Sinomastodon was once prevalent during the Early Pleistocene in South China and possibly became extinct at the end of the Early Pleistocene. Consequently, the Early Pleistocene fauna from South China is suggested to be named the Gigantopithecus-Sinomastodon fauna, to distinguish it from the typical Middle Pleistocene Ailuropoda-Stegodon fauna (s. s.).

We report a fossil specimen referable to Sinomastodontinae gen. et sp. indet. from the Neogene strata at Yanghecun locality, Xihe County, Gansu Province, China. The specimen is characterized by a brevirostrine mandible, complete pretrite trefoils, and relatively simple posttrite half lophids, showing typical features of Sinomastodon. It differs from the other known species of Sinomastodon by the following features: relatively short and wide m3 due to fewer lophid numbers, less inflated pretrite accessory central conules, poorly developed secondary trefoils and cementum, and relatively strong cingulid. All of these features indicate a bias towards pleisiomorphies of Sinomastodon, implying that this specimen is more ancestral than any known species of Sinomastodon. The symphysis of the new specimen is relatively long, which differs from the typical brevirostrine Sinomastodon, and thus we consider it a gen. et sp. indet. in the Subfamily Sinomastodontinae. In addition, the horizon in which the present specimen was found probably represents the Upper Miocene because it is lower than Pliocene strata yielding Hipparion (Proboscidipparon) pater. Generally, Sinomastodon is considered to have migrated from North America at about the time of the Miocene/Pliocene boundary, and to have been derived from a certain clade of American gomphotheres. However, the discovery of the Yanghecun specimen verifies that Sinomastodon lived in East Asia during the Late Miocene, and probably derived from Old World gomphotheres (e.g., G. wimani). The similarity between the members of the Subfamilies Sinomastodontinae and those of Cuvieroniinae is suggested to have been the result of parallel evolution.

The South American gomphotheres were previously classified in three genera and three species. Two biogeographic corridors are recognized for the dispersion of gomphotheres in South America. Cuvieronius hyodon was found mainly in the highlands and west to the Andes, and Stegomastodon platensis and Haplomastodon chimborazi were found in lowlands. The aim of the present study is to review the taxonomy of South American lowland gomphotheres. A large sample of specimens was analyzed, which included crania, teeth and postcrania from several localities in Brazil, Ecuador, Colombia, Venezuela, Argentina, and Uruguay. A review of the diagnostic characters proposed in previous taxonomic studies of South American lowland gomphotheres found that they are variable in both reputed species if large samples are considered. There is no support for more than one South American lowland gomphothere species, and it shows a significant morphological variation in some characteristics. The name for this taxon, after a taxonomic review, must be Notiomastodon platensis (Ameghino, 1888) new combination. This reduces the number of gomphothere species in South America to two genera and species. The lowland gomphothere is closely related to Cuvieronius and Rhynchotherium, and not closely related to Stegomastodon from North America, and Sinomastodon from Asia.

The systematics of gomphotheres in South America is confused, and the status of some taxa is still subject to discussion. Recently, Prado and Alberdi published a cladistic analysis of the trilophodont gomphotheres, with emphasis on the South American species. Unfortunately, the paper has problems in many points, which invalidates the phylogeny itself and the biogeographic analysis of Alberdi et al. based on it. This paper reviews the problems of the analysis of Prado and Alberdi. The main problems of their analysis are: poor taxon sampling, exclusion of non-South American species of genera present in South America, inadequate choice of outgroup, use of outdated software and methodology, and several mistakes in coding of characters. Rebuilding of the original character matrix and analysis using TNT software resulted in a new consensus tree from five equally parsimonious trees. In the analysis, Rhynchotherium , Sinomastodon , Gnathabelodon , Eubelodon and a clade containing the three South American taxa form a polytomy that have Gomphotherium as a sister group. Sinomastodon and Gnathabelodon were successive branches toward the South American taxa in Prado and Alberdi’s result. The main point of this contribution is to correct the errors in the original analysis and character coding, including polymorphism when necessary, and to use updated software and weighting strategy. The result is much less resolved than that of Prado and Alberdi, but is has a strong influence on the conclusions that those authors drew from the original phylogeny and the biogeographic implication presented in Alberdi et al.

Here we report the first discovery of a relatively complete skull of Choerolophodon from the late early Miocene Dalanggou quarry of Linxia Basin, Gansu Province in China. The skull is low and elongated, with two enamel-less and outward-and-upward curved incisors and the choerolophodont molars. These characters indicate that the new specimen can be assigned to the genus Choerolophodon, as it is quite distinct from other common gomphotheres in northern China such as Gomphotherium, Platybelodon, and Sinomastodon. The skull is also primitive in possessing laterally-expanded zygomatic arches, anteriorly-positioned orbits, weakly-developed cheek teeth cement, and retention of a P4. Prior to this discovery, Choerolophodon was found primarily from the middle-late Miocene of Africa, Eastern Europe, and western and southern Asia. The discovery of this new material expands the known temporal and spatial distributions of this taxon and helps us better understand the phylogeny, evolution, and adaptive radiation of gomphotheres in early Miocene of China.

Among the most important faunas in the Late Cenozoic, the Gigantopithecus faunas have received a good deal of attention. The Gigantopithecus fauna recently discovered in Sanhe Cave consists of more than 80 mammal species, including cf. Hominidae, Pongo sp., Hylobates sp., Sinomastodon yangziensis, Stegodon preorientalis, Cervavitus fenqii, Dicoryphochoerus ultimus and Sus xiaozhu. It is the southernmost Gigantopithecus fauna found so far in China. Its geological age is estimated to be Early Pleistocene based on the fauna and stratigraphic correlation. The significant increase in the estimated body sizes of Ailuropoda, Gigantopithecus and Tapirus shows that the Sanhe fauna is middle Early Pleistocene, later than those from Wushan and Liucheng (early Early Pleistocene). Paleomagnetic dating of the fossil-bearing strata in Sanhe Cave gives an age of approximately 1.2 Ma. The fauna is characterized by tropical-subtropical forest types, including Pongo sp., Tupaia sp., Ia sp., Typhlomys intermedius, etc., and it lacks Palaearctic types. It is a typical tropical forest fauna, suggesting an environment with a lush forest and a warm and humid climate. The discovery of the Sanhe Gigantopithecus fauna is significant for establishing the chronological stages of the Gigantopithecus faunas in China, and for discussing their origin, evolution and dynamics.

Semedo site is rich on vertebrate fossils, with huge percentage come from Ordo Proboscidea. The aim of this paper is to identify the taxonomy of Proboscidean fossils important for reconstruction on Biostratigraphy of Java. This research uses a descriptive comparative method on morphological and morphometry characters, compared to similar data from Java and others related places. Based on this research we know that in Semedo site there are several species of Proboscidean, i.e.: Sinomastodon bumiayuensis, Stegodon trigonocephalus, Stegodon ”pygmy” semedoensis, Stegodon hypsilophus, Elephas (Archidiskodon) planifrons, and Elephas Hysudrindicus. This reflects that a long environmental changes or the history of ancient environment has been recorded in Semedo, related with ecological context of hominid appearance in this region.

The objective of this paper is to reply the recent comments on our papers made by Cozzuol et al. (2011) about the phylogeny and biogeography of South American Gomphotheres. We addressed the methodological criticism and concluded that Sinomastodon is the sister group of the South American gomphotheres, as we originally proposed, despite the weighting scheme adopted to analyze the data. Therefore, we argue that our conclusions based on this phylogenetic hypothesis on the evolution and biogeography of this group is not properly refuted by Cozzoul et al.

The trilophodont gomphothere group is a monophyletic group of genera separated from other proboscideans by one synapomorphy, the presence of trefoil-shaped wear patterns on the occlusal surfaces of their teeth. These wear patterns vary from being just a single trefoil, through double trefoils, to more complex combinations of trefoils. They are present in all of the genera studied here. Our cladistic analysis makes use of 12 genera as terminal taxa. It was performed using 43 cranial, dental and postcranial morphological characters. The polarity of characters was based on outgroup criteria using the genus Phiomia as the outgroup. All characters are considered to be unordered (34 are binary and nine are multistate). The present analysis leads us to reject the hypothesis that considered Rhynchotherium as a direct ancestor of South American gomphotheres and supports the hypothesis that Sinomastodon is the sister group of Cuvieronius and Stegomastodon. High congruence between the stratigraphical record and the phylogenetic hypothesis is observed., The present work was made possible through a joint Research Project from AECI, Spain-Argentina (2000–2001); Projects PB97-1250, BTE2001-1684 and CGL2004-00400/BTE from DGICYT, Spain; grants from the Universidad Nacional del Centro, and the Project PIP-02773, CONICET and Picto 04-11503 ANPCYT, Argentina.