Haowen Tong

Haowen Tong
  • Ph D
  • Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences

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Publications (97)
Article
The Yanjinggou fauna has long been recognized as the quintessential Middle Pleistocene fauna in South China. However, the recent discovery of the Dayakou fauna in the same region has unveiled a fossil assemblage indicative of the Early Pleistocene. Our new excavations at the Dayakou pit have yielded a substantial increase in fossil materials, which...
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The trend of giant panda (Ailuropoda, Ursidae) body size evolution during the Quaternary has long been observed primarily through dental size analysis. However, exact body mass estimations have been lacking, whereas the dental size is generally viewed as a poor index for body mass estimation. In this study, we assess the accuracy of body mass predi...
Article
The typical Mammuthus-Coelodonta Faunal Complex dominated the mid- and high latitudes of continental Eurasia during the Mid-Late Pleistocene; the dominant taxa include woolly mammoth (Mammuthus primigenius), woolly rhino (Coelodonta antiquitatis), steppe bison (Bison priscus) and diverse horses (Equus spp.), etc. Recent excavations at the Early Ple...
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2.2-1.7 MaBP。随 着地层古生物工作的深入开展,盆地内也发现了若干中-晚更新世化石点;丁家堡水库全新统中发现的 象颊齿,之前被鉴定为亚洲象,新的测年数据表明其时代大于 5 万年,依据牙齿测量数据和形态特征, 本研究将其归入诺氏古菱齿象。目前已在泥河湾盆地发现百余个化石地点,鉴定出 236 种(包括未定属种) 哺乳动物,分属于 8 目、32 科和 121 属,其中 38 个属种(包括亚种)最初是以泥河湾化石材料而建立。 总而言之,泥河湾盆地哺乳动物化石以早更新世者居多,并且化石材料保存完好;有些属种在欧亚大陆 古北区第四纪哺乳动物群形成过程中发挥过重要作用,例如早期猛犸象、披毛犀、野牛及真枝角鹿等, 还有直隶狼、貉及各种真马。泥河湾盆地由于河流和断层切割以及沉积相变等原因,导致各个化...
Article
Excavations in Hualongdong (HLD), East China, have yielded abundant hominin fossils dated to 300 ka. There is a nearly complete mandible that fits well with a partial cranium, and together they compose the skull labeled as HLD 6. Thus far, detailed morphological description and comparisons of the mandible have not been conducted. Here we present a...
Article
In the Shanshenmiaozui (SSMZ) fauna from Nihewan Basin, Canis chihliensis, Nyctereutes sp., Homotherium sp., Acinonyx sp., Lynx shansius and Pachycrocuta licenti have been identified. C. chihliensis is the dominant and best represented species whose fossil materials include crania, mandibles and postcranials, including nearly complete manus and pes...
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Giant deer is among the most common animals of Mid-Late Pleistocene sites in northern China, and was one of the representative icons of the Pleistocene fauna in northern China, while the fossil materials are fairly poor and very few complete antlers were ever recovered, except those from Zhoukoudian site; on the other hand, the knowledge about the...
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The giant panda, currently restricted to a small region of central China, was once widely dispersed throughout southern China and even some parts of continental Southeast Asia during the Pleistocene epoch. However, the evolutionary process and intraspecific variation remain poorly understood in light of limited fossil records, which are usually iso...
Article
Asiatic black bears have long been recognized as key members of the Quaternary mammalian fauna in South China. Despite this, for decades, taxonomic remarks since the Early Pleistocene Ursus thibetanus have been controversial, and the intraspecific variations of it remain unresolved. We here described a nearly complete black bear skull from the late...
Article
Bailong Cave (Bailongdong), dating back to the Middle Pleistocene, is a very important Palaeolithic site in southern China with some hominin remains and substantial mammalian fossils unearthed. Cervid fossils are one of the most common elements in this site, and antler fossils from Bailong Cave were preserved in a better condition than any other Pl...
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Late Middle Pleistocene hominins in Africa displaying key modern morphologies by 315 ka are claimed as the earliest Homo sapiens. Evolutionary relationships among East Asian hominins appear complex due to a growing fossil record of late Middle Pleistocene hominins from the region, reflecting mosaic morphologies that contribute to a lack of consensu...
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The Chinese open-air site Xuchang-Lingjing (Henan) is located near the border between the Palearctic and the Oriental biozone in a lowland depression at the southern edge of the North China Plain. The site yielded a Middle Palaeolithic assemblage that includes the two fragmented, incomplete human (possibly Denisovan) skulls (Xuchang 1 and Xuchang 2...
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2.2-1.7 MaBP。 由于受河流及断层切割影响,泥河湾盆地内部不同区域尤其是桑干河两岸的地层对比,一直是个科学难题, 其根本原因在于长期以来在桑干河南岸未发现重要化石点。最近,在桑干河南岸的岑家湾台地发现了山 神庙咀化石点,目前已在此地点做过 9 次正规发掘,出土哺乳动物化石 1526 件,鉴定出 25 个种(含未定 种),其中 23 种属于狭义泥河湾动物群的成员。由此看出,山神庙咀动物群的时代应当与狭义泥河湾动 物群的时代接近,其地质年代应当不晚于 1.7 MaBP。山神庙咀遗址与小长梁遗址隔沟相望,但其化石层 稍高于小长梁文化层;山神庙咀动物群的研究结果还表明,小长梁遗址年代要比先前所测得的 1.36 MaBP 要早;小长梁周边的史前遗址的地质年代应当再做深入研究。小长梁地区的泥河...
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The Bering Land Bridge (BLB) last connected Eurasia and North America during the Late Pleistocene. Although the BLB would have enabled transfers of terrestrial biota in both directions, it also acted as an ecological filter whose permeability varied considerably over time. Here we explore the possible impacts of this ecological corridor on genetic...
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The YJW (Yangjiawan) Cave 2 of Pingxiang in Jiangxi Province is a karst cave that developed in the Permian limestone of the Changxing Formation, which is filled with clay and grit of Late Pleistocene age. Six excavations have been conducted at the site since 2015. More than ten thousand mammalian fossils have been unearthed, and the wild boar fossi...
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Background: Old World porcupines (Family: Hystricidae) are the third-largest rodents and inhabit southern Europe, Asia, and most regions of Africa. They are a typical indicator of warm climate and their distribution is restricted to tropical and subtropical zones. In China, porcupines are widely distributed in southern areas of the Yangtze River....
Article
The mainstream school of human evolution posits that Homo erectus was the earliest species to leave Africa at ∼1.85 million years (Ma) ago. Recent discoveries from the Shangchen loess-palaeosol sequence near the Lantian hominid site in northern China, however, show lithic artifacts up to 2.12 Ma, pre-dating the fossil record of H. erectus. Here we...
Chapter
Our understanding of the emergence and dispersal of the earliest tool-making hominins has been revolutionised in the last decade, with sites in eastern Africa and China pushing records of both events several hundred thousand years earlier than previously thought. In recent years, climate and environmental factors have been considered by many as pri...
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Collaborative hunting by complex social groups is a hallmark of large dogs (Mammalia: Carnivora: Canidae), whose teeth also tend to be hypercarnivorous, specialized toward increased cutting edges for meat consumption and robust p4-m1 complex for cracking bone. The deep history of canid pack hunting is, however, obscure because behavioral evidence i...
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The YJW (Yangjiawan) Cave 2 of Pingxiang in Jiangxi Province is a karst cave that developed in the Permian limestone of the Changxing Formation, which is filled with clay and grit of Late Pleistocene age. Six excavations have been conducted at the site since 2015. More than ten thousand mammalian fossils have been unearthed, and the wild boar fossi...
Article
A diversity of pliopithecoids is known from Miocene localities in Europe, but until recently, this group was relatively poorly represented in China. However, new discoveries have shown that Chinese pliopithecoids were taxonomically diverse and geographically widespread. The earliest pliopithecoids in China (and Eurasia) are Dionysopithecus and Plat...
Article
Objectives: The paleontological description and comparative analysis using discrete morphology, morphometrics (linear and geometric) and cross-sectional geometry of three femoral diaphyseal sections from the Middle Pleistocene site of Hualongdong, China. Materials and methods: The material consists of the original Hualongdong femoral fossils and...
Article
The sambar (Rusa unicolor) is among the main elements of the Oriental Realm. The sambar fossils unearthed from Bailong Cave site dated back to the Middle Pleistocene were better preserved than those from any other fossil site. We described these remains especially focusing on the deciduous teeth and petrosal bone as well as the bony labyrinth it ho...
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8 枚古人类 牙齿化石、若干石制品和骨制品及大量哺乳动物化石。但有关白龙洞遗址的形成时间却一直存在争议, 主要原因之一是动物群没有得到充分研究。基于郧西县文管所和中科院古脊椎所收藏标本,本文首次对 白龙洞历年出土的哺乳动物化石进行了综合研究;研究结果表明,白龙洞动物群由 32 种(含未定种) 哺乳动物组成,优势属种是谷氏大额牛、岩羊、野猪及各种鹿类 (水鹿、斑鹿、麂、毛冠鹿);此外, 还有豪猪、竹鼠、变异狼、貉、豺、猪獾、黑熊、大熊猫、中国硕鬣狗、锯齿剑齿虎、虎、豹、金猫、 云豹、剑齿象、苏门犀、梅氏犀、巨貘、鬣羚及水牛等;该动物群缺乏猴类动物,并且豪猪化石也很少, 这与其所处的时空位置很不相符;但总体面貌属于大熊猫-剑齿象动物群。白龙洞的大熊猫、中国硕鬣 狗、猪獾、虎及巨貘的牙齿测量数值均...
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Recent excavations at Shanshenmiaozui site in Nihewan Basin of North China uncovered a rich collection of comb-antlered deer, which includes the first discovery of the yearling antler, complete upper and lower dentitions (both deciduous and permanent), associated limb bones including the longest metapodials ever recovered. Based on toothrow length...
Article
Recent excavations at Shanshenmiaozui site in Nihewan Basin of North China uncovered a rich collection of comb-antlered deer, which includes the first discovery of the yearling antler, complete upper and lower dentitions (both deciduous and permanent), associated limb bones including the longest metapodials ever recovered. Based on toothrow length...
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Significance Human evolution through the Middle to the Late Pleistocene in East Asia has been seen as reflecting diverse groups and discontinuities vs. a continuity of form reflecting an evolving population. New Middle Pleistocene (∼300,000 y old) human remains from Hualongdong (HLD), China, provide further evidence for regional variation and the c...
Chapter
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Yangjiawan Cave 2 in Pingxiang, Jiangxi is the fossil sites that produced the most abundant fossils in Jiangxi Province. The Carnivora here are mainly represented by isolated146 teeth, yet the high abundance and diversity of the Carnivora guild can greatly contribute to reconstructing the Southern China paleofauna during the Late Pleistocene and ar...
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The mammalian fauna from the Yangjiawan caves in Jiangxi contains more than 40 species, including Stegodon and Elephas, whose age is Late Pleistocene. The 40 pieces of Stegodon fossils include such elements as DP2, DP3, DP4, M1, M3, dp3, m2 and m3, which can be included in the species S. orientalis according to their morphological characters and th...
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Elasmotherium or the Giant Siberian Unicorn is quite common in Siberia and adjacent regions during the Quaternary Period, but it only appeared in very few sites of Early Pleistocene age in northern China. In the past century, quite a number of specimens of Elasmotherium were recovered in Nihewan Basin, most of which are disarticulated postcranial b...
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The Hualong Cave was recovered in 2004, and 4 excavations have been conducted during 2006 and 2014-2016, which resulted in the discovery of several human fossils and numerous mammalian fossils. This paper is a preliminary report on the mammalian fossils unearthed during 2014-2016. Currently 43 mammalian species, including indeterminate species, bel...
Article
In southern China, the Ailuropoda-Stegodon fauna is a very general one with a relatively stable Ailuropoda and Stegodon fossil assemblage throughout the Pleistocene epoch, and cervid remains are also significant elements of this fauna. The taxonomic study on isolated cervid tooth fossils of the Late Pleistocene Ailuropoda-Stegodon fauna is not very...
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The Hualong Cave was recovered in 2004, and 4 excavations have been conducted during 2006 and 2014-2016, which resulted in the discovery of several human fossils and numerous mammalian fossils. This paper is a preliminary report on the mammalian fossils unearthed during 2014-2016. Currently 43 mammalian species, including indeterminate species, bel...
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The new fossils of Bos primigenius in this study came from two regions in Hebei Province: Nihewan Basin (one partial skull with right horn-core and one humerus) and Longhua County (one humerus, one tibia, 2 calcanea and 3 astragali). The humerus from Nihewan has a total length of 457.3 mm, which almost represents the largest aurochs individual whic...
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By at least 45,000 years before present, anatomically modern humans had spread across Eurasia [1-3], but it is not well known how diverse these early populations were and whether they contributed substantially to later people or represent early modern human expansions into Eurasia that left no surviving descendants today. Analyses of genome-wide da...
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The Early Pleistocene Shigou Site is located in Nihewan Basin, Hebei Province, North China. The geographic coordinates are 400 13' 53.8'' N, 1140 40' 19.6'' E, 830 m. In 2014 Shigou Site was excavated by the Hebei Normal University (trench A) and IVPP (trench B). This paper is a report on the excavation of trench B and the analysis on the fauna of...
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Yangjiawan Cave 2 is one of the components of Yangjiawan fossil site, which was excavated for the first time in November 2015 and October 2016. Thousands of mammalian fossils have been unearthed from Yangjiawan Cave 2, which can be referred to 31 species (including indeterminate species). Most of the fossils are teeth and few are broken bones. Sus...
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Shanshenmiaozui is an open-air fossil site of Early Pleistocene age in Nihewan Basin, China. During 2006?2011, nearly 100 fossil specimens of Mammuthus trogontherii were unearthed, including some series of tarsal, metatarsal and phalange bones of immature individuals, which represent the first discovery of hindfoot bones of immature M. trogontherii...
Article
Maludong is a Terminal Pleistocene fossil-bearing cave located on the northern edge of the Southeast Asian tropical zone, southeastern Yunnan Province. Hominins from the cave include remains with affinities to archaic hominins and others with an apparent mixture of archaic and modern traits all deriving from deposits dating from the Bølling-Allerød...
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Since the establishment of the species . Bison palaeosinensis by Teilhard de Chardin and Piveteau in 1930, no new fossils have been reported in the last few decades. Recent excavations at the Shanshenmiaozui site in Nihewan Basin of North China led to the discovery of numerous Bovini fossils, including a nearly complete mandible and articulated pos...
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For a long time, research about Nihewan Basin vertebrate fossils has focused on the evolution of major taxa and biostratigraphy; in contrast, the study of burial process and population ecology of these fossils is almost absent. Recently, scientific excavation of the Shanshenmiaozui locality provided an opportunity to undertake a taphonomy study in...
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The hominin record from southern Asia for the early Late Pleistocene epoch is scarce. Well-dated and well-preserved fossils older than ∼45,000 years that can be unequivocally attributed to Homo sapiens are lacking. Here we present evidence from the newly excavated Fuyan Cave in Daoxian (southern China). This site has provided 47 human teeth dated t...
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The mysterious loss of five calvaria of Homo erectus and three skulls of Homo sapiens sapiens, unearthed from Zhoukoudian, a world cultural heritage site, during World War II is a great loss for paleoanthropological research. Although frontal and occipital bones of Homo erectus were unearthed in 1966, additional important human fossils cannot be fo...
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The fossil tapirid records of Late Miocene and Early Pliocene were quite poor in China as before known. The recent excavations of the terminal Miocene hominoid site (between 6 and 6.5 Ma) at Shuitangba site, Zhaotong in Yunnan Province resulted in the discovery of rich tapir fossils, which include left maxilla with P2-M2 and mandibles with complete...
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Numerous juvenile specimens, including skulls and deciduous teeth of Mammuthus primigenius (Blumenbach, 1799), have been reported, which is rare among the recovered fossil records of the early mammoth species including Mammuthus meridionalis (Nesti, 1825) and Mammuthus trogontherii (Pohlig, 1885). After the publication of the juvenile mandibles and...
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Peking Man, originally named Sinanthropus pekinensis and now known as Homo erectus pekinensis, was among the best documented extinct hominids before the catastrophic loss of almost all the material in 1941. The only diagnostic specimens to survive from the original excavations are some teeth in the paleontological collections of Uppsala University,...
Article
Timing of the mammalian faunas in the Nihewan Basin, North China has provided insights into our understanding of Quaternary biochronology and biostratigraphy in East Asia. Here we contribute to this topic with detailed magnetostratigraphic investigation, coupled with mineral magnetic measurements on a fluvio-lacustrine sequence in this basin, which...
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The recently discovered macaque fossils, including isolated teeth and mandible fragments, from Tianyuan Cave and the juvenile maxilla from Wanglaopu Cave near Zhoukoudian can be identified as Macaca mulatta; they represent the first recognized occurrence of fossil M. mulatta in the far north of China. Some previously unearthed but still unpublished...
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Shanshenmiaozui in the Nihewan Basin of China is the locality that has produced the richest sample of Coelodonta nihowanensis material, including a nearly complete juvenile skull and mandible as well as various other cranial, mandibular, and postcranial bones. All of the cranial and mandibular specimens belong to individuals around 1.5 years old. O...
Article
In China, the non-Coelodonta dicerorhines are too diverse to be placed entirely in the genus Dicerorhinus. Most of the Pleistocene species should be transferred to the European genus Stephanorhinus because they differ from the Dicerorhinus species. Those differences include a much larger body size, a dolicocephalic skull, the absence of incisors, a...
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From 2006−2011, numerous Canis specimens were recovered from the newly discovered Shanshenmiaozui Locality in the Nihewan Basin. The material includes broken crania, premaxillae, maxillae, mandibles and postcranial skeleton with most of the elements preserved, and represents the most extensive and informative sample of early fossil Canis ever recov...
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Rich fossil materials of Mammuthus trogontherii have been recovered from a new locality, Shanshenmiaozui, in the Nihewan Basin during the past several years. The materials include the most complete juvenile mandible ever discovered and some postcranial bones of Mammuthus trogontherii. The geologic age of the fossil bearing strata is about 1.3 Ma ba...
Article
: The current and dominant theory about the origin of modern humans is the out-of-Africa hypothesis, which asserts that populations of Homo sapiens left Africa 100,000 years ago and replaced indigenous populations of humans in Eurasia. Many scholars equated the out-of-Africa dispersal of humans with paleoenvironmental changes. However, until now, f...
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The Rhino Cave, a Paleolithic site in Shennongjia, is the highest altitude locality of rhinoceros in the Middle-South part of China, and it is also the southernmost and the richest locality in Stephanorhinus kirchbergensis fossils which include cranium, mandibles, isolated teeth and postcranials. These materials can be referred to the species Steph...
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Fauna lists from nature reserves, game reserves and national parks are commonly used to depict modern fauna communities in synecological studies for the reconstruction of palaeocommunities and palaeoenvironments. This relies on the assumption that the modern fauna lists are truly representative samples of the communities they constitute. This study...
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We report here on the isotopic analysis of the diet of one of the oldest modern humans found in Eurasia, the Tianyuan 1 early modern human dating to approximately 40,000 calendar years ago from Tianyuan Cave (Tianyuandong) in the Zhoukoudian region of China. Carbon and nitrogen isotope analysis of the human and associated faunal remains indicate a...
Article
The Liucheng Gigantopithecus Cave is very famous for its rich mammalian fauna and Gigantopithecus fossils. The Gigantopithecus Cave Fauna has been regarded as the typical Early Pleistocene fauna in South China. The majority of the fossils unearthed has been studied and published during the past decades. The only group remaining unpublished is the r...
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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, Dicoryph...
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The Tianyuan cave near Zhoukoudian is among the most special localities in North China for its many porcupine gnawing marks on the bones, as is often the case acknowledged only in South China. But porcupines did not gnaw all the bones, that is gnawing is selective. Skulls, mandibles, tooth roots and long bones are frequently gnawed. The short bones...
Article
Today, Hystrix is a genus that only occurs in warm areas. However, during the Pleistocene and even the early Holocene, this species frequently appeared in the northern part of China, which belongs to the Palearctic Region. A total of 22 fossil Hystrix-bearing localities have been reported in North China, the time span of which covers almost all the...
Article
Giant deer with palmate brow tines were placed either in a single genus, or those from west Eurasia were placed in Megaloceros and those from the east in Sinomegaceros, which implies local evolution. In order to see whether dispersals between the two areas may have occurred, we studied the morphology and interpreted the phylogeny of these deer.The...
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Key words Xitaiping Cave, Shidu, Beijing; Late Pleistocene; mammalian fossils
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The north and south China faunas are subdivided along the line of Huaihe River-Qinling Mountains-Hengduan Mountains-Himalayas, to the north is the Palearctic Region, and to the south is the Oriental Region, which is the result of long-time evolution. Hundreds of Quaternary fossil localities have been known up to now, more than 60 of which contain w...
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Thirty-four elements of an early modern human (EMH) were found in Tianyuan Cave, Zhoukoudian, China in 2003. Dated to 42,000–39,000 calendrical years before present by using direct accelerator mass spectrometry radiocarbon, the Tianyuan 1 skeleton is among the oldest directly dated EMHs in eastern Eurasia. Morphological comparison shows Tianyuan 1...
Article
The materials of Aeretesmelanopterus from the Tianyuan Cave, a Late Pleistocene site with human fossils recovered recently near Zhoukoudian, are the richest of this species ever reported in North China. Aeretesmelanopterus is an endemic species of China, its relationship with Petaurista and Trogopterus is still controversal. Studies based on the de...
Article
The extinction of large-bodied taxa from the Pleistocene in Southeast Asia is examined. Although the chronological resolution of these extinctions is poor, and number of excavations in the region relatively few, broad characteristics of these extinctions can be described. Many taxa which became extinct appear to have been endemic to regions within...
Article
During Pleistocene Epoch, quite a number of meridionale mammals inhabited north China, such as Stegodon, Macaca, Hystrix, Dicerorhinus, Bubalus and several kinds of other probscideans. On the contrary, some boreal mammals once appeared in south China, such as Megaloceros and Ursus arctos etc. The frequent appearance of warm-adapted mammals in North...
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Key words Gigantopithecus Cave, Guangxi, China, Early Pleistocene, Chalicotheridae
Article
FOXP (FOXP1-4) is a newly defined subfamily of the forkhead box (FOX) transcription factors. A mutation in the FOXP2 forkhead domain cosegregates with a severe speech disorder, whereas several mutations in the FOXP3 forkhead domain are linked to the IPEX syndrome in human and a similar autoimmune phenotype in mice. Here we report a 1.9 A crystal st...
Article
Most of the Quaternary tapir fossils from China are isolated teeth. The purpose of this paper is to identify them and to extract systematic and evolutionary information from them. Based on morphology and W/L ratio, isolated teeth can be identified successfully. On the whole, the identification of P1, M3 and P2 is believed to be reliable, while it i...
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The Tianyuan Cave is the only human fossil-bearing site containing rich mammalian fossils found in the last decades near Zhoukoudian. Up to now more than 34 specimens of the human body have been recovered, and the mammalian fossils can be put into 29 species. Cervids dominate the fauna, and carnivores are very rare. Based on the primary examination...
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The materials of Tapirus from Renzidong, Fanchang, Anhui Province, are the best ones among the early most Pleistocene findings in China up to now, not only in richness, but also in completeness. The fossils contain complete tooth rows of both the upper and the lower, as well as most parts of the postcranial skeletons. These materials are very helpf...
Article
Under the family Rhinocerotidae, seventeen genera, sixty-two species (including four subspecies and two variants) were recognized or originally named in China. Chilotherium is the most diversified genus, which contains eighteen species and two variants. Rhinoceros sinensis, Coelodonta antiquitatis and Dicerorhinus mercki are the three most frequent...
Article
A total of 60 pieces of rhino materials have been identified from the Nanjing Man site. Morphologically, they belong to the species Dicerorhinus mercki. Among the materials, 40 pieces are broken bones, and 20 are isolated teeth or jaw bones (totally comprising 41 tooth units). Among the teeth, 74% are deciduous, most of them attached to jaws. Other...
Article
The presence of rhinoceroses on sites yielding human fossils in China. At the present time in China, 74 paleolithic sites yielding human remains have been discovered, 58 of which have rhinoceros fossils (78 %). Among the recovered fauna, rhinoceroses are the animals which show the most relationships with Man. In spite of the state of preservation,...

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