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    A Late Pliocene Hipparion houfenense fauna from Yegou, Nihewan Basin and its biostratigraphic significance
    LIU Jin-Yi, ZHANG Ying-Qi, CHI Zhen-Qing, WANG Yong, YANG Jin-Song, ZHENG Shao-Hua
    Vertebrata Palasiatica    2022, 60 (4): 278-323.   DOI: 10.19615/j.cnki.2096-9899.211209
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    Currently, there are still different views regarding the chronology of the Late Cenozoic deposits in the Nihewan Basin, which results from the contradiction between biostratigraphic correlations based on mammalian fossils and magnetostratigraphic dating results. Biostratigraphic correlations indicate that the aeolian red clay exposed in the Sanggan River canyon, the fluvio-lacustrine red clay with sands and gravels, and the sandy clay of swamp facies on both sides of the lower reaches of the Huliu River belong to the Upper Pliocene, whereas the magnetostratigraphic dating usually correlates them to the Lower Pleistocene. In October 2011, a collection of mammalian fossils was unearthed from a block of collapsed deposits at Yegou in the Nihewan Basin, which is about 300 m north of the Laowogou section that is well known for the Pliocene mammalian fossils from its lower part. The Yegou fossils are identified herein as 10 species in 9 genera: Nyctereutes tingi, N. sinensis, Pachycrocuta pyrenaica, Homotherium sp., Hipparion (Plesiohipparion) houfenense, Dicerorhinus sp., Muntiacus sp., Axis shansius, Gazella blacki, and Paracamelus sp. The fauna is quite different from the classic Early Pleistocene Nihewan Fauna in composition and provides new evidence for the existence of the Upper Pliocene in the Nihewan Basin. Based on a systematic description of the fauna, its composition and geological age are discussed, and the compositional features of large mammals of the Late Pliocene and the Early Pleistocene mammalian faunas in the Nihewan Basin are summarized.

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    A juvenile skull of the longirostrine choristodere (Diapsida: Choristodera), Mengshanosaurus minimus gen. et sp. nov., with comments on neochoristodere ontogeny
    YUAN Meng, LI Da-Qing, Daniel T. KSEPKA, YI Hong-Yu
    Vertebrata Palasiatica    2021, 59 (3): 213-228.   DOI: 10.19615/j.cnki.2096-9899.210607
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    Choristoderes were an important clade of semi-aquatic predators that occupied Laurasian freshwater ecosystems from the Middle Jurassic to the Miocene. During the Early Cretaceous, the neochoristodere lineage evolved large size and long snouts, converging on the body plan of modern crocodilians. Here, we describe a new longirostrine choristodere, Mengshanosaurus minimus gen. et sp. nov. from the Lower Cretaceous Mengyin Formation of Shandong Province, China. The holotype is the smallest reported neochoristodere individual, with a skull length of only 35 mm. The poorly ossified braincase, along with retention of a fontanel at the frontal-parietal suture, indicates this individual was a juvenile. Phylogenetic analyses recovered Mengshanosaurus as a neochoristodere, a placement supported by the presence of a single narial opening, fusion of the nasals, and expansion of the temporal fenestrae. In the Neochoristodera,Mengshanosaurus is sister to a clade consisting of Ikechosaurus, Tchoiria, Simoedosaurus, and Champsosaurus. It differs from other neochoristoderes in having the lacrimal foramen between the prefrontal and lacrimal, in addition to having large vomerine teeth (exceeding one-third the width of corresponding maxillary teeth). The closely arranged marginal teeth and large vomerine teeth suggest juvenile choristoderes may have fed on invertebrates and insects, similar to juveniles of modern crocodilians. However, the observation that very young neochoristoderes had similar skull proportions and marginal tooth shapes to adults, along with features suggesting a more fully aquatic ecology, suggest that neochoristoderes exhibited less pronounced ontogenetic niche shifts than modern crocodilians.

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    Reappraisal of the largest ctenochasmatid Moganopterus zhuiana Lü et al., 2012
    GAO Dian-Song, JIANG Shun-Xing, XU Li, CHENG Xin, YANG Li-Li, JIA Song-Hai, WANG Xiao-Lin
    Vertebrata Palasiatica    2022, 60 (3): 197-211.   DOI: 10.19615/j.cnki.2096-9899.220111
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    Moganopterus zhuiana Lü et al., 2012 was erected as a member of the Boreopteridae, which was questioned by different researchers shortly after the publication. Although the new assignment to the Ctenochasmatidae is widely accepted by pterosaur researchers, some characteristics still require a detailed description. Here, the holotype of this taxon is restudied, and some ambiguous characteristics are re-identified. The diagnosis of this taxon has been revised as the following: a large ctenochasmatid pterosaur, which can be distinguished from other members of this clade by a single autapomorphy: an elongated rod-like parietal crest that extends posterodorsally, forming an angle of about 15° with the ventral margin of the skull. This taxon can be further distinguished from other ctenochasmatids on the basis of the following combination of characteristics: straight occlusal surfaces of the upper and low jaws; presence of a low premaxillary crest confined anterior to the nasoantorbital fenestra; rostrum about two thirds of the skull length; nasoantorbital fenestra occupying slightly more than 20% of the skull length; about 100 slender teeth; and a mid-cervical length/width ratio of about 7. The wingspan of M. zhuiana has been re-estimated according to a simple regression equation for wingspan versus skull length in ctenochasmatids. It confirms that M. zhuiana, although smaller than previous thought, is still the largest known ctenochasmatid. When comparing the sizes of ctenochasmatids in the Jurassic and Cretaceous, ctenochasmatids showed a rough tendency to increase their sizes.

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    Redescription of Nochelaspis maeandrine ,the largest eugaleaspiform from the Lower Devonian of Qujing, Yunnan
    MENG Xin-Yuan, ZHU Min, GAI Zhi-Kun
    Vertebrata Palasiatica    2021, 59 (4): 257-272.   DOI: 10.19615/j.cnki.2096-9899.210727
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    The new specimens of the largest eugaleaspiform Nochelaspis maeandrine are redescribed from two localities of the Xishancun Formation in Qujing City, Yunnan Province, southwestern China. Nochelaspis is most suggestive of Yunnanogaleaspis from the same horizon, but differs in its slit-like median dorsal opening (length/width>6), much stronger inner cornual process, coarse stellate ornamentation, and the serrations along the edges of the median dorsal opening and headshield. The new findings reveal the morphological details on the ventral side of the headshield as well. The oralobranchial fenestra is covered by a large dermal ventral plate, which is decorated with dense, tiny granular tubercles, and aligned with six pairs of separated, large, and circular branchial openings. This condition is different from that of osteostracans, in which the oralobranchial fenestra is covered by numerous minute scales or larger dermal platelets, and the branchial openings are slit-shaped and covered by small skin flaps somewhat like those of elasmobranchs. However, the branchial openings of galeaspids and osteostracans are both located ventrally as in modern rays, indicating a benthic lifestyle dwelling on sandy or muddy substrates in a quiet marine environment.

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    Vayu 1.0, a new set of tools for visualizing surface meshes
    LU Jing
    Vertebrata Palasiatica    2023, 61 (1): 71-80.   DOI: 10.19615/j.cnki.2096-9899.221020
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    Vayu 1.0 is a freeware that deals with surface mesh files. It provides a set of tools for rendering, labelling, revisualizing, and reanalyzing meshes. It also offers features for VR mode and one-stop animation production. This paper describes the major features of Vayu 1.0, which includes three main panels, i.e. mesh information, keyframe editor, and shading-transform. Vayu, as an innovative freeware, contains the state-of-the-art new features which provides a fresh set of tools to accelerate future development directions in paleontology, biological sciences and beyond.

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    Reanalysis of Oculudentavis shows it is a lizard
    LI Zhi-Heng, WANG Wei, HU Han, WANG Min, YI Hong-Yu, LU Jing
    Vertebrata Palasiatica    2021, 59 (2): 95-105.   DOI: 10.19615/j.cnki.1000-3118.201020
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    The recent finding of a fossil entombed in a Late Cretaceous amber - Oculudentavis khaungraae - was claimed to represent a humming bird-sized dinosaur. Regardless of the intriguing evolutionary hypotheses about the bauplan of Mesozoic dinosaurs (including birds) posited therein, this enigmatic animal demonstrates various morphologies resembling lizards. If Oculudentavis was a bird, it challenges several fundamental morphological differences between Lepidosauria and Archosauria. Here we reanalyze the original computed tomography scan data of the holotype of Oculudentavis khaungraae (HPG-15-3). Morphological evidences demonstrated here highly contradict the avian or even archosaurian phylogenetic placement of the species. In contrast, our analysis revealed multiple skull morphologies of HPG-15-3 resembling those of squamates, including pleurodont marginal teeth, an open infratemporal fenestra, and the presence of palatal dentition. Based on these new morphological information, the phylogenetic position of Oculudentavis was analyzed in a data matrix sampling across the Diapsida. Taxon sampling of the data matrix included multiple species of lizards, birds, and major clades in Lepidosauromorpha and Archosauromorpha. In the strict consensus tree, Oculudentavis is nested within Squamata. These results show that morphology of the Oculudentavis khaungraae holotype supports a squamate rather than avian or dinosaurian affinity of the species.

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    XU Xing, GUO Yu
    Vertebrata Palasiatica    2009, 47 (4): 311-329.  
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    Recent paleontological and neontological studies on feathers and feather-like integumentary structures have improved greatly our understanding of the origin and early evolution of feathers. New observations on some non-avian dinosaur specimens preserving integumentary structures, in combination with recent paleontological and neontological data, provide additional insights into this important evolutionary issue. Five major morphogenesis events are inferred to have occurred sequentially early in feather evolution before the origin of the Aves, and they are: 1) appearance of filamentous and tubular morphology, 2) formation of follicle and barb ridges, 3) appearance of rachis, 4) appearance of planar form, and 5) formation of pennaceous barbules. These events produce several morphotypes of feathers that are common among non-avian archosaurs but are probably lost later in avian evolution, and they also produced several morphotypes of feathers that are nearly identical or identical to those of modern birds. While feathers of non-avian dinosaurs exhibit many unique features of modern feathers, some of them also possess striking features unknown in modern feathers. Several models of evolutionary origin of feathers based on developmental data suggest that the origin of feathers is a completely innovative event and the first feathers have nothing to do with reptilian scales. We believe, however, that the defining features of modern feathers might have evolved in an incremental manner rather than in a sudden way. Consequently, an evolutionary model characteristic of both transformation and innovation is more acceptable for feather evolution. The function of the first feather is inferred to be neither related to flight nor to insulation. Display or heat dissipation, among others, remains viable hypotheses for initial function of feathers. An integrative study is promising to provide much new insights into the origin of feathers.
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    On the scientific names of mastodont taxa: nomenclature, Chinese translation, and taxonomic problems
    WANG Shi-Qi, LI Chun-Xiao, ZHANG Xiao-Xiao
    Vertebrata Palasiatica    2021, 59 (4): 295-332.   DOI: 10.19615/j.cnki.2096-9899.210728
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    The mastodont-grade proboscideans represent an important stage in the evolution of the group, establishing the basic pattern of the evolution of the crown groups of proboscideans. The research on mastodons has a history of more than 400 years. The classification and nomenclature have been revised and changed many times, and the problems in their evolution were fully reflected in the history of mastodon nomenclature. In this paper, we undertook a bibliographical research into the nomenclature and etymology of various mastodont groups, reviewing 175 translated Chinese names of mastodont-grade proboscideans, including 12 taxon names higher than the genus level, 46 genera, and 117 species, covering almost all the species of the mastodont radiation. On this basis, we review the principal phylogenetic hypotheses of mastodont interrelationships, and highlight problems in the classification and nomenclature of mastodonts. The evolution of the skull and mandible of mastodons is continuous in all clades, reflecting the same parallel evolution trend; while, although the morphological characteristics of cheek teeth across all lineages are not obvious, they are relatively stable in each lineage. Choerolophodontidae is the most robust monophyletic group within the mastodonts, of which Synconolophus may be a distinct, valid genus. Miomastodon and Pliomastodon of Mammutidae may both be valid, but they are not necessarily the direct ancestor of Mammut americanum . The phylogenetic relationship between Platybelodon danovi , P. grangeri and Aphanobelodon zhaoi within the Amebelodontidae is questionable, depending on whether the lower incisor section of P. danovi is the dentine rod structure or not, while Konobelodon britti in America may be a synonym of Torynobelodon loomisi . The species assigned to Konobelodon in Asia is possibly not amebelodontids, but probably attributable to Paratetralophodon , instead; Serridentinus of Gomphotheriidae may be a valid taxon, representing a trend towards somewhat zygodonty in Gomphotheriidae that terminated with the Cuvieroniinae. The Cuvieroniinae may only include Cuvieronius and Rhynchotherium , while other brevirostrine gomphotheres in America, such as Stegomastodon may have been evolved from a lineage of amebelodonts. Notiomastodon may be related to Sinomastodon , which itself may have originated from Pliomastodon (?) zhupengensis in southern China. The name Mastodon intermedius Teilhard de Chardin & Trassaert, 1937 (now Sinomastoodon intermedius ) has the senior primary homonym Mastodon intermedius Eichwald, 1831. We suggest that Sinomastodon intermedius should be replaced with its senior synonym-Sinomastodon sendaicus (Matsumoto, 1924).

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    A new Late Triassic tetrapod locality from North China
    SHI Yu-Tai, CHEN Jian-Ye, LIU Jun
    Vertebrata Palasiatica    2023, 61 (1): 17-25.   DOI: 10.19615/j.cnki.2096-9899.220818
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    No definite tetrapod body fossil is known from the Late Triassic of North China. Here we report new tetrapod remains from the Upper Triassic Tanzhuang Formation of Jiyuan, Henan Province. Most of the specimens can be referred to the temnospondyls, and show affinity with Mastodonsaurus. Previously, Mastodonsaurus is only know from Europe. So that, this finding indicates that a clade including Mastodonsaurus and its close relatives had a wide distribution from Europe to East Asia during the Late Triassic. A possible pelvis indicates the presence of another tetrapod group. This discovery fills a tetrapod body record gap in Chinese Late Triassic.

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    Revisit of Hsianwenia wui (Cyprinidae: Schizothoracinae) from the Pliocene of Qaidam Basin
    BI Dai-Ran, WU Fei-Xiang, WANG Ning, CHANG Mee-Mann, FANG Geng-Yu
    Vertebrata Palasiatica    2022, 60 (1): 1-28.   DOI: 10.19615/j.cnki.2096-9899.211026
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    The Qaidam Basin is a key area for understanding the paleoenvironmental and faunal history of the Tibetan Plateau. The fossil schizothoracine fish, Hsianwenia wui, evolved extraordinarily thickened bones to adapt to the aridification of the Qaidam Basin during the Pliocene. However, the nature of the bone thickening itself remains elusive. To promote the further investigation of the physiological mechanism of the pachyostosis and the phylogenetic interrelationships of Hsianwenia and all relevant cyprinids, here we present a comprehensive morphological study of Hsianwenia. We have new information on the anterior part of the cranial cavity, a large supraneural 3 in the Weberian apparatus, numerous procurrent caudal fin rays supported by the preural centrum (Pu) 5, and a neural arch on Pu2. We also find the differentiated pattern of the bone-thickening: the pachyostosis exists in the endoskeleton but not in the dermal skeleton; it is more obvious in ventral bones than in dorsal ones, when the thickening is present in the dorsally and ventrally grouped endoskeletal bones (e.g., the epineural and epipleural intermuscular bones). Considering the integrity of musculoskeletal system manipulating the chewing activities, we suspect that the thickened pharyngeal jaws and the hard food processing might be associated with the unique hind protrusion (cleithral “humeral” process) of the dermal pectoral girdle of Hsianwenia.

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    SATO Tamaki, LI Chun, WU XiaoChun
    Vertebrata Palasiatica    2003, 41 (01): 17-33.  
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    Bishanopliosaurus youngi Dong, 1980 from the Jurassic Dongyuemiao Member of the Ziliujing Formation in Chongqing Municipality, China is redescribed in detail. The holotype of B. youngi is the postcranial skeleton of a young individual, but so far the most complete plesiosaurian fossil known from the Asian Jurassic. Our restudy indicates that B. youngi can be diagnosed by the bifurcated sacral ribs combined with the compressed neural spine, narrow coracoid, and the humerus with a peculiar projection. B. youngi is a valid taxon, but its phylogenetic relationships with other plesiosaurians remain uncertain mainly because of the lack of cranial features. Previous studies on the Dongyuemiao fauna and flora indicate that the sediments are of subtropical freshwater origin. The discovery of B. youngi added further information on the freshwater invasion of plesiosaurians. Freshwater plesiosaurians so far known are very fragmentary, but their wide geographic and temporal distributions suggest that the presence of plesiosaurians in freshwater environment was not uncommon in most of their history.
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    A new aardwolf-line fossil hyena from Middle and Late Miocene deposits of Linxia Basin, Gansu, China
    Henry GALIANO, Z. Jack TSENG, Nikos SOLOUNIAS, WANG Xiao-Ming, QIU Zhan-Xiang, Stuart C. WHITE
    Vertebrata Palasiatica    2022, 60 (2): 81-116.   DOI: 10.19615/j.cnki.2096-9899.211025
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    The aardwolf Proteles cristatus is the only known hyaenid, living or extinct, to exhibit an extremely reduced dentition related to its termite-specializing diet. The fossil record of extant aardwolves extends to 2 to 4 million years ago, but records that inform its evolutionary origins are essentially nonexistent. Such circumstance renders it difficult to place this unusual hyena in the broader evolutionary context of small-bodied hyaenid species in Eurasian Neogene deposits. Here we describe a new genus and species of a small-bodied hyaenid, Gansuyaena megalotis, representing the closest morphological link to aardwolves to date. This new fossil hyena is based on a skull with associated mandible, a rostrum preserving several teeth, and several referred specimens. The new specimens were discovered in Neogene deposits in Linxia Basin, Gansu Province, China. Phylogenetic analysis indicates that among early hyaenids, G. megalotis is most closely related, but unlikely ancestral, to the living aardwolf. Also recognized in this new species are the fossils previously referred to “Protictitherium” aff. P. gaillardi from Pasalar, Turkey. Additionally, “Plioviverropsguerini from Los Mansuetos, Spain is interpreted to represent a second Gansuyaena species. In addition to the living aardwolf, Proteles cristatus, our analyses suggest that the proteline lineage includes the extinct genera Gansuyaena, Mesoviverrops, and Plioviverrops. Although the precise timing and geographic location of evolutionary divergence between the aardwolf and Gansuyaena remain elusive, critical new morphological information provided by Gansuyaena specimens reinforce findings from recent genomic analyses that the aardwolf lineage has an ancient origin from small-bodied stem hyaenids prior to the appearance of large and robust bone-cracking hyaenines.

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    Reappraisal of some perissodacyl fossils from the Middle Eocene of the Lijiang Basin, Yunnan, China with a revision of tapiroid Diplolophodon
    BAI Bin
    Vertebrata Palasiatica    2023, 61 (1): 26-42.   DOI: 10.19615/j.cnki.2096-9899.220721
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    The Lijiang Fauna from the Lijiang Formation of Lijiang, western Yunnan, was dominated by 13 species of perissodactyl fossils, and its age ranged from Irdinmanhan to Sharamurunian Asian Land Mammal Ages (ALMA) according to previous research. Based on reanalysis of some perissodactyls from the Lijiang Fauna, it is suggested here that Rhodopagus yunnanensis is a junior synonym of Lijiangia zhangae, which is similar to Lophiohippus and placed in Anchilophini within the Palaeotheriidae. The enigmatic and scarce Lunania is here regarded as a palaeothere rather than a chalicothere based on its morphological similarities with Paranchilophus, and Lophiohippus probably represents the upper dentitions of Lunania. Furthermore, deperetellid Diplolophodon is revised and comprised of three species: D. similis, D. lunanensis, and D. xiangshanensis (comb. nov.). Teleolophus xiangshanensis from the Lijiang Formation is reassigned to Diplolophodon xiangshanensis. The revised perissodactyls from the Lijiang Fauna are comparable to those from the Rencun Member of the Hedi Formation of the Yuanqu Basin, and its age is confined to Sharamurunian ALMA.

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    Reappraisal of Serridentinus gobiensis Osborn & Granger and Miomastodon tongxinensis Chen: the validity of Miomastodon
    WANG Shi-Qi, ZHANG Xiao-Xiao, LI Chun-Xiao
    Vertebrata Palasiatica    2020, 58 (2): 134-158.   DOI: 10.19615/j.cnki.1000-3118.200310
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    The elephantimorph proboscideans, Serridentinus gobiensis Osborn & Granger, 1932, and Miomastodon tongxinensis Chen, 1978, from the Middle Miocene of northern China, were revised as Zygolophodon gobiensis (Osborn & Granger, 1932). However, their phylogenetic positions are still being debated because of their intermediate morphology between the typical bunodont (Gomphotheriidae) and zygodont (Mammutidae) elephantimorphs. In the present paper, we compare their dental and mandibular morphology with that of the Eurasian Z. turicensis, Gomphotherium subtapiroideum, and G. tassyi, as well as the North American Mio. merriami and G. productum. It appears that S. gobiensis and Mio. tongxinensis share with Mio. merriami the slightly more bunodont molar morphology than that of Z. turicensis, e.g., the thicker enamel, thicker pretrite crescentoids, higher interlophid enamel pillars in buccal view, and the narrower contour majorly caused by the narrower posttrite half loph(id)s. S. gobiensis and Mio. merriami also possess an “erected oval cross-sectioned mandibular tusk”, in which the cross-section is mediolaterally compressed (dorsoventral diameter being larger than the mediolateral one). Whereas, in Z. turicensis and G. productum, the mandibular tusk is “laid oval cross-sectioned”, in which the cross-section is dorsoventrally compressed (dorsoventral diameter is smaller than the mediolateral one). Therefore, it is reasonable to revive the genus Miomastodon Osborn, 1922, which contains the species that were previously attributed to Zygolophodon, but they have relatively bunodont molar morphology (i.e., the robust type of the Z. turicensis group). The mandibular tusk with erected oval cross-section seems to be a synapomorphy of Miomastodon species. Furthermore, the molar morphology of G. subtapiroideum and G. tassyi also exhibits intermediate status between the typical bunodonts and zygodonts. However, the mandibular symphysis of G. subtapiroideum and G. tassyi is stronger than that of Miomastodon, and the mandibular tusk is pyriform cross-sectioned. The validity of Miomastodon and G. subtapiroideum/tassyi obscures the boundary between the Gomphotheriidae and Mammutidae, and suggests that the evolutions of the Gomphotheriidae and Mammutidae are deeply involved in with each other, rather than straightforwardly detached. This phenomenon has been revealed by a collagen sequence analysis among Notiomastodon, Mammut, and extant elephants, which should be further studied.

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    Nothosaurus luopingensis sp. nov. (Sauropterygia) from the Anisian, Middle Triassic of Luoping, Yunnan Province, China
    SHANG Qing-Hua, LI Chun, WANG Wei
    Vertebrata Palasiatica    2022, 60 (4): 249-270.   DOI: 10.19615/j.cnki.2096-9899.220524
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    Nothosaurus luopingensis sp. nov. from Member II of the Guanling Formation (Anisian, Middle Triassic) of Luoping, Yunnan, China is described based on a specimen comprising the skull and most of the postcranial skeleton. The specimen is assigned to Nothosaurus of Eosauropterygia as suggested by a series of skull characters, such as the maxillary tooth row extending posteriorly beyond the level of the anterior margin of the upper temporal fenestra, the longitudinal diameter of the upper temporal fenestra more than twice as long as that of the orbit, and the presence of maxillary fangs. Compared with Lariosaurus, the following morphological features of the pectoral girdle and the limbs also support the assignment of the specimen to Nothosaurus, i.e., the clavicles with expanded anterolateral corners, the characteristically curved humerus with a straight preaxial angle and a postaxial concavity, the distinct deltopectoral crest on the proximal part of the humerus, no hyperphalangy in the manus, and the absence of pachyostosis in the vertebrae and ribs. On the other hand, the specimen possesses some postcranial features that were previously considered to occur mainly in Lariosaurus, such as more than three ossifications in the carpus, four sacral ribs, and an interclavicle without any trace of a posterior stem. These postcranial characters may no longer be used as the diagnostic features of Lariosaurus. Nothosaurus luopingensis is distinguished from other Nothosaurus species by a unique combination of derived characters, including that the jugal enters the orbit, the nasals are separated, the posterior end of the frontal is bifurcate, pedal digits V and IV are long and subequal in length, and the ungula phalanx is stout. Our phylogenetic analysis corroborates the monophyly of Nothosaurus and suggest that N. luopingensis is the sister group of N. yangjuanensis.

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    Cartilage on the furculae of living birds and the extinct bird Confuciusornis: a preliminary analysis and implications for flight style inferences in Mesozoic birds
    WU Qian, Jingmai K. O’CONNOR, LI Zhi-Heng, Alida M. BAILLEUL
    Vertebrata Palasiatica    2021, 59 (2): 106-124.   DOI: 10.19615/j.cnki.1000-3118.201222
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    The early evolution of flight is one of the most studied topics in vertebrate paleontology. Living birds have evolved to utilize a variety of flight styles, but studies focused on inferring flight strategies in Mesozoic birds are often contradictory and without a clear consensus, making it necessary to find additional informative characteristics that can be useful for inferences in fossils. Virtually nothing is known about the histology of the avian pectoral girdle, even though skeletal and joint tissues are key candidates to solve form-function relationships. Avian secondary cartilage found on the dermal bones of the avian skeleton is influenced by epigenetics and only forms when joints are stimulated by muscle contractions. As the only dermal bone in the avian postcranium, the furcula is a potential site for the formation of furcular secondary cartilage and merits further attention. It is still unknown whether adult living birds and fossil birds have furcular secondary cartilage. Here we present histological analyses conducted on the furcula-coracoid articulation in three living birds (Spilopelia chinensis, the Spotted dove; Passer montanus, the Eurasian tree sparrow; and Apus apus, the Common swift), taxa that utilize different flight styles, and one of the most common fossil birds of the Jehol Biota, Confuciusornis. Secondary cartilage was identified on the furculae of the living birds and of Confuciusornis, representing the first report of furcular secondary cartilage in the fossil record. Clear differences in secondary cartilage morphologies were observed in the living species, but additional data is required to establish a strong form-function relationship that could be useful for making inferences in Mesozoic birds.

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    QIU Zhan-Xiang
    Vertebrata Palasiatica    2002, 40 (04): 317-325.  
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    Chalicothere remains were only occasionally met in late Pliocene-middle Pleistocene deposits of China. Since they were usually very poorly represented, mostly by single teeth, identification was often highly tentative. Nevertheless, they were always identified as belonging to the genus Nestoritherium (formerly Circotherium). The latter was erected by Kaup in 1859 based on Falconer and Cautley’s description of anterior parts of skulls and lower jaws from the Siwaliks, India. On October 14th, 2000, the author had the opportunity to visit the fossil mammal localities in the Tianzhen County, Shanxi, accompanied by Prof. Wei Qi, who found and excavated these localities during the first years of the 1980s. Fortunately enough, some chalicothere fossils were found during the visit at the Loc. 80045, where typical Nihewan fauna were collected by Wei in 1980~ 1981. The new find of the chalicothere fossils tumed out particularly important in clarification of the affinity of the Chinese chalicothere material so far referred to Nestoritherium. The new fossils show clearly that they materially differ from the Siwaliks Nestoritherium and deserve to be classified as a new genus, for which the name Hesperotherium is proposed, alluding to the pre2extinction stage of the Chalicotheiidae. Hesperotherium gen. nov.
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    First record of Saurichthys (Actinopterygii: Saurichthyidae) from the Late Triassic of eastern Paleo-Tethys
    FANG Geng-Yu, SUN Yuan-Lin, JI Cheng, WU Fei-Xiang
    Vertebrata Palasiatica    2023, 61 (1): 1-16.   DOI: 10.19615/j.cnki.2096-9899.221013
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    The saurichthyiform fishes, characterized by a pointed rostrum and a streamlined long and slender body plan, ranked among the top predators of the ichthyofauna in the Early Mesozoic oceanic ecosystem. In a cosmopolitan pattern, these fishes rapidly radiated after the end-Permian mass extinction (EPME) and diversified morphologically and ecologically during the Middle Triassic. Thereafter, they seemingly showed a notable shrinkage from a global distribution to an occurrence basically restricted to the western Paleo-Tethys realm since the Late Triassic. Specifically, there is no saurichthyiform fossil record so far from the marine Late Triassic of South China (eastern Paleo-Tethys), where contrastingly they were highly diversified in stratigraphically older Lagerstätten (Middle Triassic Panxian-Luoping and Xingyi biotas). Here we report the discovery of Saurichthys taotie sp. nov. from the Guanling biota of Guizhou and Yunnan provinces, southwestern China. This new species is a medium-sized Saurichthys featured by subtriangular subopercles ornamented with densely arranged vertical striae, faint ornamentation on the posterior part of the skull roof, and strong longitudinal ridges decorating the anterodorsal surface of the rostrum. By marking its own group’s first occurrence in the Late Triassic of eastern Paleo-Tethyan province, Saurichthys taotie suggests that the saurichthyiform fishes were actually much more widespread than previously thought during that geological stage when they showed a considerable decline in the diversity. By still possessing some features previously only seen in its Early Triassic congeners elsewhere, Saurichthys taotie sheds new light on the evolutionary and paleobiogeographical history of saurichthyiform fishes.

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    Panxianichthys imparilis gen. et sp. nov., a new ionoscopiform (Halecomorphi) from the Middle Triassic of Guizhou, China
    XU Guang-Hui, SHEN Chen-Chen
    Vertebrata Palasiatica    2015, 53 (1): 1-15.  
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    The Ionoscopiformes are a fossil lineage of halecomorphs known only from the Mesozoic marine deposits. Because of their close relationships with the Amiiformes, the Ionoscopiformes are phylogenetically important in investigating the early evolution and biogeography of the Halecomorphi. However, fossil evidence of early ionoscopiforms was scarce; until recently,Robustichthys from the Middle Triassic Luoping Biota, eastern Yunnan,China, represents the oldest and only known ionoscopiform in the Triassic. Here we report the discovery of a new ionoscopiform, Panxianichthys imparilis gen. et sp. nov., on the basis of two well preserved specimens from the Middle Triassic Panxian Biota, western Guizhou, China. The discovery documents the second ionoscopiform in the Middle Triassic; although Panxianichthys is slightly younger than Robustichthys, it is significantly older than other members of this group from the Late Jurassic of Europe, and Early Cretaceous of North and South America. Panxianichthys possesses an important synapomorphy of the Ionoscopiformes: a sensory canal in the maxilla, but retains some primitive characters unknown in other ionoscopiforms. Results of our phylogenetic analysis recover Panxianichthys as the most primitive ionoscopiform, and provide new insight on the early evolution of this clade. The interrelationships of the Ionoscopidae have been reassessed;Quetzalichthysis regarded more closely related to Ionoscopus than to Oshunia. In addition, our analysis supports the reassessment of Furo muensteri as an ophiopsid ionoscopiform. The successive discoveries of Robustichthys andPanxianichthys from China indicate that the early diversification of the Ionoscopiformes is more rapid than previously thought.
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    Xitunaspis, a new eugaleaspid fish (Eugaleaspiformes, Galeaspida) from the Lower Devonian of Qujing, Yunnan
    SUN Hao-Ran, GAI Zhi-Kun, CAI Jia-Chen, LI Qiang, ZHU Min, ZHAO Wen-Jin
    Vertebrata Palasiatica    2022, 60 (3): 169-183.   DOI: 10.19615/j.cnki.2096-9899.220412
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    A new genus and species of the family Eugaleaspidae (Eugaleaspiformes, Galeaspida), Xitunaspis magnus gen. et sp. nov., is described from the Lower Devonian Xitun Formation in Qujing, Yunnan Province, China. The new genus displays the diagnostic characters of the Eugaleaspidae, including a slit-like median dorsal opening, no inner cornual process, developed median dorsal canals, and only three pairs of lateral transverse canals extending from lateral dorsal canals. Different from the other eugaleaspids, X. magnus possesses a large headshield with thick dermal bone and a more plesiomorphic sensory canal system. The phylogenetic analysis of the Galeaspida reveals that Xitunaspis clusters with Dunyu and Eugaleaspis to form a monophyletic clade Eugaleaspidae Liu, 1965, and has a closer relationship with Dunyu than Eugaleaspis by sharing the thick dermal bone of the headshield. The new finding represents the first convincing fossil record of the Eugaleaspiformes in the middle Lochkovian Xitun Formation and adds to our knowledge about the morphology of eugaleaspiforms and the evolutionary pattern of the sensory canal system in the Eugaleaspiformes and even Galeaspida.

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    A new pteranodontoid pterosaur forelimb from the upper Yixian Formation, with a revision of Yixianopterus jingangshanensis
    JIANG Shun-Xing, ZHANG Xin-Jun, CHENG Xin, WANG Xiao-Lin
    Vertebrata Palasiatica    2021, 59 (2): 81-94.   DOI: 10.19615/j.cnki.1000-3118.201124
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    Pterosaurs in the Jehol Biota have been found in the Yixian and Jiufotang formations. The Jingangshan bedding is in the upper part of the Yixian Formation. The first two pterosaur embryos ever discovered in the world, two archaeopterodactyloid specimens, and the questionable Yixianopterus jingangshanensis have been reported in previous literature. Here, we describe a forelimb from this horizon and confirm its phylogenetic position in the Pteranodontoidea. The holotype of Y. jingangshanensis, now housed at Benxi Geological Museum, has been examined. The diagnosis of this taxon has been revised without the consideration of the artificial parts as following, a pteranodontoid pterosaur with a distinguished combination of characters: triangular and labiolingually compressed teeth with the first two more slender and longer than the others; teeth vertical to the occlusal surface; the second wing phalanx about 93% the length of the first wing phalanx. In the Jehol Biota, archaeopterodactyloid specimens have been mainly discovered from the Yixian Formation, while tapejaroids are almost found from the Jiufotang Formation. Including the new forelimb and Y. jingangshanensis, the pteranodontoids from the Jiufotang Formation are slightly greater in number than those from the Yixian Formation in species and specimens, differing from the previous thoughts on the distribution.

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    On kannemeyeriiform dicynodonts from the Shaanbeikannemeyeria Assemblage Zone of the Ordos Basin, China
    LIU Jun
    Vertebrata Palasiatica    2022, 60 (3): 212-248.   DOI: 10.19615/j.cnki.2096-9899.220601
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    Shaanbeikannemeyeria is a common tetrapod from the lower part of the Ermaying Formation of the Ordos Basin, China. There are taxonomical questions surrounding this genus, such as the validity of the genus, and how many species are included within it. Several specimens have been collected since 1978. Shaanbeikannemeyeria first appeared from the top of the Heshanggou Formation. These specimens are described to clarify the diagnostic characters, the individual variations and the phylogenetic position of Shaanbeikannemeyeria. Only one species, S. xilougouensis, is recognized. It is characterized by the following autapomorphies: occiput strongly inclined relative to the palate such that the skull is much shorter basally than dorsally, sword tip-like premaxillary posterodorsal processes, tall and dorsally-convex cutting blade on the medial edge of the dorsal surface of the dentary, reflected lamina with a separated posteroventral process, and 15 dorsal vertebrae. This species shows variations on the cranial morphology, such as the occiput height relative to the width, the snout tip (sharp or obtuse), the shape of the orbital portion of the zygomatic arch, and the shape of caniniform process. Some variations could be ontogenetically related, such as the development of the caniniform process and tusk, the posterior extension of the maxilla on the zygomatic arch, the distance between the frontal and the premaxilla, the intertemporal bar width, and the exposing degree of the parietals. Based on postcranial bones, the second dicynodont genus (possibly Parakannemeyeria) is present in the lower Ermaying Formation.

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    Reexamination of the oldest pigeon (Aves: Columbidae) from Asia: Columba congi from the Early Pleistocene of Zhoukoudian, Beijing, China
    SHEN Wei, Thomas A. STIDHAM, LI Zhi-Heng
    Vertebrata Palasiatica    2021, 59 (3): 245-256.   DOI: 10.19615/j.cnki.1000-3118.210304
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    Columba congi is an extinct species that was described as part of the Early Pleistocene (~1.7 Ma) fauna from locality 12 of the UNESCO Zhoukoudian or “Peking Man” site in Beijing, China. Only four partial humeri of the original type series of 11 bones can be located, and the features present in those specimens do not support the original diagnosis. However, our study and redescription shows that the straight and flat margin of the pneumotricipital fossa rim (in ventral view) and the relative distal position of the dorsal supracondylar tubercle may support the continued recognition of C. congi as a valid extinct species. Columba congi appears to be the oldest fossil of Columba in Asia, and it lived during a warmer and wetter period of time of the Pleistocene with a forested Zhoukoudian. Further study of pigeons from all localities at Zhoukoudian should help to resolve questions about pigeon biogeography and evolution, including possibly the time and center of origin of the globally distributedC. livia.

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    SHANG Qing-Hua, LI Chun
    Vertebrata Palasiatica    2009, 47 (3): 178-193.  
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    A completely articulated ichthyosaur skeleton from the Guanling biota, Guizhou is described. The well preserved postcranial skeleton demonstrates that Guizhouichthyosaurus tangae,a large Triassic ichthyosaurid species previously described fromGuizhou, should be referred to Shastasaurus. Enough materials were found to make possible a satisfactory determination of the systematic position of the large Guanling ichthyosaur species, although both the genus Shastasaurus and the family Shastasauridae have long been hard to define due to the fragmentary nature of the type specimens. The postcranial characters of Shastasaurus tangaeare described in detail based on the new skeleton, the holotype ofGuizhouichthyosaurus tangae and other associated Guanling large ichthyosaur materials. The trunk is very long, with more then 60 presacral vertebrae and a ventrally bent tail. The scapula is broad sickle-shaped. The humerus is anteriorly notched, with a short shaft. The radius is nearly rectangular, with a small notch in the anterior edge, and a very slightly concave posterior edge. The ulna is much smaller than the radius, with a slightly concave anterior edge and bluntly rounded posterior and distal edges. The forefin and hindfin have four principal digits.
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    WANG XiaoLin, ZHOU ZhongHe
    Vertebrata Palasiatica    2003, 41 (01): 34-41.  
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      In this paper we report two new pterodactyloid pterosaurs from the Jiufotang Formation in western Liaoning Province of northeast China. They are Chaoyangopterus zhangi gen. et sp. nov. (Nyctosauridae) and Liaoningopterus gui gen. et sp. nov. (Anhangueridae). The former not only represents the first such record in Asia but also the earliest record and most complete skeleton of the family. Some revisions of the family are made, such as having four wing digits and well-developed manual digits I~III. Liaoningopterus is the largest pterosaur ever discovered in China; its teeth also represent the largest known from any pterosaurs. Discoveries of two more pterosaurs from the Jiufotang Formation further show that two pterosaur assemblages could be recognized in the Jehol Group, represented by the Yixian Formation and Jiufotang Formation, respectively. The pterosaur assemblage of the Jiufotang Formation shows a lot resemblance to that of the Early Cretaceous Santana Formation (Aptian/ Albian) such as the Tapejaridae and Anhangueridae. The age of the Jiufotang Formation (Aptian) is slightly older than the Santana Formation.
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    An Upper Miocene “Hipparion fauna” locality sandwiched by basalt in Hanjiaying, Nei Mongol
    WANG Qian, LIU Yan, WANG Li-Hua, Mikael FORTELIUS, ZHANG Zhao-Qun
    Vertebrata Palasiatica    2021, 59 (2): 125-137.   DOI: 10.19615/j.cnki.1000-3118.210323
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    Here we report a Hipparion fauna locality discovered in between two basalt layers near Hanjiaying Village, Jining District, Wulanchabu City, Nei Mongol. K-Ar isotopic dating of the lower and upper level of the basalt constrains the age of the fauna from 7.2 to 6.8 Ma. Compared with classical Hipparion fauna from northern China, the Hanjiaying fauna is closer to those from Baode of Shanxi, Siziwang Banner of Nei Mongol and the Linxia Basin of Gansu. It is similar to the Loc. 43, 44 and 49 from Baode by faunal composition, confirming their age to be ~7.0 Ma rather than 5.5 Ma. The high similarity with the fossils from Wulanhua, Siziwang Banner, Nei Mongol, verified the age of Wulanhua fauna at about 7 Ma. Compared with faunas from the Linxia Basin, Gansu, it is more similar to those from the upper part of the Liushu Formation, especially the Yangjiashan fauna. Based on the faunal composition and their tooth morphology, the Hanjiaying fauna could be included in the “Gazella dorcadoides” fauna, which is supposed to be at the west paleobiome in northern China during the Late Miocene.

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    LIU Di, ZHOU Zhong-He, ZHANG Yu-Guang
    Vertebrata Palasiatica    2012, 50 (1): 39-52.  
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    Body mass or weight is a crucial biological parameter for an organism. It is influenced by development, reproduction, physiology and evolution. Therefore, mass estimates for fossil species are important for many kinds of analyses. In this project, eighteen bivariate regression analyses of different measurements of the appendicular skeleton plotted against body weight in a data set of 422 individual birds, representing 229 species in 21 orders, revealed high correlations between several skeletal parameters and body mass. R-squared values of eighteen bivariate equations are ranged from 0.50 for tibiotarsal length, indicating a relative poor fit, to 0.91 for humeral diameter. To test the 18 equations empirically, they were used to predict the body weight of an additional 64 extant bird specimens, and the accuracies of various equations were compared. This predictive test showed that three parameters are generally most accurate as predictors of body mass: humerus length, ulna diameter, and tibiotarsal diameter. However, the humeral length and ulna diameter tended to give accurate results for particularly songbirds, raptors and climbing birds. The tibiotarsal diameter tended to give accurate results for terrestrial birds, such like chicken and doves. It is probable that humerus length and ulna diameters are the more accurate parameter for arboreal taxa, while tibiotarsal diameter is more accurate for terrestrial ones. Closer examination of the results showed that different measurements correlated best with body mass in different avian orders. This variation appeared to result from differences in habitat and functional morphology across the avian orders represented in the data set. The weights of some Chinese Mesozoic fossil birds were estimated using the equations generated for humeral length and tibiotarsal diameter, because ulnar diameter was frequently difficult to measure. Humeral length and tibiotarsal diameter yielded dramatically different mass estimates for some taxa, with estimates based on humerus length generally being lower. The result shows that these Early Cretaceous birds experienced a significant diversification in body weight during evolutionary process.
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    A new species of Luganoia (Luganoiidae, Neopterygii) from the Middle Triassic Xingyi Biota, Guizhou, China
    XU Guang-Hui
    Vertebrata Palasiatica    2020, 58 (4): 267-282.   DOI: 10.19615/j.cnki.1000-3118.200624
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    Neopterygii is the largest known group of ray-finned fishes today, and the Luganoiidae, recognized by its specialized skull and greatly deepened flank scales, is a stem lineage of this group in the Middle Triassic. Since its naming in 1939, the Luganoiidae has been represented solely by the marine species Luganoia lepidosteoides found near the Ladinian/Anisian boundary of the southern and eastern Switzerland, northern Italy and southeastern Spain. Here, I report the discovery of a new species of this genus, Luganoia fortuna sp. nov. based on a nearly complete specimen from the late Middle Triassic (Ladinian) marine deposits exposed near Xingyi, Guizhou, China. The discovery represents the first record of the Luganoiidae in Asia, indicating that the biogeographical distribution of this family is much wider than previously recognized. Comparative studies of the new species with the type species from Europe reveal some anatomical features previously unnoticed in Luganoia, e.g., the presence of antorbitals, absence of a plate-like quadratojugal, presence of a narrow naked region of the body at the base of the dorsal fin, presence of a small anal fin closer to the caudal fin than to the pelvic fin, and absence of postcleithra. An amended diagnosis of the genus is presented. Moreover, the distinguishable features between the Chinese and European species are highlighted in this study, and they provide valuable information on the morphological diversification of Luganoia.

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    A cuboid bone of a large Late Miocene elasmothere from Qingyang, Gansu, and its morphological significance
    ZHANG Xiao-Xiao, SUN Dan-Hui
    Vertebrata Palasiatica    2022, 60 (1): 29-41.   DOI: 10.19615/j.cnki.2096-9899.210809
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    A cuboid specimen collected from the Late Miocene stratum at Qingyang, Gansu, China is described here. The size of the Qingyang specimen is comparatively huge, even larger than the average size of Elasmotherium caucasicum collected from Nihewan, Hebei, China. The morphology of the Qingyang specimen is identical to that of other specimens of Elasmotheriini; thus, the Qingyang specimen belongs to a huge elasmothere, most probably Sinotherium. By comparison with extant rhino species, the complex of the main body and the apophysis exhibit functional significance. The angle between the frontal plate of the cuboid and the main axis of the apophysis can suggest the ecological conditions occupied by an elasmothere. From analysis of the morphology of the cuboid, Sinotherium and the more derived elasmotheres probably lived in forested or wooded environments, differing from the previous hypothesis of their paleoenvironment.

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    An overview of Triassic fishes from China
    Jin Fan
    Vertebrata Palasiatica    2006, 44 (01): 28-42.  
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    The Triassic fishes are widely distributed in China. The freshwater ichthyofaunas mainly from northern China are characterized by primitive actinopterygians, and they are closest to those from Siberia and Middle Asia, and also contain some forms similar to the fishes from other regions of Laurasia and the Gondwanan continents. Most of the known fishes from the Ordos Basin are possibly of marine. The marine ichthyofaunas from southern China are characterized by " subholosteans" , and include the modern groups of halecomorphs and the basal forms of teleosts. The Middle-Late Triassic marine fish fauna is much more diversified than the Early Triassic one, and is closely related with the ichthyofaunas in western Tethys. The Lower Yangtze region of South China is probably the cradle of some Triassic fish groups ,e. g. Saurichthyidae. The Triassic fishes in China still await comprehensive investigations.
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    The pelvic morphology of Parayunnanolepis (Placodermi, Antiarcha) revealed by tomographic data
    ZHU You-An, WANG Ya-Jing, QU Qing-Ming, LU Jing, ZHU Min
    Vertebrata Palasiatica    2023, 61 (2): 81-89.   DOI: 10.19615/j.cnki.2096-9899.221126
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    The pelvic morphology, and whether the pelvic fin is present or absent in the earliest jawed vertebrates are key in interpreting the origin of vertebrate paired fins. Parayunnanolepis xitunensis, an antiarch placoderm from the Early Devonian of Yunnan, South China, was previously described to possess the earliest evidence of both dermal and endoskeletal pelvic girdles, presumably for the attachment of the pelvic fins. Here, we redescribe the pelvic region of the holotype based on high-resolution computed tomographic data. Instead of having two large plates previously designated as dermal pelvic girdles, Parayunnanolepis possesses three pairs of lateral pelvic plates, and one large oval median pelvic plate. The paired pelvic plates are flat ventral plates, and differ from other dermal pelvic girdles in lacking a dorsal extension. There is no definitive evidence for the presence of an endoskeletal pelvic girdle in Parayunnanolepis, although the possibility cannot be ruled out. A comparison of the dermal pelvic plates in various jawed stem-gnathostomes suggests the presence of both paired and median pelvic plates is shared by different lineages and might be plesiomorphic. The jawed stem-gnathostomes may have recruited the ventral dermal skeleton of the post-thoracic body into different functional units.

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    Exceptional preservation of an extinct ostrich from the Late Miocene Linxia Basin of China
    LI Zhi-Heng, Alida M. BAILLEUL, Thomas A. STIDHAM, WANG Min, DENG Tao
    Vertebrata Palasiatica    2021, 59 (3): 229-244.   DOI: 10.19615/j.cnki.1000-3118.210309
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    Here we report a new avian fossil from the Late Miocene Linxia Basin, Northwest China, with exceptional soft-tissue preservation. This specimen preserves parts of cervical vertebrae and tracheal rings that are typically ostrich-like, but cannot be diagnosed at the species level. Therefore, the fossil is referred to Struthio sp. The new specimen was preserved in association with a partial skull of Hipparion platyodus. To explore the soft tissue preservation in a fossil deposited in a terrestrial setting, we applied a combination of analytic methods to investigate the microscopic features of the fossilized avian bone. Bacterial alterations (bone bioerosion) were revealed by light microscopy and petrographic sections under SEM imaging. Soft-tissues (fossilized remnants of endogenous blood vessels and red blood cells) were preserved in one demineralized bone fragment and also observed in the in-situ ground-section. These are the first records of soft-tissue preservation in vertebrate remains from the Late Miocene Linxia Basin. Associated geological and sedimentological evidence combined with our new data provide insights into the postmortem taphonomic conditions of this ostrich specimen. A seasonal monsoon might have facilitated the microbial erosion penecontemporaneous with the burial of the specimen. This study encourages interdisciplinary research involving morphology, sedimentology, geochemistry, and histological soft-tissue analyses to better understand the Late Miocene faunal turnovers, climates, and fossil preservation in the Liushu Formation in northwestern China.

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    David W. E. HONE, XU Xing, WANG De-You
    Vertebrata Palasiatica    2010, 48 (1): 19-26.  
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     A single large theropod tooth was recovered from the middle Upper Cretaceous Majiacun Formation, Henan Province, China. The morphology of the tooth strongly suggests that it belongs to a baryonychine spinosaurid theropod based on the following features: long, conical tooth with a sub-circular cross-section, slight recurvature along the length of the tooth and numerous fine denticles on both anterior and posterior carinae. This tooth therefore may represent the first record of the baryonychines in Asia, and the first in the Late Cretaceous. This suggests that the baryonychines were more widely distributed both geographically and temporally than previously thought. Based on the gross morphology and inferred ecology of spinosaurids as a whole, we note that these animals are surprisingly rare in the fossil record compared to other theropod clades. This suggests that in life they may have been genuinely rare animals, perhaps as a result of their extreme morphological specialisation.
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    Development and applications of paleontological computed tomography
    WANG Yan-Fang, WEI Cun-Feng, QUE Jie-Min, ZHANG Wen-Ding, SUN Cui-Li, SHU Yan-Feng, HOU Ye-Mao, ZHANG Jiu-Chang, SHI Rong-Jian, WEI Long
    Vertebrata Palasiatica    2019, 57 (1): 84-92.   DOI: 10.19615/j.cnki.1000-3118.170921
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    The traditional serial grinding method used to investigate the internal structure of fossils cannot be readily applied to valuable fossil specimens due to its destructive and time-consuming nature. Computed tomography (CT) is an ideal non-destructive technique for investigating the internal structure of fossils, in which thousands of serial images are obtained and used to produce an accurate reconstruction of the internal morphology. This paper reviews the design, development and applications of the first CT system in China dedicated exclusively to scanning fossils. The 225 kV three-dimensional (3D) fossil micro-CT (225-3D-μCT) is capable of high-resolution volumetric imaging, with a resolution up to 5 μm, and can accommodate specimens measuring up to 100 mm in diameter and 100 mm in length. The 450 kV ordinary fossil CT (450-TY-ICT) can produce high signal-to-noise ratio (SNR) images of specimens ranging up to 800 mm in diameter and 1000 mm in length, with a resolution up to 200 μm. Two paleontological CT facilities represent a high-performance platform offering the functional diversity needed to meet the demands of studying fossils at a variety of different scales. The two machines have become indispensable for paleontological research in China.

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    Using Bayesian tip-dating method to estimate divergence times and evolutionary rates
    ZHANG Chi
    Vertebrata Palasiatica    2021, 59 (4): 333-341.   DOI: 10.19615/j.cnki.2096-9899.210516
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    Bayesian tip dating is a recently developed method to estimate divergence times and evolutionary rates. It overcomes several drawbacks in traditional stepwise approach. However, it also requires more knowledge about statistics. This paper hierarchically explains the theory and computation in the Bayesian tip-dating approach, and divides the whole process into prior for the divergence times, prior for the evolutionary rates, model for the character changes and Markov chain Monte Carlo algorithm, which are key components in this method. The aim is to provide a general guidance for paleontologists in empirical data analyses.

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    First Middle Devonian galeaspid from the Haikou Formation in Yunnan Province
    MENG Xin-Yuan, Zhu Min, WANG Jun-Qing, PAN Zhao-Hui, GAI Zhi-Kun
    Vertebrata Palasiatica    2022, 60 (3): 184-196.   DOI: 10.19615/j.cnki.2096-9899.220613
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    The Early Emsian Event (E’Em Event), the most significant bio-event for Chinese Devonian vertebrates, has significantly impacted on the diversity of galeaspids. The endemic Galeaspida almost became extinct after the Mid-Emsian Event (M’Em Event). Only few galeaspid taxa survived from these events, such as Clarorbis apponomedianus from the Eifelian of Guangxi, South China, and an indeterminate galeaspid from the Frasnian of Ningxia, Northwest China. Here, we report the first Middle Devonian galeaspid, Dongfangaspis sp., from the Haikou Formation in Wuding, Yunnan Province. The new material is more suggestive of the type species of Dongfangaspis, D. major, than Laxaspis and Polybranchiaspis in its suborbicular headshield with small inner cornual process, broad and nearly aequilate ventral rim, and about 45 pairs of branchial fossae. Dongfangaspis bears the largest number of branchial fossae ever recorded in galeaspids, which probably play an important role in Dongfangaspis surviving from the E’Em and M’Em events. The new finding represents the second Middle Devonian fossil record of galeaspids, and extends the chronological range of Dongfangaspis from the Pragian (Early Devonian) to the Eifelian (Middle Devonian).

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    A small-sized dinocephalosaurid archosauromorph from the Middle Triassic of Yunnan, southwestern China
    WANG Wei, LEI Hong, LI Chun
    Vertebrata Palasiatica    2024, 62 (1): 13-32.   DOI: 10.19615/j.cnki.2096-9899.231013
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    Among numerous marine reptiles discovered in the Triassic eastern Tethys, today’s Southern China, Dinocephalosaurus is a bizarre animal comparable to European Tanystropheus in developing a prominently long neck. These two taxa are respectively assigned to Dinocephalosauridae and Tanystropheidae, and the two families and other basal members collectively form an early-diverging clade of Archosauromorpha. Here we report a new archosauromorph specimen, IVPP V18579, excavated from the lower Middle Triassic (Anisian), from Luoping, Yunnan in southwestern China. Compared with all the hitherto known dinocephalosaurids and tanystropheids, this skeletally mature individual is exclusively similar to Dinocephalosaurus in a number of characteristics, particularly with the long posterodorsal process of the premaxilla extending posteriorly beyond the level of the external nares, the concave posterior margin of the anteroposteriorly broad quadrate, and the strongly expanded distal end of the chevron in most of the caudal vertebrae. However, this reptile is much smaller than Dinocephalosaurus and different from Dinocephalosaurus and the other dinocephalosaurid, Pectodens, in many aspects, such as an anteriorly tapering long rostrum, the dentition composed of short conical teeth with less heterodonty, relatively but obviously tall neural spines of the axis and the anterior cervical vertebrae. Our phylogenetic analysis suggests that the new archosauromorph is a dinocephalosaurid, and then we erect Austronaga minuta gen. et sp. nov. based on this specimen. Detailed comparisons in osteological anatomy and the discussion about its potential aquatic adaptation of this new taxon are also provided.

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    Taxonomic revision of the holotype of Proboselaphus watasei Matsumoto, 1915 (Bovidae, Artiodactyla) from Chuanyu area, China
    NISHIOKA Yuichiro, KOHNO Naoki, KUDO Yuichiro
    Vertebrata Palasiatica    2021, 59 (3): 200-212.   DOI: 10.19615/j.cnki.1000-3118.210322
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    A taxonomic revision of Proboselaphus watasei Matsumoto, 1915 (Bovidae, Artiodactyla) from the Pleistocene of Chuanyu area, China demonstrates that this genus and species names are invalid. The holotype with a skull and mandibles was recently rediscovered in the fossil collection by Nobuo Naora, which is housed in the National Museum of Japanese History, Chiba Prefecture, Japan. Proboselaphus watasei was described as a closely-related species to living nilgai, or Boselaphus tragocamelus, in South Asia, based on general characteristics of pecorans, such as small bony horn-cores and hypsodont cheek teeth. However, the cranial and dental morphologies re-examined in the present study clearly show that the holotype has cervid-specific characteristics: e.g., the fronto-parietal surface curving dorsally, the basioccipital with a triangular outline, and molars with isolated anterior and posterior lobes. The molars of the holotype are comparable to those of Cervus unicolor, in having strong accessary structures (or spurs, cingulums/cingulids, and styles/stylids), and are as large as those ofCervus cf. C. unicolor from the Pleistocene deposits in southern China. This taxonomic change suggests that any crown-boselaphins had not dispersed into East Asia since the Pleistocene.

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    Jin Fan
    Vertebrata Palasiatica    1996, 34 (02): 102-122.  
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    The study of the Late Mesozoic deposits (generally called "Jehol Group" except for the uppermost Sunjiawan Formation) of western Liaoning was initiated early in this century, but the stratigraphic sequence of the Jehol Group, its correlation with the corresponding non-marine strata in northern China and the age assignment have long been controversial issues. The present paper is to deal with these problems mainly based on the data of fossil fishes. The Late Mesozoic lithostratigraphic succession in western Liaoning is adapted as follows (Table l): the Yixian (excluding the former Jingangshan Beds and Tuhulu Beds), Jiufotang, Shahai / Zhanglaogongtun, Fuxin and Sunjiawan formations in ascending order, and certain irregular or diachronous phenomena of lithostratigraphic nomerclature are recognized among the separate basins in this region, e.g., the current Yixian Formation in the western LingyuafrSanshijiazi and middle Chaoyang- KazuoJianchang basins can only becorrelated with the Yixian volcanic rocks and its base sedimentary horizon in the eastern Fuxin-Yixian-Jinzhou Basin, but the Jiufotang Formation in the western and middle basins is equivalent to the Jingangshan and Tuhulu beds plus the Jiufotang Formation in the eastern basin after the litho- and biostratigraphic principles. The igneous rocks over the Yixian Formation has long been ignored or misplaced, and its reasonable position in the sequence should be between the Jiufotang and Fuxin formations. The Jehol Group can be coincidentally correlated with the corresponding strata in other or basins of northern China on the basis of fossil fishes (Table 2). The loweß part (Yixian and Jiufotang formations) of the Jehol Group and its equivalents in northern China were generally originated in fluviolacustrine environment while the upper part (Shahai and Fuxin formations) deposited in deltaic and lacustroswamp facies. The Lycoptera and Kuyangichthys ichthyofaunas (Table 3) show a Late Jurassic and Early Cretaceous aspect, not a few forms are closely related with, •or quite similar to fishes found in the Upper Jurassic of Kazakhstan in central Asia, in the Purbeck-Wea1den of western Europe, and in the Lower Cretaceous of Kitakyushu in Japan. Based on the evidence of the fish faunae, together with the data afforded by other fossil groups, event stratigraphy and radiometric dating etc., the age of the Jehol Group and its corresponding strata in northern China is assigned to late Late Jurassic-mid Early Cretaceous (about Portlandian-Barremian).
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    Climate change and evolution of early lagomorphs (Mammalia): a study perspective based on new materials of Ordolagus from Nei Mongol (northern China)
    Chiara ANGELONE, ZHANG Zhao-Qun
    Vertebrata Palasiatica    2021, 59 (2): 138-168.   DOI: 10.19615/j.cnki.1000-3118.210325
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    The Early Oligocene is a critical time for global climate changes in the Cenozoic. This epoch witnessed severe mammalian faunal turnovers known as “Grande Coupure” in Europe and “Mongolian Remodelling” in Asia. However, insights about morphological changes in Oligocene mammal lineages have not been explored in detail. One of the least diversified groups of recent mammals, lagomorpha, is globally common in the fossil records, especially in Asia. During the Oligocene, many Eocene archaic lagomorph taxa died out and were replaced by more advanced forms. New findings from Nei Mongol and re-examination of the specimens from older collections enabled a revision of a common Asian lagomorph genus, Ordolagus, which possibly has a close affinity with the Middle-Late Eocene genus Gobiolagus. In Nei Mongol, we recognized the presence of Ordolagus during the basal Early Oligocene. Comparisons with coeval and slightly older lagomorph taxa from Asia and North America show that Ordolagus attained some salient tooth morphological characters (i.e., development of anteroconid on p3, full hypselodonty of cheek teeth, and lingual connection of trigonid and talonid on p4-m2), which are also the key features of modern leporids. The appearance of those morphologic features in Ordolagus is coeval to major global or regional climatic changes. Further investigations on Asian early lagomorphs compared with the study of other small mammals and local climatic factors will be essential to refine the role of lagomorphs as palaeoclimatic proxies.

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