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    20 July 2021, Volume 59 Issue 3
    A new species of Pteronisculus from the Middle Triassic (Anisian) of Luoping, Yunnan, China, and phylogenetic relationships of early actinopterygian fishes
    REN Yi, XU Guang-Hui
    2021, 59(3):  169-199.  DOI: 10.19615/j.cnki.2096-9899.210518
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    Actinopterygii, the largest group of extant vertebrates, includes Cladistia, Actinopteri (Chondrostei plus Neopterygii) and closely related fossil taxa. The extinct genus Pteronisculus belongs to a stem lineage of actinopterygian fishes represented by 11 species from the Early Triassic of Madagascar, Europe and North America, and a single species from the early Middle Triassic of China. Here, we report the discovery of a new species of this genus, Pteronisculus changae, on the basis of five well-preserved specimens from the Middle Triassic (Anisian) marine deposits exposed in Luoping, eastern Yunnan, China. The discovery documents the second convincing species of Pteronisculus in the Middle Triassic and the largest stem actinopterygian fish in the Luoping Biota, having a maximum total length of up to 295 mm. The new species possesses a toothed lacrimal, which is characteristic ofPteronisculus, but it is easily distinguished from other species of the genus by some autapomorphies, e.g., a medial process at the middle portion of the intertemporal, 21 supraneurals, and 83 lateral line scales. The results of our cladistic analysis provide new insights into the relationships of early actinopterygians and recover Pteronisculus as a sister taxon of the Carboniferous rhadinichthyid Cyranorhis at the actinopterygian stem. Based on the body form, teeth and other features, it can be deduced that Pteronisculus changae is likely a relatively fast-swimming predator, feeding on planktonic invertebrates and smaller or younger fishes known to occur in the same biota. As one of the youngest species of the genus, the new species provides additional evidence to suggest that the diversity of Pteronisculus is higher than previously thought and that the eastern Paleotethys Ocean likely constituted a refuge for species of this genus during the early Middle Triassic.

    Taxonomic revision of the holotype of Proboselaphus watasei Matsumoto, 1915 (Bovidae, Artiodactyla) from Chuanyu area, China
    NISHIOKA Yuichiro, KOHNO Naoki, KUDO Yuichiro
    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.

    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
    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.

    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
    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.

    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
    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.