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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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    Morphometric analysis of the cervical vertebral series in extant birds with implications for Mesozoic avialan feeding ecology
    LIU Bi-Ying, Thomas A. STIDHAM, WANG Xiao-Ping, LI Zhi-Heng, ZHOU Zhong-He
    Vertebrata Palasiatica    2024, 62 (2): 99-119.   DOI: 10.19615/j.cnki.2096-9899.240305
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    The inference of Mesozoic avialan bird diets previously relied on traditional methods such as morphological comparisons among taxa and direct evidence such as identifiable stomach contents. However, the application of these approaches has been limited because of uncommon preservation of relevant fossil evidence. We searched for additional informative characteristics to help develop new methods to assess the diet of fossil birds. In particular, the morphology of the avialan neck is highly modularized and plays roles in multiple functions including food acquisition. The structure of and variation among the cervical vertebrae likely reflects the demands of feeding ecology in fossil and extant birds because the avialan neck evolved to, at least in part, replace the forelimbs by assisting with activities such as cranioinertial feeding and other ecological functions. Here, we utilize morphometric and statistical analyses to establish an initial quantitative relationship between cervical morphology and dietary modes in both extant and extinct birds. This morphometric framework derived from the cervical morphology of living birds is used as a basis to estimate the diet categories of five taxa of Mesozoic birds. The results indicate that there is a quantitative correlation between cervical morphology differentiation and their interrelated feeding modes. The enantiornithine taxa examined exhibit cervical morphologies similar to extant insectivorous or carnivorous birds. The ornithurine species show cervical morphologies that are more aligned with generalist or herbivorous birds, and exhibit preliminary morphological features tied to aquatic adaptions. These findings are consistent in part with other direct fossil evidence, as well as hypotheses developed from other skeletal comparisons. Therefore, the cervical vertebral series, as a skeletal system closely linked to food acquisition, can serve as one of the valuable metrics to provide information for inferring the diet of long extinct Mesozoic birds.

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    Reappraisal of Bothriolepis sinensis Chi, 1940 from the Tiaomachien Formation, Hunan, China
    LUO Yan-Chao, ZHU Min, LU Li-Wu, PAN Zhao-Hui
    Vertebrata Palasiatica    2023, 61 (4): 261-276.   DOI: 10.19615/j.cnki.2096-9899.230901
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    Bothriolepis sinensis Chi 1940, mainly based on anterior median dorsal plates from the Middle Devonian Tiaomachien Formation of Hunan, is the first Paleozoic vertebrate taxon erected in China. Although additional materials of B. sinensis from the type locality were described by Lu in 1988, its morphology and phylogeny remain poorly understood. In this study, we complemented the morphology of the skull and trunk armor of B. sinensis based on Chi’s specimens housed in the Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, and several previously undescribed specimens in the Geological Museum of China. Bothriolepis sinensis differs from other Bothriolepis in the following combination of characteristics: enlarged supraotic thickening, length/width ratio of head shield 1.4-1.6, broad orbital fenestra (greater than 1/3 of the head shield width), and fan-shaped preorbital recess. The phylogenetic analysis did not place B. askinae in the most basal position of the genus and revealed that B. sinensis and B. kwangtungensis consistently from a monophyletic group characterized by their slender proximal segment of the pectoral fin (length/width ratio greater than 7). A majority of Chinese Bothriolepis species (B. niushoushanensis , B. lochangensis , B. tungseni , B. kwangtungensis and B. sinensis ) were clustered in a clade characterized by the pectoral pit-line on the ventral central plate 1 extending to the ventral central plate 2. The paleogeographic reconstruction using the data from the DeepBone platform showed that Bothriolepis had its oldest occurrences in South China and East Gondwana in Eifelian, dispersed rapidly worldwide, and then diversified across the coasts of the Rheic Ocean.

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    A giant bamboo rat from the latest Miocene of Yunnan
    Lawrence J. FLYNN, LI Qiang, Jay KELLEY, Nina G. JABLONSKI, JI Xue-Ping, Denise F. SU, WANG Xiao-Ming
    Vertebrata Palasiatica    2023, 61 (4): 277-283.   DOI: 10.19615/j.cnki.2096-9899.230710
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    The Shuitangba subbasin lignite deposits of the Zhaotong Basin in northern Yunnan Province have produced vertebrate fossils of terminal Miocene age. We conducted test wet screening of fossiliferous sediment in 2014 to increase representation of small mammals. This effort produced four teeth of a very large bamboo rat, much larger than the previously known bamboo rat present at Shuitangba, and representing a new species. This new species is characterized by its molars being remarkably larger than those of other known species of Miorhizomys , and being hypsodont with cementum, and less anterorposteriorly compressed. The age of this new species from Shuitangba is in the range of 6.2 to 6.7 Ma. It appears that diverse bamboo rats of the extinct genus Miorhizomys were present in the Late Miocene of Yunnan, somewhat before the 6 Ma appearance of extant Rhizomys to the north in the vicinity of Shanxi Province.

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    New findings of Xiyuichthys (Xiushuiaspidae, Galeaspida) from the Silurian of Jiangxi Province and Tarim Basin
    SHAN Xian-Ren, LIN Xiang-Hong, ZHANG Yu-Meng, LI Xu-Tong, GAI Zhi-Kun
    Vertebrata Palasiatica    2023, 61 (4): 245-260.   DOI: 10.19615/j.cnki.2096-9899.230904
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    New findings of the early Silurian Xiyuichthys (Xiushuiaspidae, Galeaspida), Xiyuichthys lixiensis sp. nov. and X. zhangi are described from the Qingshui Formation in Jiangxi Province and the Tataertag Formation in Tarim Basin respectively. X. lixiensis sp. nov. is characterized by the partially serrated lateral margin of the headshield and the ornamentation composed of extremely coarse granular tubercles (one tubercle per square millimetre). The complete early Silurian biostratigraphic sequence in northwestern Jiangxi warrants the erection of a standard section for the correlation of Silurian shallow marine red beds in South China and Tarim blocks. Thus, the finding of X. lixiensis from the Qingshui Formation (Silurian Lower Red Beds) in Jiujiang of Jiangxi bears important biostratigraphic significance. It can directly compare to X. zhangi from the Tataertag Formation in Tarim Basin on the specific level, which corroborates the correlations between the Tataertag Formation in Tarim Block and the Silurian Lower Red Beds in South China. Fossil records suggest that Xiushuiaspidae have a relatively broader stratigraphic range, but exhibit distinct composition at different stratigraphic horizons, with Xiyuichthys and Changxingaspis arising in the Silurian Lower Red Beds (Qingshui, Tangchiawu, and Tataertag formations) and Xiushuiaspis occurring in the Silurian Upper Red Beds (Xikeng Formation).

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    An egg clutch of the Stalicoolithidae discovered in Wuning, Jiangxi, China
    ZHOU Ming-Xiao, YAN Yun, QIU Wen-Jiang, FANG Kai-Yong, ZHU Xu-Feng, WANG Qiang, WANG Xiao-Lin
    Vertebrata Palasiatica    2023, 61 (4): 317-325.   DOI: 10.19615/j.cnki.2096-9899.230519
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    Here we describe an incomplete dinosaur clutch with three broken eggs and seven prints discovered in Wuning County, Jiangxi Province, that can be referred to Coralloidoolithus shizuiwanensis based on the following features: the eggs are nearly spheroid and arranged tightly and irregularly in the clutch, the eggshell thickness ranges 2.76-2.97 mm, the horizontal accretion lines are almost evenly distributed throughout the eggshell, and the secondary eggshell units are distributed in the medial and outer zones of the columnar layer. This egg clutch of Coralloidoolithus shizuiwanensis represents the first discovery of dinosaur eggs in Wuning County, and shows the age of the strata containing the dinosaur eggs in this area should be Late Cretaceous.

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    A skull of Early Pleistocene Paracamelus gigas (Artiodactyla, Mammalia) from Luotuo Hill in Dalian, Northeast China
    DONG Wei, LIU Wen-Hui, BAI Wei-Peng, LIU Si-Zhao, WANG Yuan, LIU Jin-Yuan, JIN Chang-Zhu
    Vertebrata Palasiatica    2024, 62 (1): 47-68.   DOI: 10.19615/j.cnki.2096-9899.230616
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    Originated in North America in the Middle Eocene, camelids were a successful group with very large diversity. But the camels emigrated to the Old World from North America, probably during the middle stage of the Middle Miocene, and did not radiate much as those in North America, represented by only two genera Paracamelus and Camelus. The former was considered as giving rise to the latter, but the detailed relationship of the Old World camelines was controversial. The new camel material unearthed from Layer 4 in the Jinyuan Cave at Luotuo Hill in Dalian, Liaodong peninsula in Northeast China, was described and referred to as Paracamelus gigas. Its dentition length is slightly longer than that of Camelus knoblochi but evidently larger than that of C. ferus and C. dromedarius. Based on the fossil records and morphometric evidences, P. gigas originated from a form similar to P. alexejevi in the Late Pliocene in the Old World, instead of from Megatylopus gigas of North America and then migrated into Asia as previously thought. The morphometric similarities between the Early Pleistocene Dalian specimens and those of the Middle and Late Pleistocene C. knoblochi indicate that P. gigas probably gave rise to C. knoblochi as formerly postulated and likely in the late Early Pleistocene by reduction or simplifying of P3 and P4, disappearance of p3 and shortening of dentition length. P. gigas inhabited in the forest steppe environment of Liaodong peninsula from 1.1 to 1.52 Ma based on paleomagnetic dating and pollen evidence.

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    A possible new amphicyonid from the Miocene of the Linxia Basin
    JIANGZUO Qi-Gao, GAO Yuan, Alberto VALENCIANO, LU Dan, WANG Shi-Qi
    Vertebrata Palasiatica    2024, 62 (2): 156-164.   DOI: 10.19615/j.cnki.2096-9899.240320
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    Here we report a new form of amphicyonid from an uncertain locality in the Linxia Basin. The derived dental traits imply an affinity to Magericyon, previously known from Europe and possibly southern Asia. The specimen suggests a higher diversity of amphicyonids in eastern Asia than previously thought, and more discovery with stratigraphic information will be needed to elucidate the evolution of Amphicyonidae in eastern Asia.

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    Decline in phylogenetic diversity of Arthrodira (stem-group Gnathostomata) correlates with major Devonian bioevents
    XUE Qin-Yuan, YU Yi-Lun, PAN Zhao-Hui, ZHU You-An, ZHU Min
    Vertebrata Palasiatica    2024, 62 (1): 1-12.   DOI: 10.19615/j.cnki.2096-9899.231124
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    Arthrodira, the most diverse subgroup of placoderms, or jawed stem gnathostomes, is noted for their occupation of high trophic levels, especially in the later stage of their evolutionary history. Despite the relatively complete fossil record of arthrodires, the results of previous studies on the arthrodire diversity, based on counting the numbers of taxa in each time interval (raw or taxic diversity), correlate poorly with major Devonian bioevents. Here, we assemble a new, exhaustive dataset of arthrodires, comprising 450 species of 219 genera. Most taxa are integrated into a supertree integrating the results from various phylogenetic investigations. Our analysis of the phylogenetic diversity, accounting for the presence of ghost lineages, reveals a very different pattern compared to the raw diversity. The phylogenetic diversity of arthrodires exhibited a typical early burst pattern, peaking in the Early Devonian (Lochkovian-Pragian Boundary), and followed by declines that aligned well with several major bioevents. Near each of the first four events, the arthrodire diversity experienced only minor drops and generally persisted at high levels. The later three events, particularly the Frasnian-Famennian Boundary and Devonian-Carboniferous Boundary Events, led to steep declines in arthrodire diversity, from which they never recovered before their complete extinction in the end-Devonian. All these declines were not evident in the raw or taxic diversity pattern, except that related to the Frasnian-Famennian Boundary Event.

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    A taxonomical revision of ‘Dongfangaspis qujingensis’ from the Lower Devonian of Qujing, Yunnan Province
    SHAN Xian-Ren, ZHU Min, LI Qiang, GAI Zhi-Kun
    Vertebrata Palasiatica    2024, 62 (2): 85-98.   DOI: 10.19615/j.cnki.2096-9899.240321
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    The affinity of ‘Dongfangaspis qujingensis’, initially assigned to Dongfangaspis but later to Laxaspis, has long been controversial. However, the taxonomical revision raises a new problem of junior homonym since the type species of Laxaspis is L. qujingensis. Here, we describe some new materials of ‘Dongfangaspis qujingensis’ and Damaspis vartus from the Xishancun Formation (early Lochkovian, Early Devonian) in Qujing, Yunnan Province. ‘Dongfangaspis qujingensis’ strikingly resembles Damaspis vartus in the slightly longer headshield, bifurcated ends of the lateral transverse canals, unconnected V-shaped posterior supraorbital canals, and at least seven pairs of lateral transverse canals issuing from the lateral dorsal canal. These similarities indicate that ‘D. qujingensis’ is more suggestive of Damaspis than Dongfangaspis and Laxaspis. Therefore, we propose to remove ‘Dongfangaspis qujingensis’ from Laxaspis to Damaspis. The new specimens of Damaspis vartus reveal five long lateral transverse canals on the right side, corroborating that the asymmetric sensory canal system in the holotype is the intraspecific variation.

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    First report of Hispanodorcas from the Late Miocene of China
    WU Yong, WANG Shi-Qi, LIANG Zhi-Yong, GUO Ding-Ge, SUN Bo-Yang, LIU Long, DUAN Kai, CHEN Guo-Zhong
    Vertebrata Palasiatica    2024, 62 (2): 135-155.   DOI: 10.19615/j.cnki.2096-9899.240123
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    As a small to middle-sized bovid, Hispanodorcas had previously only been found in the pan-Mediterranean region and South Asia. Its taxonomic classification at the tribe level has been a subject of debate, with possible associations to Antilopini, Reduncini, or Oiocerini. Here, we report on the first discovery of Hispanodorcas in East Asia, H. longdongica sp. nov. from the Daidian Locality in China, dating to the early Baodean age (~8-7 Ma). The new material consists of five skulls with varying states of preservation and provides the most complete osteological information on Hispanodorcas to date. It features a long, slender, and posteriorly curved horncore with a weak homonymous twist and both laterodorsal and medioventral grooves, which is characteristic of Hispanodorcas. This new species is characterized by having the smallest size amongst all known Hispanodorcas species, a weakly curved brain case in the facial region, and poorly developed posterior and anterior basilar tuberosities. These primitive characteristics suggest that H. longdongica may represent an early evolutionary stage of this genus. Furthermore, they indicate that Hispanodorcas might have directly evolved from the Gazella stock. The homonymous twist in the horncore, which aligns with Oiocerini, may be a case of homoplasy.

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    Reassessment of Trilophodon connexus Hopwood, 1935 and attributing it to the Choerolophodontidae
    LI Chun-Xiao, CHEN Jin, WANG Shi-Qi
    Vertebrata Palasiatica    2024, 62 (1): 33-46.   DOI: 10.19615/j.cnki.2096-9899.230917
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    Trilophodon connexus Hopwood, 1935 has long been considered a typical species of Gomphotherium in China. However, due to the unknown state of the mandibular symphysis and tusks, there is no definite evidence to assign “T. connexus” to Gomphotherium. Here we describe and reevaluate a hemimandible from the Halamagai Formation, Ulungur region, northern Junggar Basin, which was previously identified as Gomphotherium cf. G. shensiensis. The mandibular symphysis is deeply troughed and lacks mandibular tusks; therefore, it undoubtedly belongs to the Choerolophodontidae. Further comparison revealed that the cheek tooth morphology is identical to that of the type specimen of Trilophodon connexus. The characteristic features include high bunodonty, elongation of the m3 with four lophids, an only weakly chevroned lophid 2, enlargement of the posterior pretrite central conule 2, unfused state of the pretrite mesoconelet 2 (if present) and anterior pretrite central conule 2, as well as the absence of ptychodonty, choerodonty, and cementodonty. Therefore, T. connexus Hopwood, 1935 is a choerolophodontid rather than a species of Gomphotherium. Based on the above features, we provisionally refer to it as “Choerolophodonconnexus. “Choerolophodonconnexus is characterized by the following features: weak or absent ptychodonty, choerodonty, and loph chevron (which were all strong in the typical species of Choerolophodon), as well as multiplication of the lophids in the m3, which were similar to that of the North American Gnathabelodon. Therefore, Gnathabelodon might represent a distinct lineage within the Choerolophodontidae, and may be derived from the East Asian “Choerolophodonconnexus.

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    Micromammal fossils from the basal part of the Jiaozigou Formation in Yagou area, Linxia Basin, Gansu Province
    WANG Ban-Yue, QIU Zhan-Xiang
    Vertebrata Palasiatica    2023, 61 (4): 284-316.   DOI: 10.19615/j.cnki.2096-9899.230927
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    Renewed fieldwork in 2003 produced a rich micromammal assemblage from the basal part of the Jiaozigou Formation in the Yagou area of Linxia Basin. 17 genera distributed among 13 families of three orders were recovered. The micromammal fauna is a typical Oligocene assemblage for Central and Eastern Asia. 14 genera (~82% of the fauna) are common in the Oligocene of Asia. Of them four genera and four species of Eucricetodon are restricted to the Oligocene, one genus (Bagacricetodon ) is restricted to Late Oligocene and Glis and Eomyodon made their first appearances in the Late Oligocene. Based on this micromammal composition, the basal part of the Jiaozigou Formation in the Yagou area could be mainly of Late Oligocene in age, which is in accordance with the conclusion based on large mammal fossils. In comparison with the other Late Oligocene micromammal faunas in Central and East Asia, the Yagou Fauna is slightly older than the Ulan III biozone of Nei Mongol and biozone C of Mongolia, because it has two Eocene genera and lacks more advanced genera. This is roughly in accordance with the recent palaeomagnetic interpretation for the Maogou section, where the lower boundary of the Jiaozigou Formation was correlated with Chron C10r (~29 Ma). The presence of large number of xerophilous zapodines, ctenodactylids, cricetids and lagomorphs combined with fossorial Tsaganomys and the lithology of the fossil-bearing deposits, composed of gypsiferous reddish brown mudstone, tend to show a semiarid woodland-shrubland habitat during the Late Oligocene in Yagou area. In Late Oligocene more frequent faunal interchange might have occurred between Asia and Europe (4 genera commonly shared) rather than between Asia and America (only 1 genus shared), partly because of the disappearance of the Turgai Strait.

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    First histochemical examination of a Miocene ostrich eggshell with the oldest mineral-bound peptides
    WU Qian, PAN Yan-Hong, LI Zhi-Heng, ZHOU Zhong-He, Alida M. BAILLEUL
    Vertebrata Palasiatica    2024, 62 (2): 120-134.   DOI: 10.19615/j.cnki.2096-9899.240329
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    Because ancient proteins have a higher preservation potential than ancient DNA, proteomic studies can help shed light on the biology of some extinct biological groups that are beyond the reach of the field of ancient DNA. The oldest peptide discovered so far is part of the protein struthiocalcin (SCA-1) involved in eggshell mineralization and found within an ostrich egg from the Late Miocene Linxia Basin of Northwest China. It was originally hypothesized that SCA-1 was evenly distributed within the eggshell and was able to enter the fossil record for so long, because it was bound to calcite crystals. We conducted histological, scanning electron microscopy and Raman spectroscopic analyses on this same fossil egg to test if any protein or organic matter could be observed within specific regions of the eggshell and indeed bound to calcite crystals. Our results show that the eggshell is made entirely of calcite except at the base layer, which is made of mammillary knobs at least partially made of apatite. These knobs were secondarily phosphatized during diagenesis. After decalcification of this material, the fossilized mammillary knobs showed fibrous residues consistent in location and morphology with remnants of original organic material forming a network. This network was similar to the organic matrix observed in an extant ostrich eggshell with this same method. The results here suggest that SCA-1 may have been concentrated at the mammillary knobs, rather than evenly throughout the eggshell. Phosphatization may be another taphonomic process that favors organic preservation in deep-time. The paleoclimate and taphonomic environment of the Linxia Basin may have provided favorable conditions for the molecular preservation of this egg. More in-depth histochemical and mineralogical analyses will certainly increase our understanding of organic and ancient protein preservation in this basin.

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    New suoid remains (Mammalia, Artiodactyla) from the Late Miocene of Haritalyangar, India
    Vertebrata Palasiatica    2024, 62 (1): 69-84.   DOI: 10.19615/j.cnki.2096-9899.231120
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    We report here a number of mandibular, maxillary and dental fossil remains of Indian Suoidea from the Middle Siwalik of Haritalyangar area in Bilaspur District of Himachal Pradesh, North India. Haritalyangar is well known for the diversity of the Late Miocene fauna and the hominoids and other primates. The suoid remains were collected by one of the authors (ARS) during different field seasons and their localities were plotted on the map, along with those of the proboscidean reported recently, including the hominoid localities. The fossil localities are spread over the ‘Lower Alternations’ and the ‘Upper Alternations’ from Makkan Khad to Sir Khad. We have assigned the fossils into three genera, Propotamochoerus (P. hysudricus), Hippopotamodon (H. sivalense) and Yunnanochoerus (Y. dangari). Propotamochoerus hysudricus represents the most common suid in the Middle Siwaliks. The new remains of the suoid Yunnanochoerus dangari further documents this rare palaeochoerid only known in the Haritalyangar area by a few fossils. The new suoid remains show clear affinities with the Nagri fauna of the Pakistan Siwaliks. Biochronological correlations with the Potwar Plateau based on suoids indicate an age bracket of ~10-9 Ma for the ‘Lower Alternations’ of Haritalyangar, close to the bracket mostly recently proposed on the basis of magnetic polarity stratigraphy.

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