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New fossils of small and medium-sized bovids from the Early Site of Shanshenmiaozui in Nihewan Basin, North China
TONG Hao-Wen, ZHANG Bei, CHEN Xi, WANG Xiao-Min
Vertebrata Palasiatica    2022, 60 (2): 134-168.   DOI: 10.19615/j.cnki.2096-9899.220413
Abstract   (181 HTML34 PDF(pc) (11536KB)(259)  

Shanshenmiaozui site in Nihewan Basin in North China is a recently discovered Early Pleistocene site which yields rich and diverse mammalian fossils. In the fauna, the small and medium-sized bovid fossils are well represented and can be referred to the following taxa: Spirocerus wongi, Gazella sinensis, Ovis shantungensis and Megalovis piveteaui respectively, among which G. sinensis is the dominate species. S. wongi and G. sinensis are mainly represented by horn-cores and partial skull bones as well as mandibles; in addition, metacarpal and/or metatarsal bones were also recognized for all of the four species. The horn-cores are easy to be identified to the species level, while the dentitions and the postcranial bones underwent a series of examinations and comparisons before getting properly determined and referred to the most approximate taxa. Among the postcranial bones, the metapodials, especially to the metacarpal bones special attentions were paid, which are crucial not only for taxonomic identification, but also for phylogenetic and paleoecological reconstructions; the previously misidentified metapodial specimens in Nihewan fauna were reconsidered in this paper. In the SSMZ fauna, the bovid guild is dominated by Gazella and Bison, which indicates steppe was the most important biome in Nihewan Basin during Early Pleistocene.


Fig. 1 Location of Shanshenmiaozui (SSMZ) site, with distributional map of living antelopes and related fossil forms of the Pleistocene Epoch in China
Data for the extant taxa are from Jiang ( 2004 ), Smith and Xie ( 2008 ). Fossil localities:
1. Xiashagou of Nihewan; 2. Yushe; 3. Dongdoubi of Yuxian; 4. Jinyuan Cave; 5. Zhoukoudian Loc.1;6. Xujiayao; 7. Mashandong; 8. Salawusu; 9. Xifeng of Qingyang; 10. Loufangzi; 11. Jinniushan;12. Miaohoushan; 13. Haimao of Dalian; 14. Gulongshan; 15. Xiaogushan; 16. Chicheng; 17. Dingcun;18. Xihoudu; 19. Tianshuigou of Dali; 20. Wenxi; 21. Pinglu; 22. Danangou of Yuxian; 23. Bajiazui;24. Gonghe; 25. Yangguo; 26. Tuozidong; 27. Tunliu; 28. Banpo; 29. Linyi; 30. Yuanmou; 31. Heshui;32. Xingtai; 33. Xiaochangcun; 34. Wuwang of Linyi; 35. Gengjiagou; 36. Luhuo; 37. Zhoukoudian Loc.15;38. Upper Cave; 39. Guxiangtun; 40. Zhoujiayoufang; 41. Dali Man site; 42. Aba; 43. Chifeng;44. Rouyuan; 45. Yanjiagang; 46. Yuhongcun of Dali; 47. Lingjing
Extracts from the Article
In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021), Ochotonoides complicidens (Tong et al., 2021), Ochotona youngi (Tong et al., 2021), Nyctereutes sp., Canis chihliensis (Tong et al., 2012, 2021), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012; Tong and Chen, 2016; Chen and Tong, 2017; Tong et al., 2021), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014), Elasmotherium peii (Tong et al., 2014, 2018), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021), Nipponicervus elegans (Tong et al., 2021), Eucladoceros boulei (Tong and Zhang, 2019), Elaphurus bifurcatus (Tong et al., 2021), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna.
Gazelles are adapted to open and arid environment, therefore, their range, both fossil and extant, only limited to the northern part in China, and the southernmost occurrences are Xuchang Man site at Lingjing in Henan Province (Dong and Li, 2008) and the Tuozi Cave in Jiangsu Province (Dong et al., 2007) ( Fig. 1). As to the “ Gazella kaihuaensis” from Zhejiang Province (Jin et al., 1994), it’s an incorrect identification because of the following evidences: larger size, lack of a pedicle, horn far away from orbit, horn-cores with smooth surface rather than with longitudinal grooves, supraorbital foramen too small, small angle between horn-core and cranial roof, none of which coincides with the characters of a Gazella. The partial skull bone described by Jin et al. (1994) should belong to a serow ( Capricornis).
# Calculated by the first author of this paper; mean values in parentheses. ...
Late Pleistocene Artiodactyla (Mammalia) from the Lingjing site, Xuchang, Henan Province (China)
1
2008
... Gazelles are adapted to open and arid environment, therefore, their range, both fossil and extant, only limited to the northern part in China, and the southernmost occurrences are Xuchang Man site at Lingjing in Henan Province (Dong and Li, 2008 ) and the Tuozi Cave in Jiangsu Province (Dong et al., 2007 ) ( Fig. 1 ). As to the “ Gazella kaihuaensis” from Zhejiang Province (Jin et al., 1994 ), it’s an incorrect identification because of the following evidences: larger size, lack of a pedicle, horn far away from orbit, horn-cores with smooth surface rather than with longitudinal grooves, supraorbital foramen too small, small angle between horn-core and cranial roof, none of which coincides with the characters of a Gazella. The partial skull bone described by Jin et al. ( 1994 ) should belong to a serow ( Capricornis). ...
1
2007
... Gazelles are adapted to open and arid environment, therefore, their range, both fossil and extant, only limited to the northern part in China, and the southernmost occurrences are Xuchang Man site at Lingjing in Henan Province (Dong and Li, 2008 ) and the Tuozi Cave in Jiangsu Province (Dong et al., 2007 ) ( Fig. 1 ). As to the “ Gazella kaihuaensis” from Zhejiang Province (Jin et al., 1994 ), it’s an incorrect identification because of the following evidences: larger size, lack of a pedicle, horn far away from orbit, horn-cores with smooth surface rather than with longitudinal grooves, supraorbital foramen too small, small angle between horn-core and cranial roof, none of which coincides with the characters of a Gazella. The partial skull bone described by Jin et al. ( 1994 ) should belong to a serow ( Capricornis). ...
Late Pleistocene mammalian fauna from the Mashandong, Chaoyang, Liaoning Province
3
2009
... Measurements of the horn-cores of Spirocerus wongi, compared with related species (mm)
# Calculated by the first author of this paper; mean values in parentheses. ...
New remains of Mammuthus trogontherii from the Early Pleistocene Nihewan beds at Shanshenmiaozui, Hebei
1
2012
... In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1 ) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021 ), Ochotonoides complicidens (Tong et al., 2021 ), Ochotona youngi (Tong et al., 2021 ), Nyctereutes sp., Canis chihliensis (Tong et al., 2012 , 2021 ), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012 ; Tong and Chen, 2016 ; Chen and Tong, 2017 ; Tong et al., 2021 ), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014 ), Elasmotherium peii (Tong et al., 2014 , 2018 ), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021 ), Nipponicervus elegans (Tong et al., 2021 ), Eucladoceros boulei (Tong and Zhang, 2019 ), Elaphurus bifurcatus (Tong et al., 2021 ), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017 ). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna. ...
On newborn calf skulls of Early Pleistocene Mammuthus trogontherii from Shanshenmiaozui in Nihewan Basin, China
1
2016
... In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1 ) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021 ), Ochotonoides complicidens (Tong et al., 2021 ), Ochotona youngi (Tong et al., 2021 ), Nyctereutes sp., Canis chihliensis (Tong et al., 2012 , 2021 ), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012 ; Tong and Chen, 2016 ; Chen and Tong, 2017 ; Tong et al., 2021 ), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014 ), Elasmotherium peii (Tong et al., 2014 , 2018 ), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021 ), Nipponicervus elegans (Tong et al., 2021 ), Eucladoceros boulei (Tong and Zhang, 2019 ), Elaphurus bifurcatus (Tong et al., 2021 ), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017 ). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna. ...
Juvenile skulls and other postcranial bones of Coelodonta nihowanensis from Shanshenmiaozui, Nihewan Basin, China
1
2014
... In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1 ) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021 ), Ochotonoides complicidens (Tong et al., 2021 ), Ochotona youngi (Tong et al., 2021 ), Nyctereutes sp., Canis chihliensis (Tong et al., 2012 , 2021 ), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012 ; Tong and Chen, 2016 ; Chen and Tong, 2017 ; Tong et al., 2021 ), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014 ), Elasmotherium peii (Tong et al., 2014 , 2018 ), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021 ), Nipponicervus elegans (Tong et al., 2021 ), Eucladoceros boulei (Tong and Zhang, 2019 ), Elaphurus bifurcatus (Tong et al., 2021 ), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017 ). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna. ...
New fossils of Eucladoceros boulei (Artiodactyla, Mammalia) from Early Pleistocene Nihewan Beds, China
1
2019
... In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1 ) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021 ), Ochotonoides complicidens (Tong et al., 2021 ), Ochotona youngi (Tong et al., 2021 ), Nyctereutes sp., Canis chihliensis (Tong et al., 2012 , 2021 ), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012 ; Tong and Chen, 2016 ; Chen and Tong, 2017 ; Tong et al., 2021 ), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014 ), Elasmotherium peii (Tong et al., 2014 , 2018 ), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021 ), Nipponicervus elegans (Tong et al., 2021 ), Eucladoceros boulei (Tong and Zhang, 2019 ), Elaphurus bifurcatus (Tong et al., 2021 ), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017 ). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna. ...
A preliminary report on the excavations at the Early Pleistocene fossil site of Shanshenmiaozui in Nihewan Basin, Hebei, China
1
2011
... The SSMZ site lies at the neighboring hill of Xiaochangliang, a well-known Paleolithic Site in Nihewan Basin. Based on the stratigraphic correlation in the field, the fossil-bearing sand-silt bed at SSMZ site is a little higher than the cultural layer at XCL site (Tong et al., 2011 ; Liu et al., 2012 ), whose paleomagnetic age is about 1.36 Ma BP (Zhu et al., 2001 ). On the other hand, the SSMZ fauna is very similar with the classical Nihewan fauna in faunal composition, which means the SSMZ fauna should have an approximate age as the latter based on faunal correlation (Tong et al., 2021 ), whose recent dating result is 2.2-1.7 Ma (Liu et al., 2012 ). ...
New remains of Canis chihliensis (Mammalia, Carnivora) from Shanshenmiaozui, a Lower Pleistocene Site in Yangyuan, Hebei
1
2012
... In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1 ) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021 ), Ochotonoides complicidens (Tong et al., 2021 ), Ochotona youngi (Tong et al., 2021 ), Nyctereutes sp., Canis chihliensis (Tong et al., 2012 , 2021 ), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012 ; Tong and Chen, 2016 ; Chen and Tong, 2017 ; Tong et al., 2021 ), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014 ), Elasmotherium peii (Tong et al., 2014 , 2018 ), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021 ), Nipponicervus elegans (Tong et al., 2021 ), Eucladoceros boulei (Tong and Zhang, 2019 ), Elaphurus bifurcatus (Tong et al., 2021 ), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017 ). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna. ...
New fossils of Stephanorhinus kirchbergensis and Elasmotherium peii from the Nihewan Basin
1
2014
... In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1 ) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021 ), Ochotonoides complicidens (Tong et al., 2021 ), Ochotona youngi (Tong et al., 2021 ), Nyctereutes sp., Canis chihliensis (Tong et al., 2012 , 2021 ), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012 ; Tong and Chen, 2016 ; Chen and Tong, 2017 ; Tong et al., 2021 ), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014 ), Elasmotherium peii (Tong et al., 2014 , 2018 ), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021 ), Nipponicervus elegans (Tong et al., 2021 ), Eucladoceros boulei (Tong and Zhang, 2019 ), Elaphurus bifurcatus (Tong et al., 2021 ), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017 ). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna. ...
New fossils of Bison palaeosinensis (Artiodactyla, Mammalia) from the steppe mammoth site of Early Pleistocene in Nihewan Basin, China
1
2017
... In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1 ) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021 ), Ochotonoides complicidens (Tong et al., 2021 ), Ochotona youngi (Tong et al., 2021 ), Nyctereutes sp., Canis chihliensis (Tong et al., 2012 , 2021 ), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012 ; Tong and Chen, 2016 ; Chen and Tong, 2017 ; Tong et al., 2021 ), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014 ), Elasmotherium peii (Tong et al., 2014 , 2018 ), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021 ), Nipponicervus elegans (Tong et al., 2021 ), Eucladoceros boulei (Tong and Zhang, 2019 ), Elaphurus bifurcatus (Tong et al., 2021 ), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017 ). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna. ...
New postcranial bones of Elasmotherium peii from Shanshenmiaozui in Nihewan Basin, northern China
2
2018
... In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1 ) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021 ), Ochotonoides complicidens (Tong et al., 2021 ), Ochotona youngi (Tong et al., 2021 ), Nyctereutes sp., Canis chihliensis (Tong et al., 2012 , 2021 ), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012 ; Tong and Chen, 2016 ; Chen and Tong, 2017 ; Tong et al., 2021 ), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014 ), Elasmotherium peii (Tong et al., 2014 , 2018 ), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021 ), Nipponicervus elegans (Tong et al., 2021 ), Eucladoceros boulei (Tong and Zhang, 2019 ), Elaphurus bifurcatus (Tong et al., 2021 ), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017 ). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna. ...

# Calculated by the first author of this paper; mean values in parentheses. ...
Chronological significance of the mammalian fauna from the Early Pleistocene Shanshenmiaozui site in Nihewan Basin, northern China
10
2021
... In the past decades, much more explorations and excavations have been conducted, and almost all of them were took place on the southern bank of the Sanggan River, and the Shanshenmiaozui Site ( Fig. 1 ) is among the most productive sites in mammalian fossils. Up to now, 25 species, including undetermined species, have been recognized and/or published: Allactaga sibirica (Tong et al., 2021 ), Ochotonoides complicidens (Tong et al., 2021 ), Ochotona youngi (Tong et al., 2021 ), Nyctereutes sp., Canis chihliensis (Tong et al., 2012 , 2021 ), Homotherium sp., Acinonyx sp., Lynx sp., Panthera sp., Pachycrocuta licenti, Mammuthus trogontherii (Tong, 2012 ; Tong and Chen, 2016 ; Chen and Tong, 2017 ; Tong et al., 2021 ), Hipparion sp., Equus sanmeniensis, Coelodonta nihowanensis (Tong and Wang, 2014 ), Elasmotherium peii (Tong et al., 2014 , 2018 ), Sus lydekkeri, Paracamelus gigas (Tong et al., 2021 ), Nipponicervus elegans (Tong et al., 2021 ), Eucladoceros boulei (Tong and Zhang, 2019 ), Elaphurus bifurcatus (Tong et al., 2021 ), Spirocerus wongi, Gazella sinensis, Ovis shantungensis, Megalovis piveteaui, Bison palaeosinensis (Tong et al., 2017 ). Among the 25 taxa mentioned above, 23 of which once appeared in the classical Nihewan fauna (=Xiashagou fauna), which account for 92% of the SSMZ fauna. ...

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