Liushu Formation Explained

Liushu Formation
Age:	Late Miocene ~
Period:	Late Miocene
Type:	Geological formation
Prilithology:	Red claystone, Siltstone
Otherlithology:	Mudstone
Region:	Linxia Basin, Gansu Province.

The Liushu Formation is a geological formation in Gansu province, China that spans up to 100 m thick and is widely distributed within the Linxia Basin, with a paleomagnetic age between 11 and 6.4 mya (Upper/Late Miocene).

Geology

The formation is divided into three parts representing three different ages, generally referred to as the "upper part" (youngest, at 6-7 mya), the "middle part" (intermediate, at 9 to 7.6 mya, also known as the Dashengou Fauna), and the "lower part" (oldest).^[1]

Paleobiota

Birds

Genus	Species	Locality	Stratigraphic position	Material	Notes	Images
Gansugyps ^[2]	G. linxiaensis	Yangwapuzijifang, Guanghe County	Upper part	A preserved skeleton	An extinct old-world vulture
Linxiavis	L. inaquosus^[3]	Baiwang, Gansu Province	Middle part–Upper part	Articulated and associated elements of the wings, shoulder girdle, vertebrae, and hind limb	An extinct sandgrouse
Miosurnia^[4]	M. diurna	Linxia Basin of Gansu Province	Upper part	A nearly complete articulated bird skeleton, lacking the right forelimb and left manual digits	A diurnal surniin bird that belongs to the owl family.
Mioneophron^[5]	M. longirostris	Baihua Village, Zhuangheji Town, Guanghe County	Upper part	A nearly complete skeleton	An extinct old-world vulture.
Panraogallus	P. hezhengensis	Zhuangkeji Township, Gansu Province	Upper part	A skeleton	A phasianid bird
Sinoergilornis	S. guangheensis	Langwagou Village, Zhuangkeji Town, Guanghe County	Upper part	A three-dimensionally preserved distal left tarsometatarsus and a nearly complete set of pedal phalanges	An eogruid
Struthio^[6]	S. linxiaensis	Yangwapuzijifang, Guanghe County	Upper part	An incomplete pelvis and synsacrum	An extinct ostrich

Mammals

Artiodactyla

Genus	Species	County	Stratigraphic position	Material	Notes	Images
Chleuastochoerus^[7]	C. linxiaensis	Yangwapuzi, Guanghe County	Upper part	The cranium of a male individual and a juvenile female	A Hyotheriinae suidae.
	C. stehlini		Middle to Upper part	Cranium skull and mandible
Dicrocerus	D. sp.	Houshan locality, Guanghe County	Middle to Upper part.		An ancient deer
Dorcadoryx	D. sp.	Baihuacun and Langwagou locality.	Middle to Upper part.		An early caprini relative.
Eostyloceros^[8]	E. hezhengensis	Gaojiashan, Hezheng County	Upper part	An adult skull with its cranial appendages, lacking the muzzle, the left dentition, and the basioccipital part
Euprox^[9]	E. furcatus		Upper part	An adult skull and pair of near-complete antlers.	The largest known Eupox species ever discovered
Gazella^[10]	G. dorcadoides	Houshan and Baihuacun locality, Guanghe County	Upper part.		Early relatives of gazelles.
	G. gaudryi		Middle to Upper part
	G. paotehensis			Multiple skulls.
Hezhengia	H. bohlini	Houshan, Bajia, Shuanggongbei, and Zhongmajia locality, Guanghe County	Middle part		A bovid
Honanotherium	H. schlosseri	Houshan locality, Guanghe County	Middle part		An ancient giraffid
Miotragocerus	M. gregarius	Houshan locality, Guanghe County	Middle part		An extinct antelope
Paleotragus	P. microdon	Houshan locality, Guanghe County	Upper part		An ancient giraffid
Protoryx	P. sp.	Houshan locality, Guanghe County	Upper part		An ancient bovid
Samotherium	S. boissieri	Houshan locality, Guanghe County	Middle part	A cranium skull.	An ancient giraffid
Schansitherium	S. tafeli	Dashengou and Yangjiashan, Gansu County	Upper part	A cranium skull.	A four-horned giraffid
Shaanxispira	S. linxiaensis.	Panyang and Songjianao locality.	Lower to Middle part	A cranium skull.	An ancient bovid
Sinotragus	S. wimani	Songshugou locality, Guanghe County	Upper part		An ancient bovid
Tsaidamotherium ^[11]	T. brevirostrum	Yancaiping locality, Hezheng County.	Lower part.	A cranium skull.	An ovibovinid caprinae .
Urmiatherium^[12]	U. intermedium	Huaigou village, Gansu province	Upper part	A male adult skull	An early caprinae.

Carnivora

Caniformia

Genus	Species	County	Stratigraphic position	Material	Notes	Images
Melodon	M. majori		Middle part		A musteloid
Parataxidea	P. sinensis	Houshan locality, Guanghe County	Upper part	A fossil skull with maxillary teeth.	a badger-like mustelid
Promephitis	P. hootoni		Middle part		A musteloid
	P. sp.
Sinictis	S. sp.		Middle part		A musteloid
Ursavus^[13]	U. tedfordi.	Huaigou village, Guanghe County	Upper part	A cranium skull in association with the mandible	An early bear

Feliformia

Genus	Species	County	Stratigraphic position	Material	Notes	Images
Amphimachairodus	A. giganteus	Houshan, Guanghe County	Upper part to Middle part		A homotheriine felid.
	A. hezhengensis^[14]		Upper part.	A nearly complete cranium skull
Dinocrocuta	D. gigantea	Houshan locality, Guanghe County	Middle part	A complete cranium skull.	A gigantic hyena
Felis	F. sp.	Guanghe County	Upper part to Middle part		A felid.
Hyaenictitherium	H. hyaenoides	Houshan locality, Guanghe County	Upper part to Middle part		A hyaenid
	H. wongii
Ictitherium	I. sp.	Guanghe County	Upper part to Middle part		A hyaenid
Metailurus	M. major	Guanghe County	Upper part		A metalurin felid.

Perissodactyla

The Liushu formation displays a lineage of Elasmotheriine rhinoceros throughout its stratigraphy. Iranotherium is restricted to the middle part of the formation but remains absent in the overlying layers, the only Elasmotheriine that can be found in the upper layers is Sinotherium.

Chalicotheres

Genus	Species	Locality	Stratigraphic position	Material	Notes	Images
Ancylotherium^[15]	A. sp.	Duikang locality, Guanghe County.	Lower part	A juvenile cranium skull.	A shizotheriine chalicothere
Nestoritherium	N. linxiaense	Houshan locality, Guanghe County	Lower and Middle parts	A skull without the occipital surface, right zygomatic arch, premaxilla, and anterior nasal bone	A chalicothere

Equids

Genus	Species	Locality	Stratigraphic position	Material	Notes	Images
Hipparion	H. chiai	Guanghe County	Middle part		An ancient horse
	H. coelophyes		Upper part
	H. dongxiangense^[16]		Lower part
	H. weihoensis		Middle part

Rhinocerotids

Genus	Species	Locality	Stratigraphic position	Material	Notes	Images
Acerorhinus	A. hezhengensis		Middle part	Skulls based on sexual dimorphism	A hornless rhino
Chilotherium	C. wimani	Zhongmajia, Guanghe County	Middle part	Several skull specimens.	A tusked rhino
Iranotherium	I. morgani^[17]	Guanghe County	Middle part	Two skulls based on sexual dimorphism and an isolated mandible	An elasmotheriine with a unique sexual dimorphism, ancestral to Sinotherium
Ningxiatherium^[18]	N. euryrhinus.	Guonigou in Nalesi Township, Dongxiang County	Lower part	A completely adult skull with both cheek tooth rows
Parelasmotherium	P. linxiaense	Dongxiang County	Lower part	A near complete skull	A grazing elasmotheriine
	P. simplum		Lower part
Sinotherium	S. lagrelii^[19]	Guanghe County	Upper part	A partial skull	An elasmotheriine, ancestral to Elasmotherium. It displays the important transition from nose-horned to forehead-horned rhinos.

Rodents

Genus	Species	County	Member	Material	Notes	Images
Huerzelerimys^[20]	H. asiaticus	Niuzhawan, Sanhe Village, Hezheng County	Upper part.	A complete skull with a mandible and 5 cervical .	An early murinae rodent.
Hystrix^[21]	H. brevirostra	Maijiaji Village, Hezheng County.	Upper part.	A nearly complete skull.	An early porcupine.
Pararhizomys	P. hipparionum		Middle part		A rodent
	P. parvulus^[22]	Xiayangwan, Guanghe County.	Middle part	An anterior part of a skull with two I2s, left M1‒3 and right M1‒2.

Proboscidea

Genus	Species	Locality	Member	Material	Notes	Images
Tetralophodon	T. exoletus	Houshan locality, Guanghe County	Middle part		A gomphothere
Konobelodon^[23]	K. robustus	Zhongmajia and Guonigou, Guanghe County	Middle and Upper parts	Multiple specimens comprising a skull, pelvis bone, and incomplete mandibles	An amebelodont

Reptiles

Notes and References

1. Web site: Vertebrata PalAsiatica, Vol. 42, 2004----Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences. 2020-12-21. english.ivpp.cas.cn.
2. Zhang, Zihui & Xiaoting, Zheng & Zheng, Guangmei & Hou, Lianhai. (2010). A new Old World vulture (Falconiformes: Accipitridae) from the Miocene of Gansu Province, northwest China. Journal of Ornithology. 151. 401-408. 10.1007/s10336-009-0468-1.
3. Li. Zhiheng. Stidham. Thomas A.. Deng. Tao. Zhou. Zhonghe. 2020-03-31. Evidence of Late Miocene Peri-Tibetan Aridification From the Oldest Asian Species of Sandgrouse (Aves: Pteroclidae). Frontiers in Ecology and Evolution. 8. 59. 10.3389/fevo.2020.00059. 2296-701X. free.
4. Li, Z., Stidham, T.A., Zheng, X., Wang, Y., Zhao, T., Deng, T., & Zhou, Z. (2022). Early evolution of diurnal habits in owls (Aves, Strigiformes) documented by a new and exquisitely preserved Miocene owl fossil from China. Proceedings of the National Academy of Sciences of the United States of America, 119.
5. li, Zhiheng & Clarke, Julia & Zhou, Zhonghe & Deng, Tao. (2016). A new Old World vulture from the late Miocene of China sheds light on Neogene shifts in the past diversity and distribution of the Gypaetinae. The Auk. 133. 615-625. 10.1642/AUK-15-240.1.
6. Hou, L., Zhou, Z., Zhang, F. et al. A Miocene ostrich fossil from Gansu Province, northwest China. Chin.Sci.Bull. 50, 1808–1810 (2005). https://doi.org/10.1360/982005-575
7. Hou S, Deng T. A new species of Chleuastochoerus (Artiodactyla: Suidae) from the Linxia Basin, Gansu Province, China. Zootaxa. 2014 Oct 13;3872(5):401-39. doi: 10.11646/zootaxa.3872.5.1. PMID 25544095.
8. Deng, Tao, Wang, Shi-Qi, Shi, Qin-Qin, Li, Yi-Kun & Li, Yu, 2014, A new species of Eostyloceros (Cervidae, Artiodactyla) from the Late Miocene of the Linxia Basin in Gansu, China, Zootaxa 3893 (3), pp. 363-381
9. Hou, Sukuan. (2014). A new species of Euprox (Cervidae, Artiodactyla) from the upper Miocene of the Linxia Basin, Gansu Province, China, with interpretation of its paleoenvironment. Zootaxa. 3911. 43-62. 10.11646/zootaxa.3911.1.2.
10. Shi, Qinqin. (2023). Fossil bovids from the Linxia Basin of Gansu Province, China, and their implications for regional biostratigraphy, palaeogeography and palaeoecology. Palaeogeography, Palaeoclimatology, Palaeoecology. 619. 111543. 10.1016/j.palaeo.2023.111543.
11. SHI QinQin, "New species of Tsaidamotherium (Bovidae, Artiodactyla) from China sheds new light on the skull morphology and systematics of the genus." SCIENCE CHINA Earth Sciences 2014, Vol. 57 Issue (2): 258-266 DOI: 10.1007/s11430-013-4722-2
12. Shi, Qinqin & Wang, Shiqi & Chen, Shaokun & Li, Yi-kun & Palasiatica, Vertebrata. (2016). The first discovery of Urmiatherium (Bovidae, Artiodactyla) from Liushu Formation, Linxia Basin. Vertebrata Palasiatica.
13. QiuZ, X., Deng, T., & WangB, Y. (2014). A Late Miocene Ursavus skull from Guanghe, Gansu, China.
14. Jiangzuo Q, Werdelin L, Sanisidro O, Yang R, Fu J, Li S, Wang S, Deng T. Origin of adaptations to open environments and social behaviour in sabretoothed cats from the northeastern border of the Tibetan Plateau. Proc Biol Sci. 2023 Apr 26;290(1997):20230019. doi: 10.1098/rspb.2023.0019. Epub 2023 Apr 19. PMID 37072045; PMCID: PMC10113030.
15. Chen, Shaokun & Deng, Tao & Pang, Li-Bo & He, Wen & Chen, Shan-Qin. (2012). A juvenile skull of Ancylotherium (Mammalia, Perissodactyla, Chalicotheriidae) from the Pliocene of China. Geobios. 45. 527-534. 10.1016/j.geobios.2012.06.002.
16. Deng. Tao. 2007-06-12. Skull of Parelasmotherium (Perissodactyla, Rhinocerotidae) from the upper miocene in the Linxia Basin (Gansu, China). Journal of Vertebrate Paleontology. en. 27. 2. 467–475. 10.1671/0272-4634(2007)27[467:SOPPRF]2.0.CO;2. 130213090 . 0272-4634.
17. Deng. Tao. 2005-06-27. https://doi.org/10.1671/0272-4634(2005)025[0442:NDOIMP2.0.CO;2 New discovery of Iranotherium morgani (Perissodactyla, Rhinocerotidae) from the late Miocene of the Linxia Basin in Gansu, China, and its sexual dimorphism]. Journal of Vertebrate Paleontology. 25. 2. 442–450. 10.1671/0272-4634(2005)025[0442:NDOIMP]2.0.CO;2. 85820005 . 0272-4634.
18. Deng, Tao. (2008). A new elasmothere (Perissodactyla, Rhinocerotidae) from the Late Miocene of the Linxia Basin in Gansu, China. Geobios. 41. 719-728. 10.1016/j.geobios.2008.01.006.
19. Deng. Tao. Wang. ShiQi. Hou. SuKuan. 2013-05-01. A bizarre tandem-horned elasmothere rhino from the Late Miocene of northwestern China and origin of the true elasmothere. Chinese Science Bulletin. en. 58. 15. 1811–1817. 10.1007/s11434-012-5574-4. 2013ChSBu..58.1811D. 1861-9541. free.
20. Wang B Y, Qiu Z X, Li L Z, 2020. A Late Miocene Huerzelerimys (Rodentia: Muridae)skull from Hezheng, Gansu, China. Vertebrata PalAsiatica, 58(2): 120–133
21. Wang B Y, Qiu Z X, 2020. New Hystrix (Hystricidae, Rodentia) from the Neogene ofLinxia Basin, Gansu, China. Vertebrata PalAsiatica, 58(3): 204–220
22. Wang B Y, 2022. A new species of Pararhizomys (Tachyoryctoidinae, Muroidea)from Linxia Basin of Gansu Province. Vertebrata PalAsiatica, 60(4): 271–277
23. Wang, Shiqi & Shi, Qinqin & He, Wen & Chen, ShanQin & Yang, XiangWen. (2016). A new species of the tetralophodont amebelodontine Konobelodon Lambert, 1990 (Proboscidea, Mammalia) from the Late Miocene of China. Geodiversitas. 38. 65-97. 10.5252/g2016n1a4.