Jiufotang Formation

Jiufotang Formation
Stratigraphic range: Aptian
Jiufotang Formation; Stratigraphic range: Aptian
Type	Geological formation
Unit of	Jehol Group
Sub-units	Shangheshou Bed, Meileyingzi Bed
Underlies	Fuxin Formation, Shahai Formation
Overlies	Yixian Formation
Location
Region	Liaoning
Country	China

The Jiufotang Formation (Chinese: 九佛堂组, pinyin: jiǔfótáng zǔ) is an Early Cretaceous geological formation in Chaoyang, Liaoning which has yielded fossils of feathered dinosaurs, primitive birds, pterosaurs, and other organisms (see Jehol Biota). It is a member of the Jehol group. The exact age of the Jiufotang has been debated for years, with estimates ranging from the Late Jurassic to the Early Cretaceous. He et al. (2004) used argon - isotope radiometry to confirm biostratigraphic age estimates. They confirmed an Early Cretaceous, Aptian age for the Jiufotang Formation, 120.3 +/-0.7 million years ago. Fossils of Microraptor and Jeholornis are from the Jiufotang.[1]

Fauna

Choristoderans

Genus	Species	Notes
Philydrosaurus	Philydrosaurus proseilus	Monjurosuchid
Ikechosaurus	Ikechosaurus pijiagouensis	Simoedosaurid
Liaoxisaurus	Liaoxisaurus chaoyangensis	Simoedosaurid

Fish

Genus	Species	Notes	Images
Jinanichthys	Jinanichthys longicephalus	An osteoglossiform.	Lycoptera Sinamia
Lycoptera	Lycoptera davidi	An osteoglossiform.
Peipiaosteus	Peipiaosteus pani	A fish related to Sturgeons.
Protosephurus	Protosephurus liui	A Paddlefish.
Sinamia	Sinamia zdanskyi	A Bowfin.

Mammals

Several mammal specimens have been found from the Jiufotang, but only one has been formally described and named.

Genus	Species	County	Member	Abundance	Notes
Liaoconodon	Liaoconodon hui[2]			Nearly complete skeleton	A eutriconodont
Unnamed mammal	Unnamed			Partial remains including a foot and fragments of the skull	A mammal similar to Eomaia and Sinodelphys from the Yixian Formation. Found as stomach contents of Microraptor zhaoianus.

Ornithischians

Genus	Species	Member	Material	Notes	Images
Chuanqilong[3]	C. chaoyangensis[3]			An ankylosaurine.[3]
Psittacosaurus	Psittacosaurus meileyingensis	Meileyingzi Bed	"[Four] individuals, [two] complete skulls."[4]	A ceratopsian.	Psittacosaurus meileyingensis Psittacosaurus mongoliensis
Psittacosaurus	Psittacosaurus mongoliensis	Shangheshou and Meileyingzi Beds		A ceratopsian.

Pterosaurs

Genus	Species	Notes	Images
Chaoyangopterus	C. zhangi	A chaoyangopterid.[5]	Liaoningopterus Nurhachius Nemicolopterus
Eoazhdarcho	E. liaoxiensis	A chaoyangopterid.[5]
Guidraco	G. venator	A pteranodontoid.[6]
"Huaxiapterus"	"H." benxiensis	In need of new generic name.
"Huaxiapterus"	"H." corollatus	A tapejarid, in need of new generic name, probably the same genus as "H." benxiensis.
Ikrandraco	I. avatar	A pteranodontoid.[6]
Jidapterus	J. edentus	A chaoyangopterid.[5]
Liaoningopterus	L. gui	A pteranodontoid.[6]
Liaoxipterus	L. brachyognathus	An istiodactylid.[5]
Linlongopterus	L. jennyae	A pteranodontoid.[6]
Nemicolopterus	N. crypticus	A tapejaromorph. Previously and still occasionally thought to be a Tapejarid. Most likely a juvenile, hence small size.
Nurhachius	N. ignaciobritoi	An istiodactylid.[5]
Shenzhoupterus	S. chaoyangensis	A chaoyangopterid.
Sinopterus	S. dongi	A tapejarid.[5]
Sinopterus	S. jii	Previously known as Huxiapterus jii.

Vesperopterylus	V. lamadongensis	An anurognathid.

Saurischians

A large titanosaur is present in the formation.[7]

Enantiornithines

Genus	Species	Material	Notes	Images
Alethoalaornis	Alethoalaornis agitornis[8]		An enantiornithean	Longipteryx Rapaxavis Sinornis
Boluochia	Boluochia zhengi	"Partial skull [and] partial postcranium."[9]	A longipterygid
Cathayornis	Cathayornis aberransis		An entantiornithine
	Cathayornis caudatus		Nomen dubium[10]
	Cathayornis yandica	"Partial skeleton."[9]	An enantiornithean
Cuspirostrisornis	Cuspirostrisornis houi		A possible avisaurid
Dapingfangornis	Dapingfangornis sentisorhinus		An enantiornithean
Eocathayornis	Eocathayornis walkeri	"Partial skeleton."[9]	An enantiornithean
Gracilornis	Gracilornis jiufotangensis		An enantiornithean
Huoshanornis	Huoshanornis huji		An enantiornithean
Largirostrornis	Largirostrornis sexdentoris		An enantiornithean
Longchengornis	Longchengornis sanyanensis		An enantiornithean
Longipteryx	Longipteryx chaoyangensis		A longipterygid
Rapaxavis	Rapaxavis pani		A longipterygid
Shangyang	Shangyang graciles		An enantiornithean
Sinornis	Sinornis santensis	"Partial skull [and] many nearly complete postcranial skeletons."[11]	An enantiornithean
Xiangornis	Xiangornis shenmi		An enantiornithean

Euornithines

Genus	Species	County	Material	Notes	Images
Aberratiodontus	Aberratiodontus wui			Junior synonym of Yanornis martini[12]	Yixianornis grabaui
Chaoyangia	Chaoyangia beishanensis		"Partial skeleton."[11]	An ornithuran
Jianchangornis	Jianchangornis microdonta	Jianchang		An ornithuran
Parahongshanornis[13]	Parahongshanornis chaoyangensis			A Hongshanornithid
Schizooura	Schizooura lii	Jianchang		An ornithuran with long, forked tail feathers
Songlingornis	Songlingornis linghensis			An ornithuran.
Yanornis	Yanornis martini			An ornithuran.
Yixianornis	Yixianornis grabaui			An ornithuran.

Misc theropods

Color key

Taxon	Reclassified taxon	Taxon falsely reported as present	Dubious taxon or junior synonym	Ichnotaxon	Ootaxon	Morphotaxon

Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.

Genus	Species	County	Member	Material	Notes	Images
Confuciusornis	C. sanctus				A primitive short-tailed bird (pygostylian). Also present in the Yixian Formation	Confuciusornis sanctus Jeholornis prima Microraptor gui Sinotyrannus kazuoensis
Confuciusornis	C. jianchangensis	Liaoning
Dalianraptor	D. cuhe				Nomen dubium, named from a forged artificially assembled specimen.
Jeholornis[1]	J. prima[1]				A basal avialan, also present in the Yixian Formation[14]
Jeholornis[1]	J. palmapenis[15]				A basal avialan
Kompsornis	K. longicaudatus			"Nearly complete specimen".[16]	A long tailed avialan, related to Jeholornis.
Microraptor	M. gui		Shangheshou Bed	Extremely common[7]	Could be synonym of Microraptor zhaoianus
Microraptor	M. zhaoianus		Shangheshou Bed[7]	"At least [three] partial semiarticulated postcrania and several skulls."[17] Feathered integument is also preserved.[17]	A dromaeosaurid.
Omnivoropteryx	O. sinousaorum				An omnivoropterygiform
Sapeornis	S. angustis			"Nearly complete skeleton without skull."[18]	An omnivoropterygiform, a primitive short-tailed bird
Sapeornis	S. chaoyangensis				An omnivoropterygiform, a primitive short-tailed bird
Shenshiornis	Shenshiornis primita[19]				An omnivoropterygiform
Similicaudipteryx	S. yixianensis				An oviraptorosaur
Sinotyrannus[20]	S. kazuoensis[20]				A tyrannosauroid[20]
Zhongjianornis	Z. yangi				A primitive short-tailed bird[21]

References

He, H.Y.; Wang, X.L.; Zhou, Z.H.; Wang, F.; Boven, A.; Shi, G.H.; Zhu, R.X. (2004). "Timing of the Jiufotang Formation (Jehol Group) in Liaoning, northeastern China, and its implications". Geophysical Research Letters. 31 (13): 1709. Bibcode:2004GeoRL..3112605H. doi:10.1029/2004GL019790.
Meng, J.; Wang, Y.; Li, C. (2011). "Transitional mammalian middle ear from a new Cretaceous Jehol eutriconodont". Nature. 472 (7342): 181–185. Bibcode:2011Natur.472..181M. doi:10.1038/nature09921. PMID 21490668.
Han, F.; Zheng, W.; Hu, D.; Xu, X.; Barrett, P.M. (2014). "A New Basal Ankylosaurid (Dinosauria: Ornithischia) from the Lower Cretaceous Jiufotang Formation of Liaoning Province, China". PLoS ONE. 9 (8): e104551. Bibcode:2014PLoSO...9j4551H. doi:10.1371/journal.pone.0104551. PMC 4131922. PMID 25118986.
"Table 22.1," in Weishampel, et al. (2004). Page 479.
Wang, X.; Zhou, Z. (2006). "Pterosaur assemblages of the Jehol Biota and their implication for the Early Cretaceous pterosaur radiation". Geological Journal. 41 (3–4): 405–418. doi:10.1002/gj.1046.
Rodrigues, Taissa; Jiang, Shunxing; Cheng, Xin; Wang, Xiaolin; Kellner, Alexander W.A. (2015). "A new toothed pteranodontoid (Pterosauria, Pterodactyloidea) from the Jiufotang Formation (Lower Cretaceous, Aptian) of China and comments on Liaoningopterus gui Wang and Zhou, 2003". Historical Biology. 27 (6): 782–795. doi:10.1080/08912963.2015.1033417.
Xu, X.; Norell, M.A. (2006). "Non-Avian dinosaur fossils from the Lower Cretaceous Jehol Group of western Liaoning, China". Geological Journal. 41 (3–4): 419–437. doi:10.1002/gj.1044.
Li, L., Hu, D.-Y., Duan, Y., Gong, E.-P. and Hou, L.-H. (2007). "Alethoalaornithidae fam. nov., a new family of enantiornithine bird from the Lower Cretaceous of western Liaoning." Acta Palaeontologica Sinica, 46(3): 365-372. (http://www.cqvip.com/qk/90074X/200703/25642898.html) [in Chinese]
"Table 11.1," in Weishampel, et al. (2004). Page 212.
O'Connor, J.; Dyke, G. (2010). "A reassessment of Sinornis santensis and Cathayornis yandica (Aves: Enantiornithes)". Records of the Australian Museum. 62: 7–20. doi:10.3853/J.0067-1975.62.2010.1540.
"Table 11.1," in Weishampel, et al. (2004). Page 214.
Cau, A.; Arduini, P. (2008). "Enantiophoenix electrophyla gen. et sp. nov. (Aves, Enantiornithes) from the Upper Cretaceous (Cenomanian) of Lebanon and its phylogenetic relationships". Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale in Milano. 149: 293–324.
Li, Li; Jing-Qi Wang; Shi-Lin Hou (2011). "A new ornithurine bird (Hongshanornithidae) from the Jiufotang Formation of Chaoyang, Liaoning, China" (PDF). Vertebrata PalAsiatica. 49 (2): 195–200.
Li, D.; Sulliven, C.; Zhou, Z.; Zhang, Z. (2010). "Basal birds from China: a brief review". Chinese Birds. 1 (2): 83–96. doi:10.5122/cbirds.2010.0002.
O'Connor, J.K.; Sun, C.; Xu, X.; Wang, X.; Zhou, Z. (2012). "A new species of Jeholornis with complete caudal integument". Historical Biology. 24: 29–41. doi:10.1080/08912963.2011.552720.
Xuri Wang; Jiandong Huang; Martin Kundrát; Andrea Cau; Xiaoyu Liu; Yang Wang; Shubin Ju (2020). "A new jeholornithiform exhibits the earliest appearance of the fused sternum and pelvis in the evolution of avialan dinosaurs". Journal of Asian Earth Sciences. in press: Article 104401. doi:10.1016/j.jseaes.2020.104401.
"Table 10.1," in Weishampel, et al. (2004). Page 198.
"Table 11.1," in Weishampel, et al. (2004). Page 216.
Hu, D.; Li, L.; Hou, L.; Xu, X. (2010). "A new sapeornithid bird from China and its implication for early avian evolution". Acta Geologica Sinica (English Edition). 84 (3): 472–482. doi:10.1111/j.1755-6724.2010.00188.x.
Ji, Q.; Ji, S.-A.; Zhang, L.-J. (2009). "First large tyrannosauroid theropod from the Early Cretaceous Jehol Biota in northeastern China". Geological Bulletin of China. 28 (10): 1369–1374.
Zhou, Z.; Zhang, F.; Li, Z. (2010). "A new Lower Cretaceous bird from China and tooth reduction in early avian evolution". Proceedings of the Royal Society B. 277 (1679): 219–227. doi:10.1098/rspb.2009.0885. PMC 2842673. PMID 19586952.