Jiufotang Formation

Jiufotang Formation
Jiufotang Formation
Stratigraphic range: Aptian; ~122–118.9 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
Type	Geological formation
Unit of	Jehol Group
Sub-units	Shangheshou & Meileyingzi Beds
Underlies	Fuxin & Shahai Formations
Overlies	Yixian Formation
Lithology
Primary	Mudstone
Other	Sandstone
Location
Coordinates	41°12′N 119°24′E / 41.2°N 119.4°E
Approximate paleocoordinates	44°24′N 119°30′E / 44.4°N 119.5°E
Region	Liaoning
Country	China
Type section
Named for	Jiufotang, Pingfangzi, Liaoning
Named by	Ryuji Endo
Year defined	1934
	Jiufotang Formation (China) Jiufotang Formation (Liaoning)

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. New uranium-lead dates reveal the formation is deposited in the Aptian stage of the Early Cretaceous.^[2] Fossils of Microraptor and Jeholornis are from the Jiufotang.^[3]

Fossil content

Choristoderans

Genus	Species	Notes
Philydrosaurus	P. proseilus	Monjurosuchid
Ikechosaurus	I. pijiagouensis	Simoedosaurid
Liaoxisaurus	L. chaoyangensis	Simoedosaurid
Hyphalosaurus	H. baitaigouensis^[4]	Hyphalosaurid

Fish

Genus	Species	County	Notes
Caeruleum	C. miraculum^[5]	Weichang	A lamprey
Caeruleum	C. gracilis^[6]	Weichang	A lamprey
Jinanichthys	J. longicephalus		An osteoglossomorph
Lycoptera	L. davidi		An osteoglossomorph
Peipiaosteus	P. pani		A fish related to sturgeons
Protopsephurus	P. liui		A paddlefish
Sinamia	S. zdanskyi		An amiiform

Mammaliamorphs

Several mammaliamorph specimens have been found from the Jiufotang, but only a few have been formally described and named.

Genus	Species	Material	Notes
Cokotherium	C. jiufotangensis^[7]	Partial skeleton	A eutherian
Fossiomanus	F. sinensis^[8]	A nearly complete skeleton	A tritylodontid cynodont, likely the youngest known non-mammalian synapsid
Liaoconodon	L. hui^[9]	A 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	County	Member	Material	Notes	Images
Chuanqilong^[10]	C. chaoyangensis^[10]	Liaoning		A nearly complete skeleton missing only the distal portion of the caudal series.	A basal ankylosaurid, sister taxon to Liaoningosaurus.^[10]
Liaoningosaurus^[11]	L. paradoxus^[11]					Liaoningosaurus paradoxus
Psittacosaurus	Psittacosaurus meileyingensis		Meileyingzi Bed	"[Four] individuals, [two] complete skulls."^[12]	A ceratopsian.	Psittacosaurus meileyingensis Psittacosaurus mongoliensis
Psittacosaurus	Psittacosaurus mongoliensis		Shangheshou and Meileyingzi Beds		A ceratopsian.

Pterosaurs

Genus	Species	County	Member	Material	Notes	Images
Chaoyangopterus	C. zhangi				A chaoyangopterid^[13]	Liaoningopterus Nemicolopterus Nurhachius Sinopterus and Huaxiadraco
Eoazhdarcho	E. liaoxiensis				A chaoyangopterid^[13]
Feilongus	F. youngi		Jianshangou Bed		A boreopterid
Forfexopterus	F. jeholensis				A ctenochasmatid
Gladocephaloideus	G. jingangshanensis		Jingangshan Bed		A ctenochasmatid
Guidraco	G. venator				A pteranodontoid^[14]
Hongshanopterus	H. lacustris				An istiodactylid
Huaxiadraco	H. corollatus			Multiple specimens	A tapejarid^[15]
"Huaxiapterus"	"H." benxiensis				A tapejarid; synonymized with Huaxiadraco in 2023
Ikrandraco	I. avatar				A pteranodontoid^[14]
Jidapterus	J. edentus				A chaoyangopterid^[13]
Liaoningopterus	L. gui				A pteranodontoid^[14]
Liaoxipterus	L. brachyognathus				An istiodactylid^[13]
Linlongopterus	L. jennyae				A pteranodontoid^[14]
Meilifeilong	M. youhao	Jianchang		A nearly complete skeleton and a partial skull	A chaoyangopterid^[16]
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^[13]
Pangupterus	P. liui				A pterodactyloid
Shenzhoupterus	S. chaoyangensis				A chaoyangopterid
Shenzhoupterus	S. sanyainus				A chaoyangopterid^[17]
Sinopterus	S. dongi				A tapejarid^[13]
Sinopterus	S. jii				Previously known as Huxiapterus jii
Vesperopterylus	V. lamadongensis				An anurognathid

Saurischians

A large titanosaur is present in the formation.^[18]

Enantiornithines

Genus	Species	County	Material	Notes	Images
Alethoalaornis	A. agitornis^[19]			An enantiornithean	Longipteryx Rapaxavis Sinornis Yuanchuavis
Boluochia	B. zhengi		"Partial skull [and] partial postcranium."^[20]	A longipterygid
Brevirostruavis	B. macrohyoideus		"an extremely elongate bony hyoid element (only slightly shorter than the skull), combined with a short cranial rostrum."^[21]	An enantiornithine
Cathayornis	C. aberransis			An enantiornithine
	C. caudatus			Nomen dubium^[22]
	C. yandica		"Partial skeleton."^[20]	An enantiornithean
Cuspirostrisornis	C. houi			A possible avisaurid
Dapingfangornis	D. sentisorhinus			An enantiornithean
Eocathayornis	E. walkeri		"Partial skeleton."^[20]	An enantiornithean
Gracilornis	G. jiufotangensis			An enantiornithean
Imparavis^[23]	I. attenboroughi	Jianchang	An articulated, nearly complete specimen
Huoshanornis	H. huji			An enantiornithean
Largirostrornis	L. sexdentoris			An enantiornithean
Longchengornis	L. sanyanensis			An enantiornithean
Longipteryx	L. chaoyangensis			A longipterygid
Musivavis	M. amabilis		"A nearly complete and articulated skeleton preserved in a single slab."^[24]	An euenantiornithean
Rapaxavis	R. pani			A longipterygid
Shangyang	S. graciles			An enantiornithean
Sinornis	S. santensis		"Partial skull [and] many nearly complete postcranial skeletons."^[25]	An enantiornithean
Xiangornis	X. shenmi			An enantiornithean
Yuanchuavis	Y. kompsosoura			An pengornithid

Euornithines

Genus	Species	County	Material	Notes	Images
Aberratiodontus	A. wui			Junior synonym of Yanornis martini^[26]	Yanornis martini Yixianornis grabaui
Chaoyangia	C. beishanensis		"Partial skeleton."^[25]	A basal euornithean
Jianchangornis	J. microdonta	Jianchang		A basal euornithean
Mengciusornis	M. dentatus			A schizoourid
Parahongshanornis^[27]	P. chaoyangensis			A hongshanornithid
Piscivoravis	P. lii			An ornithuromorph
Schizooura	S. lii	Jianchang		A schizoourid with long, forked tail feathers
Similiyanornis	S. brevipectus			An ornithuromorph
Songlingornis	S. linghensis			An ornithuromorph
Yanornis	Y. martini			An ornithuromorph
Yixianornis	Y. grabaui			An ornithuromorph
Zhongjianornis	Z. yangi			A basal euornithean^[28]

Non-ornithothoracean 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
Chongmingia^[29]	C. zhengi			A partial skeleton with preserved soft tissues	A jinguofortisid avialan	Confuciusornis sanctus Jeholornis prima Microraptor gui Sinotyrannus kazuoensis Wulong bohaiensis
Confuciusornis	C. sanctus				A primitive short-tailed avialan (pygostylian). Also present in the Yixian Formation
Confuciusornis	C. jianchangensis	Liaoning
Cratonavis^[30]	C. zhui	Jianchang		A complete skeleton with preserved feathers	A jinguofortisid avialan
Dalianraptor	D. cuhe				Nomen dubium, named from a forged artificially assembled specimen.
Jeholornis^[3]	J. prima^[3]				A jeholornithiform avialan, also present in the Yixian Formation^[31]
Jeholornis^[3]	J. palmapenis^[32]				A jeholornithiform avialan
Kompsornis	K. longicaudatus			"Nearly complete specimen".^[33]	A jeholornithiform avialan
Microraptor	M. gui		Shangheshou Bed	Extremely common^[18]	Could be synonym of Microraptor zhaoianus
Microraptor	M. zhaoianus		Shangheshou Bed^[18]	"At least [three] partial semiarticulated postcrania and several skulls."^[34] Feathered integument is also preserved.^[34]	A dromaeosaurid.
Neimengornis	N. rectusmim				A jeholornithiform avialan
Omnivoropteryx	O. sinousaorum				An omnivoropterygiform avialan
Sapeornis	S. angustis			"Nearly complete skeleton without skull."^[35]	An omnivoropterygiform, a primitive short-tailed avialan
Sapeornis	S. chaoyangensis				An omnivoropterygiform, a primitive short-tailed avialan
Shenshiornis	Shenshiornis primita^[36]				An omnivoropterygiform avialan
Similicaudipteryx	S. yixianensis				An oviraptorosaur
Sinotyrannus^[37]	S. kazuoensis^[37]				A tyrannosauroid^[37]
Wulong^[38]	W. bohaiensis^[38]	Chaoyang, Liaoning^[38]		Complete articulated skeleton (D2933).^[38]	A dromaeosaurid

Crustaceans

Genus	Species	County	Member	Material	Notes	Images
Eosestheria	Eosestheria middendorfii^[39]				A clam shrimp

Plants

Genus	Species	Notes
Czekanowskia	Czekanowskia rigida^[40]	A member of Czekanowskiales, Dominant
Ginkgo	Ginkgo digitata^[40]	A member of Ginkgoales
Elatocladus	Elatocladus pinnatus^[40]	Conifer foliage
Pityolepis	Pityolepis ovatus^[40]	Conifer cone
Leptostrobus	Leptostrobus sinensis^[40]	A member of Czekanowksiales, occasionally misspelled "Leptosthrobus"
Baiera	Baiera valida^[40]	A member of Ginkgoales
Baiera	Baiera cf. gracilis^[40]	A member of Ginkgoales
Ginkgoites	Ginkgoites trancata^[40]	A member of Ginkgoales, dominant
Ginkgoites	Gingkgoites trancatus^[40]	A member of Ginkgoales, dominant
Podozamites	Podozamites sp.^[39]	Conifer foliage
Sphenopteris	Sphenopteris sp.^[39]	Seed plant foliage

References

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