Jiufotang Formation

Jiufotang Formation
Jiufotang Formation
Stratigraphic range: Aptian; ~
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	41.2°N 119.4°E
Approximate paleocoordinates	44.4°N 119.5°E
Region	Liaoning
Country	China
Type section
Named for	Jiufotang, Pingfangzi, Liaoning
Named by	Ryuji Endo
Year defined	1934[1]
	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

Fish

Genus	Species	Notes
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[4]	Partial skeleton	A eutherian
Fossiomanus	F. sinensis[5]	A nearly complete skeleton	A tritylodontid cynodont, likely the youngest known non-mammalian synapsid
Liaoconodon	L. hui[6]	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[7]	C. chaoyangensis[7]	Liaoning		A nearly complete skeleton missing only the distal portion of the caudal series.	A basal ankylosaurid, sister taxon to Liaoningosaurus.[7]
Liaoningosaurus[8]	L. paradoxus[8]					Liaoningosaurus paradoxus
Psittacosaurus	Psittacosaurus meileyingensis		Meileyingzi Bed	"[Four] individuals, [two] complete skulls."[9]	A ceratopsian.	Psittacosaurus meileyingensis Psittacosaurus mongoliensis
Psittacosaurus	Psittacosaurus mongoliensis		Shangheshou and Meileyingzi Beds		A ceratopsian.

Pterosaurs

Genus	Species	Member	Material	Notes	Images
Chaoyangopterus	C. zhangi			A chaoyangopterid[10]	Liaoningopterus Nemicolopterus Nurhachius Sinopterus and Huaxiadraco
Eoazhdarcho	E. liaoxiensis			A chaoyangopterid[10]
Feilongus	F. youngi	Jianshangou Bed		A boreopterid
Forfexopterus	F. jeholensis			A ctenochasmatid
Gladocephaloideus	G. jingangshanensis	Jingangshan Bed		A ctenochasmatid
Guidraco	G. venator			A pteranodontoid[11]
Hongshanopterus	H. lacustris			An istiodactylid
Huaxiadraco	H. corollatus		Multiple specimens	A tapejarid[12]
"Huaxiapterus"	"H." benxiensis			A tapejarid; synonymized with Huaxiadraco in 2023
Ikrandraco	I. avatar			A pteranodontoid[11]
Jidapterus	J. edentus			A chaoyangopterid[10]
Liaoningopterus	L. gui			A pteranodontoid[11]
Liaoxipterus	L. brachyognathus			An istiodactylid[10]
Linlongopterus	L. jennyae			A pteranodontoid[11]
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[10]
Pangupterus	P. liui			A pterodactyloid
Shenzhoupterus	S. chaoyangensis			A chaoyangopterid
Shenzhoupterus	S. sanyainus			A chaoyangopterid[13]
Sinopterus	S. dongi			A tapejarid[10]
Sinopterus	S. jii			Previously known as Huxiapterus jii

Vesperopterylus	V. lamadongensis			An anurognathid

Saurischians

A large titanosaur is present in the formation.[14]

Enantiornithines

Genus	Species	Material	Notes	Images
Alethoalaornis	Alethoalaornis agitornis[15]		An enantiornithean	Longipteryx Rapaxavis Sinornis Yuanchuavis
Boluochia	Boluochia zhengi	"Partial skull [and] partial postcranium."[16]	A longipterygid
Brevirostruavis	Brevirostruavis macrohyoideus	"an extremely elongate bony hyoid element (only slightly shorter than the skull), combined with a short cranial rostrum."[17]	An enantiornithine
Cathayornis	Cathayornis aberransis		An enantiornithine
	Cathayornis caudatus		Nomen dubium[18]
	Cathayornis yandica	"Partial skeleton."[16]	An enantiornithean
Cuspirostrisornis	Cuspirostrisornis houi		A possible avisaurid
Dapingfangornis	Dapingfangornis sentisorhinus		An enantiornithean
Eocathayornis	Eocathayornis walkeri	"Partial skeleton."[16]	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
Musivavis	Musivavis amabilis	"A nearly complete and articulated skeleton preserved in a single slab."[19]	An euenantiornithean
Rapaxavis	Rapaxavis pani		A longipterygid
Shangyang	Shangyang graciles		An enantiornithean
Sinornis	Sinornis santensis	"Partial skull [and] many nearly complete postcranial skeletons."[20]	An enantiornithean
Xiangornis	Xiangornis shenmi		An enantiornithean
Yuanchuavis	Yuanchuavis kompsosoura		An pengornithid

Euornithines

Genus	Species	County	Material	Notes	Images
Aberratiodontus	A. wui			Junior synonym of Yanornis martini[21]	Yanornis martini Yixianornis grabaui
Chaoyangia	C. beishanensis		"Partial skeleton."[20]	A basal euornithean
Jianchangornis	J. microdonta	Jianchang		A basal euornithean
Mengciusornis	M. dentatus			A schizoourid
Parahongshanornis[22]	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[23]

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[24]	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[25]	C. zhui	Jianchang		A complete skeleton with preserved feathers	A jinguofortisid avialam
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[26]
Jeholornis[3]	J. palmapenis[27]				A jeholornithiform avialan
Kompsornis	K. longicaudatus			"Nearly complete specimen".[28]	A jeholornithiform avialan
Microraptor	M. gui		Shangheshou Bed	Extremely common[14]	Could be synonym of Microraptor zhaoianus
Microraptor	M. zhaoianus		Shangheshou Bed[14]	"At least [three] partial semiarticulated postcrania and several skulls."[29] Feathered integument is also preserved.[29]	A dromaeosaurid.
Neimengornis	N. rectusmim				A jeholornithiform avialan
Omnivoropteryx	O. sinousaorum				An omnivoropterygiform avialan
Sapeornis	S. angustis			"Nearly complete skeleton without skull."[30]	An omnivoropterygiform, a primitive short-tailed avialan
Sapeornis	S. chaoyangensis				An omnivoropterygiform, a primitive short-tailed avialan
Shenshiornis	Shenshiornis primita[31]				An omnivoropterygiform avialan
Similicaudipteryx	S. yixianensis				An oviraptorosaur
Sinotyrannus[32]	S. kazuoensis[32]				A tyrannosauroid[32]
Wulong[33]	W. bohaiensis[33]	Chaoyang, Liaoning[33]		Complete articulated skeleton (D2933).[33]	A dromaeosaurid

Crustaceans

Genus	Species	County	Member	Material	Notes	Images
Eosestheria	Eosestheria middendorfii[34]				A clam shrimp

Plants

Genus	Species	Notes
Czekanowskia	Czekanowskia rigida[35]	A member of Czekanowskiales, Dominant
Ginkgo	Ginkgo digitata[35]	A member of Ginkgoales
Elatocladus	Elatocladus pinnatus[35]	Conifer foliage
Pityolepis	Pityolepis ovatus[35]	Conifer cone
Leptostrobus	Leptostrobus sinensis[35]	A member of Czekanowksiales, occasionally misspelled "Leptosthrobus"
Baiera	Baiera valida[35]	A member of Ginkgoales
Baiera	Baiera cf. gracilis[35]	A member of Ginkgoales
Ginkgoites	Ginkgoites trancata[35]	A member of Ginkgoales, dominant
Ginkgoites	Gingkgoites trancatus[35]	A member of Ginkgoales, dominant
Podozamites	Podozamites sp.[34]	Conifer foliage
Sphenopteris	Sphenopteris sp.[34]	Seed plant foliage

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