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°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

Fish

Genus	Species	County	Notes
Caeruleum^[4]	C. miraculum	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^[5]	Partial skeleton	A eutherian
Fossiomanus	F. sinensis^[6]	A nearly complete skeleton	A tritylodontid cynodont, likely the youngest known non-mammalian synapsid
Liaoconodon	L. hui^[7]	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^[8]	C. chaoyangensis^[8]	Liaoning		A nearly complete skeleton missing only the distal portion of the caudal series.	A basal ankylosaurid, sister taxon to Liaoningosaurus.^[8]
Liaoningosaurus^[9]	L. paradoxus^[9]					Liaoningosaurus paradoxus
Psittacosaurus	Psittacosaurus meileyingensis		Meileyingzi Bed	"[Four] individuals, [two] complete skulls."^[10]	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^[11]	Liaoningopterus Nemicolopterus Nurhachius Sinopterus and Huaxiadraco
Eoazhdarcho	E. liaoxiensis				A chaoyangopterid^[11]
Feilongus	F. youngi		Jianshangou Bed		A boreopterid
Forfexopterus	F. jeholensis				A ctenochasmatid
Gladocephaloideus	G. jingangshanensis		Jingangshan Bed		A ctenochasmatid
Guidraco	G. venator				A pteranodontoid^[12]
Hongshanopterus	H. lacustris				An istiodactylid
Huaxiadraco	H. corollatus			Multiple specimens	A tapejarid^[13]
"Huaxiapterus"	"H." benxiensis				A tapejarid; synonymized with Huaxiadraco in 2023
Ikrandraco	I. avatar				A pteranodontoid^[12]
Jidapterus	J. edentus				A chaoyangopterid^[11]
Liaoningopterus	L. gui				A pteranodontoid^[12]
Liaoxipterus	L. brachyognathus				An istiodactylid^[11]
Linlongopterus	L. jennyae				A pteranodontoid^[12]
Meilifeilong	M. youhao	Jianchang		A nearly complete skeleton and a partial skull	A chaoyangopterid^[14]
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^[11]
Pangupterus	P. liui				A pterodactyloid
Shenzhoupterus	S. chaoyangensis				A chaoyangopterid
Shenzhoupterus	S. sanyainus				A chaoyangopterid^[15]
Sinopterus	S. dongi				A tapejarid^[11]
Sinopterus	S. jii				Previously known as Huxiapterus jii

Vesperopterylus	V. lamadongensis				An anurognathid

Saurischians

A large titanosaur is present in the formation.^[16]

Enantiornithines

Genus	Species	Material	Notes	Images
Alethoalaornis	Alethoalaornis agitornis^[17]		An enantiornithean	Longipteryx Rapaxavis Sinornis Yuanchuavis
Boluochia	Boluochia zhengi	"Partial skull [and] partial postcranium."^[18]	A longipterygid
Brevirostruavis	Brevirostruavis macrohyoideus	"an extremely elongate bony hyoid element (only slightly shorter than the skull), combined with a short cranial rostrum."^[19]	An enantiornithine
Cathayornis	Cathayornis aberransis		An enantiornithine
	Cathayornis caudatus		Nomen dubium^[20]
	Cathayornis yandica	"Partial skeleton."^[18]	An enantiornithean
Cuspirostrisornis	Cuspirostrisornis houi		A possible avisaurid
Dapingfangornis	Dapingfangornis sentisorhinus		An enantiornithean
Eocathayornis	Eocathayornis walkeri	"Partial skeleton."^[18]	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."^[21]	An euenantiornithean
Rapaxavis	Rapaxavis pani		A longipterygid
Shangyang	Shangyang graciles		An enantiornithean
Sinornis	Sinornis santensis	"Partial skull [and] many nearly complete postcranial skeletons."^[22]	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^[23]	Yanornis martini Yixianornis grabaui
Chaoyangia	C. beishanensis		"Partial skeleton."^[22]	A basal euornithean
Jianchangornis	J. microdonta	Jianchang		A basal euornithean
Mengciusornis	M. dentatus			A schizoourid
Parahongshanornis^[24]	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^[25]

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^[26]	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^[27]	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^[28]
Jeholornis^[3]	J. palmapenis^[29]				A jeholornithiform avialan
Kompsornis	K. longicaudatus			"Nearly complete specimen".^[30]	A jeholornithiform avialan
Microraptor	M. gui		Shangheshou Bed	Extremely common^[16]	Could be synonym of Microraptor zhaoianus
Microraptor	M. zhaoianus		Shangheshou Bed^[16]	"At least [three] partial semiarticulated postcrania and several skulls."^[31] Feathered integument is also preserved.^[31]	A dromaeosaurid.
Neimengornis	N. rectusmim				A jeholornithiform avialan
Omnivoropteryx	O. sinousaorum				An omnivoropterygiform avialan
Sapeornis	S. angustis			"Nearly complete skeleton without skull."^[32]	An omnivoropterygiform, a primitive short-tailed avialan
Sapeornis	S. chaoyangensis				An omnivoropterygiform, a primitive short-tailed avialan
Shenshiornis	Shenshiornis primita^[33]				An omnivoropterygiform avialan
Similicaudipteryx	S. yixianensis				An oviraptorosaur
Sinotyrannus^[34]	S. kazuoensis^[34]				A tyrannosauroid^[34]
Wulong^[35]	W. bohaiensis^[35]	Chaoyang, Liaoning^[35]		Complete articulated skeleton (D2933).^[35]	A dromaeosaurid

Crustaceans

Genus	Species	County	Member	Material	Notes	Images
Eosestheria	Eosestheria middendorfii^[36]				A clam shrimp

Plants

Genus	Species	Notes
Czekanowskia	Czekanowskia rigida^[37]	A member of Czekanowskiales, Dominant
Ginkgo	Ginkgo digitata^[37]	A member of Ginkgoales
Elatocladus	Elatocladus pinnatus^[37]	Conifer foliage
Pityolepis	Pityolepis ovatus^[37]	Conifer cone
Leptostrobus	Leptostrobus sinensis^[37]	A member of Czekanowksiales, occasionally misspelled "Leptosthrobus"
Baiera	Baiera valida^[37]	A member of Ginkgoales
Baiera	Baiera cf. gracilis^[37]	A member of Ginkgoales
Ginkgoites	Ginkgoites trancata^[37]	A member of Ginkgoales, dominant
Ginkgoites	Gingkgoites trancatus^[37]	A member of Ginkgoales, dominant
Podozamites	Podozamites sp.^[36]	Conifer foliage
Sphenopteris	Sphenopteris sp.^[36]	Seed plant foliage

References

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