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	41.2°N 119.4°E / 41.2; 119.4
Approximate paleocoordinates	44.4°N 119.5°E / 44.4; 119.5
Region	Liaoning
Country	China
Type section
Named for	Jiufotang, Pingfangzi, Liaoning
Named by	Ryuji Endo
Year defined	1934[1]
Jiufotang Formation (China) Show map of China Jiufotang Formation (Liaoning) Show map of 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	County	Member	Abundance	Notes
Philydrosaurus	P. proseilus				Monjurosuchid
Ikechosaurus	I. pijiagouensis				Simoedosaurid
Liaoxisaurus	L. chaoyangensis				Simoedosaurid
Hyphalosaurus	H. baitaigouensis[4]				Hyphalosaurid

Fish

Genus	Species	County	Member	Notes	Images
Caeruleum	C. miraculum[5]	Weichang		A lamprey
	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	County	Member	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 mongoliensis		Shangheshou and Meileyingzi Beds

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
	S. sanyainus				A chaoyangopterid[17]
Sinopterus	S. dongi				A tapejarid[13]
	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	Member	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	Member	Material	Notes	Images
Aberratiodontus	A. wui				Junior synonym of Yanornis martini[26]	Gansus zheni Shuilingornis angelai Yanornis martini Yixianornis grabaui
Chaoyangia	C. beishanensis			"Partial skeleton."[25]	A basal euornithean
Gansus[27]	G. zheni			Multiple complete, articulated skeletons	A basal euornithean
Jianchangornis	J. microdonta	Jianchang			A basal euornithean
Mengciusornis	M. dentatus				A schizoourid
Parahongshanornis[28]	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
Shuilingornis[29]	S. angelai				A gansuid
Songlingornis	S. linghensis				An ornithuromorph
Yanornis	Y. martini				An ornithuromorph
Yixianornis	Y. grabaui				An ornithuromorph
Zhongjianornis	Z. yangi				A basal euornithean[30]

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[31]	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
	C. jianchangensis	Liaoning
Cratonavis[32]	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[33]
	J. palmapenis[34]				A jeholornithiform avialan
Kompsornis	K. longicaudatus			"Nearly complete specimen".[35]	A jeholornithiform avialan
Microraptor	M. gui		Shangheshou Bed	Extremely common[18]	Could be synonym of Microraptor zhaoianus
	M. zhaoianus		Shangheshou Bed[18]	"At least [three] partial semiarticulated postcrania and several skulls."[36] Feathered integument is also preserved.[36]	A dromaeosaurid.
Neimengornis	N. rectusmim				A jeholornithiform avialan
Omnivoropteryx	O. sinousaorum				An omnivoropterygiform avialan
Sapeornis	S. angustis			"Nearly complete skeleton without skull."[37]	An omnivoropterygiform, a primitive short-tailed avialan
	S. chaoyangensis
Shenshiornis	Shenshiornis primita[38]				An omnivoropterygiform avialan
Similicaudipteryx	S. yixianensis				An oviraptorosaur
Sinotyrannus[39]	S. kazuoensis[39]				A tyrannosauroid[39]
Wulong[40]	W. bohaiensis[40]	Chaoyang, Liaoning[40]		Complete articulated skeleton (D2933).[40]	A dromaeosaurid

Crustaceans

Genus	Species	County	Member	Material	Notes	Images
Eosestheria	Eosestheria middendorfii[41]				A clam shrimp

Plants

Genus	Species	County	Member	Material	Notes	Images
Czekanowskia	Czekanowskia rigida[42]				A member of Czekanowskiales, Dominant
Ginkgo	Ginkgo digitata[42]				A member of Ginkgoales
Elatocladus	Elatocladus pinnatus[42]				Conifer foliage
Pityolepis	Pityolepis ovatus[42]				Conifer cone
Leptostrobus	Leptostrobus sinensis[42]				A member of Czekanowksiales, occasionally misspelled "Leptosthrobus"
Baiera	Baiera valida[42]				A member of Ginkgoales
	Baiera cf. gracilis[42]
Ginkgoites	Ginkgoites trancata[42]				A member of Ginkgoales, dominant
	Gingkgoites trancatus[42]
Podozamites	Podozamites sp.[41]				Conifer foliage
Sphenopteris	Sphenopteris sp.[41]				Seed plant foliage

See also

• Yixian Formation
• List of dinosaur-bearing rock formations

References

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Further reading

• Wang, Y; Evans, SE (2011). "A gravid lizard from the Cretaceous of China and the early history of squamate viviparity". Naturwissenschaften. 98 (9): 739–743. Bibcode:2011NW.....98..739W. doi:10.1007/s00114-011-0820-1. PMID 21766177. S2CID 8017857.