Jiufotang Formation
Geological formation in Liaoning, China
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The Jiufotang Formation (Chinese: 九佛堂组; pinyin: jiǔfótáng zǔ; previously transliterated as Chiufotang Formation[2]) 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.[3] Fossils of Microraptor and Jeholornis are from the Jiufotang.[4] The fossil layers containing lagersatte conditions date from 124.44 to 121.09 Ma.
| Jiufotang Formation | |
|---|---|
| Stratigraphic range: Barremian–Albian | |
| 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) | |
Fossil content
Choristoderans
| Genus | Species | County | Member | Abundance | Notes |
|---|---|---|---|---|---|
| Philydrosaurus | P. proseilus | Monjurosuchid | |||
| Ikechosaurus | I. pijiagouensis | Simoedosaurid | |||
| Liaoxisaurus | L. chaoyangensis | Simoedosaurid | |||
| Hyphalosaurus | H. baitaigouensis[5] | Hyphalosaurid | |||
Fish
| Genus | Species | County | Member | Notes | Images |
|---|---|---|---|---|---|
| Caeruleum | C. miraculum[6] | Weichang | A lamprey |  | |
| C. gracilis[7] | Weichang | A lamprey | |||
| Hebeimyzon[8] | H. weichangensis | 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[9] | Partial skeleton | A eutherian | ||
| Fossiomanus | F. sinensis[10] | A nearly complete skeleton | A tritylodontid cynodont, likely the youngest known non-mammalian synapsid | ||
| Liaoconodon | L. hui[11] | 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 | Locality | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|---|
|
C. chaoyangensis[12] |
Liaoning | A nearly complete skeleton missing only the distal portion of the caudal series. |
A basal ankylosaurid, sister taxon to Liaoningosaurus.[12] |
   | |||
| Liaoningosaurus[13] | L. paradoxus[13] | Baicaigou locality | Liaoning | A nearly complete, articulated skeleton | A basal ankylosaurid. | ||
|
P. meileyingensis |
Meileyingzi Bed |
"[Four] individuals, [two] complete skulls."[14] |
A ceratopsian. | ||||
|
P. mongoliensis |
Shangheshou and Meileyingzi Beds |
||||||
Pterosaurs
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
C. zhangi |
A chaoyangopterid[15] |
       | ||||
|
E. liaoxiensis |
A chaoyangopterid[15] | |||||
|
F. youngi |
Jianshangou Bed |
A boreopterid | ||||
| Forfexopterus | F. jeholensis | A ctenochasmatid | ||||
|
G. jingangshanensis |
Jingangshan Bed |
A ctenochasmatid | ||||
| Guidraco |
G. venator |
|||||
| Hongshanopterus | H. lacustris | An istiodactylid | ||||
| Huaxiadraco | H. corollatus | Multiple specimens | A tapejarid[17] | |||
|
"H." benxiensis |
A tapejarid; synonymized with Huaxiadraco in 2023 | |||||
| Ikrandraco |
I. avatar |
|||||
|
J. edentus |
A chaoyangopterid[15] | |||||
|
L. gui |
||||||
|
L. brachyognathus |
An istiodactylid[15] | |||||
|
L. jennyae |
||||||
| Meilifeilong | M. youhao | Jianchang | A nearly complete skeleton and a partial skull | A chaoyangopterid[18] | ||
|
N. crypticus |
A tapejaromorph; previously and still occasionally thought to be a tapejarid. Most likely a juvenile, hence small size. | |||||
|
N. ignaciobritoi |
An istiodactylid[15] | |||||
| Pangupterus | P. liui | A pterodactyloid | ||||
|
S. chaoyangensis |
A chaoyangopterid | |||||
| S. sanyainus | A chaoyangopterid[19] | |||||
|
S. dongi |
A tapejarid[15] | |||||
|
S. jii |
Previously known as Huxiapterus jii | |||||
|
V. lamadongensis |
An anurognathid | |||||
Saurischians
Skeletal[20] and dental[21] remains are known, the latter was assigned to an indeterminate titanosauriform.
Enantiornithines
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
Alethoalaornis |
A. agitornis[22] |
   .png)    | ||||
|
B. zhengi |
"Partial skull [and] partial postcranium."[23] |
A longipterygid | ||||
| Brevirostruavis | B. macrohyoideus | "an extremely elongate bony hyoid element (only slightly shorter than the skull), combined with a short cranial rostrum."[24] | An enantiornithine | |||
|
C. aberransis |
An enantiornithine | |||||
|
C. caudatus |
Nomen dubium[25] | |||||
|
C. yandica |
"Partial skeleton."[23] |
An enantiornithean | ||||
|
C. houi |
A possible avisaurid | |||||
|
D. sentisorhinus |
An enantiornithean | |||||
|
E. walkeri |
"Partial skeleton."[23] |
An enantiornithean | ||||
|
G. jiufotangensis |
An enantiornithean | |||||
| Imparavis[26] | I. attenboroughi | Jianchang | An articulated, nearly complete specimen | |||
|
H. huji |
An enantiornithean | |||||
|
L. sexdentoris |
An enantiornithean | |||||
|
L. sanyanensis |
An enantiornithean | |||||
|
L. chaoyangensis |
A longipterygid | |||||
| Musivavis | M. amabilis | "A nearly complete and articulated skeleton preserved in a single slab."[27] | An euenantiornithean | |||
| Plumadraco[28] | P. bankoorum | Near Xiaotaizi Village, Jianchang County | A complete, articulated specimen preserved in a single slab primarily in dorsal aspect with feathers preserved around the head, body, wings, and tail | A bohaiornithid | ||
|
R. pani |
A longipterygid | |||||
|
S. graciles |
An enantiornithean | |||||
|
S. santensis |
"Partial skull [and] many nearly complete postcranial skeletons."[29] |
An enantiornithean | ||||
|
X. shenmi |
An enantiornithean | |||||
|
Y. kompsosoura |
An pengornithid | |||||
Euornithines
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
A. wui |
Junior synonym of Yanornis martini[30] |
      | ||||
|
C. beishanensis |
"Partial skeleton."[29] |
A basal euornithean | ||||
|
I. huchzermeyeri |
Multiple complete, articulated skeletons |
A possible gansuid basal euornithean. Some authors consider Gansus zheni, another species named from the same locality, to be a junior synonym.[31][32] | ||||
|
J. microdonta |
A basal euornithean | |||||
| Kunpengornis[33] | K. anhuimusei | Jianchang | A nearly complete and articulated skeleton | A piscivorous euornithean | ||
|
M. dentatus |
||||||
|
P. chaoyangensis |
||||||
|
P. lii |
||||||
|
S. lii |
A schizoourid with long, forked tail feathers | |||||
|
S. brevipectus |
An ornithuromorph | |||||
|
S. angelai |
A gansuid | |||||
|
S. linghensis |
An ornithuromorph | |||||
|
Y. martini |
An ornithuromorph | |||||
|
Y. grabaui |
A near complete skeleton |
An ornithuromorph | ||||
|
Z. yangi |
A complete skeleton |
A basal euornithean[36] | ||||
Non-ornithothoracean theropods
A large theropod tooth representing a distinct taxon from Sinotyrannus is known.[21]
Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
| Changzhousaurus[37] | C. sinensis | Jianchang | A nearly complete skeleton with extensive plumage, preserved on a slab and counterslab | A paravian of uncertain affinities derived from a pennaraptoran |       | |
| Chongmingia[38] | C. zhengi | A partial skeleton with preserved soft tissues | A jinguofortisid avialan | |||
|
C. sanctus |
A primitive short-tailed avialan (pygostylian). Also present in the Yixian Formation | |||||
|
C. jianchangensis |
||||||
| Cratonavis[39] | C. zhui | Jianchang | A complete skeleton with preserved feathers | A jinguofortisid avialan | ||
|
D. cuhe |
Nomen dubium, named from a forged artificially assembled specimen. | |||||
|
J. prima[4] |
A jeholornithiform avialan, also present in the Yixian Formation[40] | |||||
|
J. palmapenis[41] |
A jeholornithiform avialan | |||||
|
K. longicaudatus |
"Nearly complete specimen".[42] |
A jeholornithiform avialan | ||||
|
M. gui |
Shangheshou Bed |
Extremely common[20] |
Could be synonym of Microraptor zhaoianus | |||
|
M. zhaoianus |
Shangheshou Bed[20] |
"At least [three] partial semiarticulated postcrania and several skulls."[43] Feathered integument is also preserved.[43] |
A dromaeosaurid. | |||
|
N. rectusmim |
A jeholornithiform avialan | |||||
|
O. sinousaorum |
An omnivoropterygiform avialan | |||||
|
S. angustis |
"Nearly complete skeleton without skull."[44] |
An omnivoropterygiform, a primitive short-tailed avialan | ||||
|
S. chaoyangensis |
||||||
|
Shenshiornis primita[45] |
An omnivoropterygiform avialan | |||||
|
S. yixianensis |
||||||
|
S. kazuoensis[46] |
||||||
| Wulong[47] | W. bohaiensis[47] | Chaoyang, Liaoning[47] | Complete articulated skeleton (D2933).[47] | A dromaeosaurid | ||
Crustaceans
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
| Eosestheria | Eosestheria middendorfii[48] | A clam shrimp |
Plants
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
| Czekanowskia | C. rigida[49] | A member of Czekanowskiales, Dominant | ||||
| Ginkgo | G. digitata[49] | A member of Ginkgoales | ||||
| Elatocladus | E. pinnatus[49] | Conifer foliage | ||||
| Pityolepis | P. ovatus[49] | Conifer cone | ||||
| Leptostrobus | L. sinensis[49] | A member of Czekanowksiales, occasionally misspelled "Leptosthrobus" | ||||
| Baiera | B. valida[49] | A member of Ginkgoales | ||||
| B. cf. gracilis[49] | ||||||
| Ginkgoites | G. trancata[49] | A member of Ginkgoales, dominant | ||||
| G. trancatus[49] | ||||||
| Podozamites | P. sp.[48] | Conifer foliage | ||||
| Sphenopteris | S. sp.[48] | Seed plant foliage | ||||