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Paleobiota of the Yixian Formation

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Ordosemys liaoxiensis and Psittacosaurus preserved together

The Yixian Formation (simplified Chinese: 义县组; traditional Chinese: 義縣組; pinyin: Yìxiàn zǔ) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans about 1.6 million years during the early Cretaceous period (Barremian-Aptian stage). It is known for its fossils, listed below.

The Yixian Formation is divided into the following subunits: (ages from Zhong et al., 2021[1] unless otherwise noted)

  • Upper Yixian (124.1 Ma) ("Jingangshan Bed") ("Dawangzhangzi Bed")
  • Upper Lava Unit
  • Jianshangou Unit (125.46 Ma)
  • Lower Lava Unit
  • Lujiatun Unit (125.76–125.68 Ma)

Sauropods

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Genus Species County Member Material Notes Images

Dongbeititan

D. dongi

Beipiao Jianshangou Bed

Partial postcrania[2]

A basal titanosauriform

Euhelopus

cf. Euhelopus

cf. E. sp.

Lujiatun Bed

Isolated teeth[3][4]

A basal titanosauriform

Liaoningotitan

L. sinensis

nearly complete skeleton[5]

A basal titanosauriform

Ruixinia R. zhangi Beipiao A partial articulated skeleton[6] A somphospondylan titanosauriform

Sauropoda spp.

Lujiatun & Jianshangou Beds

Isolated vertebrae and teeth[7][4]

One or more species of sauropods, possibly titanosaurs, have been recovered from the Yixian Formation, though most remain undescribed[3]

Ornithischians

[edit]

The dinosaurs listed below were compiled in a survey by Xu and Norell in 2006, unless otherwise noted.[7]

Ankylosaurs

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Genus Species County Member Material Notes Images

Liaoningosaurus

L. paradoxus

Yixian

Dawangzhangzi Bed

"Partial skull [and] skeleton."[8]

A small ankylosaur
Liaoningosaurus

Ceratopsians

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Genus Species County Member Material Notes Images

Liaoceratops

L. yanzigouensis

Beipiao

Lujiatun Bed

"[Two] nearly complete skulls, juvenile to adult."[9]

A ceratopsian

Liaoceratops
Psittacosaurus

Psittacosaurus

P. lujiatunensis

Beipiao

Lujiatun & Jianshangou Beds

Thousands of specimens in the Lujiatun Bed, somewhat less common in the Jianshangou Bed

A ceratopsian

Ornithopods

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Genus Species County Member Material Notes Images

Bolong

B. yixianensis

Jianshangou Bed

A nearly complete skeleton.

An iguanodontian ornithopod

Bolong
Jeholosaurus
Jinzhousaurus
Changmiania[10] C. liaoningensis Lujiatun Bed Two nearly complete skeletons A basal ornithopod that may have dug burrows

Jeholosaurus

J. shangyuanensis

Beipiao

Lujiatun Bed

"Skull with associated vertebrae and hindlimb, skull with cervical vertebrae."[11]

An ornithopod

Jinzhousaurus

J. yangi

Yixian

Dawangzhangzi Bed

"Nearly complete skull and skeleton."[12]

An ornithopod

Theropods

[edit]

The dinosaurs listed below were compiled in a survey by Xu and Norell in 2006, unless otherwise noted.[7]

Avialans

[edit]
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
Abitusavis[13] A. lii Ningcheng "Nearly complete skeleton."[13]
Changchengornis
Confuciusornis sanctus
Eogranivora
Hongshanornis
Liaoningornis
Liaoxiornis
Shanweiniao
Yixianosaurus

Archaeorhynchus

A. spathula

Dawangzhangzi Bed

An early, advanced short-tailed bird (ornithuran)[14]

Changchengornis[15]

C. hengdaoziensis[15]

Jianshangou Bed

"Nearly complete skeleton."[15]

A confuciusornithid

Confuciusornis

C. chuonzhous

Junior synonym of Confuciusornis sanctus[16]

C. dui[17]

Jianshangou Bed

"[Two] skeletons."[17]

C. feducciai[18]

Jianshangou Bed

C. sanctus[17]

Jianshangou and Dawangzhangzi Beds

"Hundreds of complete skeletons."[17]

A primitive short-tailed bird (pygostylian), also present in the Jiufotang Formation

C. suniae

Junior synonym of Confuciusornis sanctus[16]

Dalingheornis

D. liweii

Dawangzhangzi Bed

An Enantiornithine[19]

Eoenantiornis

E. buhleri

Jianshangou Bed

"Nearly complete skeleton."[20]

An entantiornithine

Eogranivora

E. edentulata

Dawangzhangzi Bed

"nearly complete skeleton with skull"[21]

An ornithuromorph

Iteravis

I. huchzermeyeri

An ornithuromorph

Jinzhouornis

J. yixianensis

Jianshangou Bed

A pygostylian, a junior synonym of Confuciusornis sanctus[22]

J. zhangjiyingia

Jianshangou Bed

Junior synonym of C. sanctus

Jixiangornis

J. orientalis

Jianshangou Bed

A basal avialan, also present in the Jiufotang Formation[23]

Hongshanornis

H. longicresta

Jianshangou and Dawangzhangzi Beds[24]

4 specimens[24]

An ornithuromorph[25]

Jibeinia

J. luahera

Previously regarded as Yixian, horizon reassigned to the Huajiying Formation by Jin et al., 2008.[26]

Liaoningornis

L. longidigitris

Lujiatun Bed

"Partial skeleton."[27]

An enantiornithine.[28]

Liaoxiornis

L. delicatus

Dawangzhangzi Bed

Considered a nomen dubium because the name is based on remains of a hatchling entantiornithine and cannot be confidently matched with the adult form.[29]

Lingyuanornis

L. parvus

Dawangzhangzi Bed

Junior objective synonym of Liaoxiornis delicatus, a nomen dubium.[29]

Longicrusavis

L. houi

Dawangzhangzi Bed

An ornithuromorph[25]

Longirostravis

L. hani

bed unknown

An enantiornithine

Protopteryx

P. fengningensis

Previously regarded as Yixian, horizon reassigned to the Huajiying Formation by Jin et al., 2008.[26]

Mirusavis

M. parvus

Partial skeleton.[30]

An immature female enantiornithine preserved with medullary bone tissue.[30]

Sapeornis[31] S. chaoyangensis Nearly comeplete skeleton.[31] A subadult omnivoropterygid with preserved feathers.[31]

Shanweiniao

S. cooperorum

Dawangzhangzi Bed

An enantiornithine.[32]

Vescornis

V. hebeiensis

Previously regarded as Yixian, horizon reassigned to the Huajiying Formation by Jin et al., 2008.[26]

Yixianosaurus Y. longimanus Dawangzhangzi Bed An anchiornithid

Zhongornis

Z. haoae

Dawangzhangzi Bed

A primitive avialan with a short but non-pygostylian tail[33]

Compsognathids

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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

Huaxiagnathus

H. orientalis

Beipiao

Jianshangou Bed

A compsognathid

Sinocalliopteryx
Sinosauropteryx prima

Sinocalliopteryx

S. gigas

Beipiao

Jianshangou Bed

A large compsognathid

Sinosauropteryx

S. prima

Beipiao & Lingyuan

Jianshangou & Dawangzhangzi Beds

Four nearly complete specimens[34][35]

A compsognathid. The first non-avian dinosaur found with feather impressions.

S.? sp.

Beipiao

Jianshangou Bed

One specimen (GMV 2124) with partial skull, postcrania, and preserved feathers

An unnamed coelurosaur originally described as a specimen of Sinosauropteryx, later found to be a new genus and species.[36]

Dromaeosaurs

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Genus Species County Member Material Notes Images

Changyuraptor[37]

C. yangi

Jianchang

A complete articulated specimen with extensive feathering.[37]

A dromaeosaurid

Changyuraptor
Graciliraptor
Sinornithosaurus millenii
Tianyuraptor
Zhenyuanlong
Zhongjianosaurus

Graciliraptor

G. lujiatunensis

Beipiao

Lujiatun Bed

An incomplete specimen.

A dromaeosaurid

Sinornithosaurus

S. haoiana

Yixian

Dawangzhangzi Bed

Might be a junior synonym of Sinornithosaurus millenii.

S. millenii

Beipiao

Jianshangou Bed

Three specimens with skulls and postcrania, two with preserved feathers.[38] Feathered integument is also preserved.[38][39]

A dromaeosaurid, the first ever found with feather impressions.

Tianyuraptor

T. ostromi

Lingyuan

Dawangzhangzi Bed

A dromaeosaurid

Zhenyuanlong[40]

Z. suni

Jianchang

One nearly complete skeleton with preserved feathers, including long tail feathers and large wing feathers.

A dromaeosaurid

Zhongjianosaurus[41]

Z. yangi

Lingyuan

A partial skeleton includes an arm, legs, vertebrae, ribs, and most of the tail.[41]

A microraptorian dromaeosaurid with very long legs.[41]

Unnamed dromaeosaurid

Specimen NGMC 91

Lingyuan

Dawangzhangzi Bed

Complete specimen with feather imprints

A juvenile dromaeosaurid that may be a specimen of Sinornithosaurus millenii.[42][43] Informally nicknamed "Dave".



Ornithomimosaurs

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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

Hexing

H. qingyi

Beipiao

Lujiatun

A primitive ornithomimosaur[44]

Shenzhousaurus

Shenzhousaurus

S. orientalis

Beipiao

Lujiatun Bed

"Articulated skull and incomplete skeleton."[45]

A primitive ornithomimosaur

Oviraptorosaurs

[edit]
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

Caudipteryx

C. dongi

Beipiao

Jianshangou Bed

"[One] partial postcranium with feather imprints."[46]

Oviraptorosaurs

Caudipteryx zoui
Incisivosaurus
Protarchaeopteryx

C. zoui

Beipiao

Jianshangou Bed

Common, only over a small region

C. sp.[47]

Incisivosaurus

I. gauthieri

Beipiao

Lujiatun Bed

"[One] skull [and a] partial cervical."[46]

An oviraptorosaur

Ningyuansaurus

N. wangi

Complete skeleton with feather impressions.[48]

An oviraptorosaur

Protarchaeopteryx

P. robusta

Beipiao

Jianshangou Bed

"Nearly complete skull and skeleton, at least [two] individuals."[49]

A possible relative of Incisivosaurus

Similicaudipteryx

S. yixianensis

Two specimens

An oviraptorosaur, also present in the Jiufotang Formation[50]

Xingtianosaurus

X. ganqi

Liaoning

Dakangpu Bed

One nearly complete specimen (IVPP V13390), minus the head

An oviraptorosaur[51]

Therizinosaurs

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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

Beipiaosaurus

B. inexpectus

Beipiao

Jianshangou Bed

"Incomplete skeleton with integument."[52]

A therizinosaur

Beipiaosaurus
Jianchangosaurus
Jianchangosaurus J. yixianensis Jianchang A nearly complete juvenile specimen with feather impressions.[53] A therizinosaur

Troodontids

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Genus Species County Member Material Notes Images
Daliansaurus D. liaoningensis Beipiao Lujiatun Bed A very complete specimen.[54] A troodontid
Daliansaurus
Jianianhualong
Liaoningvenator
Mei
Sinovenator
Sinusonasus

Jianianhualong

J. tengi

Beipiao

Dawangzhangzi Bed

A very complete specimen with feather impressions.[55]

A troodontid

Liaoningvenator L. curriei Beipiao A very complete specimen. A troodontid

Mei

M. long

Beipiao

Lujiatun Bed

A small troodontid fossilized in a sleeping position

Sinovenator

S. changii

Beipiao

Lujiatun Bed

Three specimens with partial skulls and postcrania.[56][57] Hundreds of additional specimens known but undescribed.

A troodontid

Sinusonasus

S. magnodens

Beipiao

Lujiatun Bed

A troodontid

Undescribed deinonychosaur/possible troodontid

Dawangzhangzi Bed

One specimen with feather imprints

A possible troodontid.[58]

Undescribed troodontid

Specimen CAGS-IG01-004

Lujiatun Bed

Skull and partial skeleton

A troodontid similar to Mei.[59]

Tyrannosaurs and other theropods

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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

Dilong

D. paradoxus

Beipiao

Lujiatun & Jianshangou Bed

Rare

A tyrannosauroid

Dilong
Yutyrannus

Grallator

lower Jianshangou Bed

Footprints belonging to the Grallator form taxon, belonging to a small theropod. Of known Yixian theropods, the footprints most closely match Caudipteryx.[60]

Raptorex

R. kriegsteini

A tyrannosaurid, probably from higher Mongolian strata.

Scansoriopteryx

S. heilmanni

Initially reported from the Dawangzhangzi Bed, probably from the Daohugou Beds instead.[61]

Yutyrannus

Y. huali

Unknown

Three nearly complete skeletons of various ages

A large tyrannosauroid[62]

Unnamed theropod

Not yet described

Yixian

Lujiatun Bed

Partial post-cranial skeleton[63]

A possible carnosaur[64]

Pterosaurs

[edit]

The pterosaurs listed below were compiled in a survey by Wang and Zhou in 2006, unless otherwise noted.[65]

Genus Species County Member Material Notes Images

Beipiaopterus

B. chenianus

Jianshangou Bed

A ctenochasmatid
Feilongus
Gladocephaloideus
Haopterus
Moganopterus
Ningchengopterus

Boreopterus

B. cuiae

Jianshangou Bed

A boreopterid

Cathayopterus

C. grabaui

Dawangzhangzi Bed

A ctenochasmatid

Elanodactylus[66]

E. prolatus

Jianshangou Bed[67]

A ctenochasmatid
Eopteranodon E. lii[68] Beipiao A tapejarid
E. yixianensis[69] Beipiao Jianshangou Bed A tapejarid

Eosipterus

E. yangi

Jianshangou Bed

A ctenochasmatid

Gegepterus[70]

G. changae[67]

Jianshangou Bed[67]

A ctenochasmatid

Gladocephaloideus

G. jingangshanensis

Jianshangou Bed

A ctenochasmatid
Haopterus

H. gracilis

Jianshangou Bed

An ornithocheirid
Luchibang[71] L. xingzhe One nearly complete articulated skeleton (ELDM 1000) An istiodactylid, the most complete known

Moganopterus

M. zhuiana

Lujiatun bed?

A ctenochasmatid

Ningchengopterus

N. liuae

Jianshangou or Dawangzhangzi bed

A pterodactyloid[72]
Nurhachius N. sp. Jingangshan Member[73] An incomplete specimen[73] An istiodactylid

Pterofiltrus

P. qiui

Jianshangou Bed

A ctenochasmatid[67]
Yixianopterus Y. jingangshanensis An istiodactyliform

Zhenyuanopterus

Z. longirostris

Jianshangou Bed

A boreopterid.[74] May be adult form of Boreopterus.[75]

Misc. reptiles

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Choristoderans

[edit]
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

Hyphalosaurus

H. lingyuanensis

Dawangzhangzi Bed

Hyphalosaurus baitaigouensis
Monjurosuchus splendens

H. baitaigouensis

Jingangshan Bed

Thousands of specimens

Monjurosuchus

M. splendens

Dawangzhangzi Bed

Sinohydrosaurus

S. lingyuanensis

Dawangzhangzi Bed

Junior synonym of Hyphalosaurus lingyuanensis

Lizards

[edit]
Genus Species County Member Material Notes Images

Dalinghosaurus

D. longidigitus

Jianshangou Bed

A lizard related to shinisaurs
Dalinghosaurus
Xianglong

Liushusaurus

L. acanthocaudata[76]

Jianshangou or Dawangzhangzi bed

A lizard related to geckos and skinks

Xianglong

X. zhaoi

Jianshangou Bed

A gliding lizard related to chameleons and dragon lizards

Yabeinosaurus

Y. tenuis

Dawangzhangzi & Jingangshan Beds

A lizard related to geckos and skinks

Turtles

[edit]
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

Manchurochelys

M. liaoxiensis[77]

Jianshangou Bed

Reclassified as Ordosemys liaoxiensis[78]

Ordosemys

M. manchoukuoensis[16]

Jingangshan Bed

Ordosemys

O. liaoxiensis[78]

Jianshangou Bed

Mammals

[edit]
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.

The mammals listed below were compiled in a survey by Meng and colleagues in 2006, unless otherwise noted.[79]

Genus Species County Member Material Notes Images

Acristatherium

A. yanensis[80]

Beipiao

Lujiatun Bed

A eutherian
Eomaia
Gobiconodon zofiae
Jeholodens
Repenomamus robustus
Yanoconodon

Akidolestes

A. cifellii

Lingyuan

Dawangzhangzi Bed

A spalacotheriid "symmetrodont"
Ambolestes[81] A. zhoui Nincheng Jianshangou Bed A eutherian

Chaoyangodens[82]

C. lii

Lingyuan

Dawangzhangzi Bed

A eutriconodont

Eomaia

E. scansoria

Lingyuan

Dawangzhangzi Bed

A tribosphenidan, possibly a eutherian or stem-therian

Gobiconodon

G. zofiae

Beipiao

Lujiatun Bed

A gobiconodont

G. luoianus[83]

Lujiatun Bed

A gobiconodont; junior synonym of G. zofiae

Jeholodens

J. jenkinsi

Chaoyang

Jianshangou Bed

A eutriconodont

Juchilestes[84]

J. liaoningensis

Beipiao

Lujiatun Bed

An amphidontid

Jueconodon[85]

J. cheni

A eutriconodont

Maotherium

M. sinensis

Chaoyang

Jianshangou Bed

A zhangeotheriid "symmetrodont"

M. asiaticus[86]

Beipiao

Lujiatun Bed

A zhangeotheriid "symmetrodont"

Meemannodon

M. lujiatunensis

Beipiao

Lujiatun Bed

A gobiconodont
Origolestes O. lii A zhangeotheriid "symmetrodont"

Repenomamus

R. giganticus

Beipiao

Lujiatun Bed

A gobiconodont

R. robustus

Beipiao

Lujiatun Bed

A large gobiconodont, which ate small dinosaurs

Sinobaatar

S. lingyuanensis

Lingyuan

Jianshangou and Dawangzhangzi Beds

A multituberculate

Sinodelphys

S. szalayi

Lingyuan

Dawangzhangzi Bed

A therian, initially interpreted as a metatherian and later reinterpreted as a eutherian

Yanoconodon[87]

Y. allini[87]

Fengning[87]

A eutricondodont

Zhangheotherium

Z. quinquecuspidens

Chaoyang

Jianshangou Bed

A zhangeotheriid "symmetrodont"

Amphibians

[edit]
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

Callobatrachus

C. sanyanensis

Sihetun

Jianshangou Bed

Junior synonym of Liaobatrachus

Liaobatrachus

Laccotriton

L. subsolanus

Paozhanggou

Fengshan Bed

A urodelan

Liaobatrachus

L. grabaui

Sihetun

Jianshangou Bed

A spadefoot toad

L. zhaoi

Beipiao & Heitizigou

Lujiatun Bed & Jianshangou Bed

A spadefoot toad

Mesophryne

M. beipiaoensis

Beipiao & Heitizigou

Lujiatun Bed & Jianshangou Bed

Junior synonym of Liaobatrachus

Sinerpeton

S. fengshanensis

Paozhanggou

Fengshan Bed

A urodelan

Fish

[edit]
Genus Species County Member Abundance Notes Images

Hybodontoidea indet.

Family, genus and species indeterminate

An elasmobranch known from a single dorsal fin spine[88]
Lycoptera
Mesomyzon
Protopsephurus
Sinamia

Lycoptera

L. davidi

Jianshangou & Dawangzhangzi Beds

Also present in the Jiufotang Formation & Huajiying Formation

An osteoglossiform

L. muroii

Jingangshan Bed

L. sinensis

Jianshangou Bed

Mesomyzon

M. mengae

A lamprey

Peipiaosteus

P. pani

Jianshangou & Dawangzhangzi Bed

Also present in the Jiufotang Formation

A peipiaosteid fish related to sturgeons

Protopsephurus

P. liui

Dawangzhangzi Bed

Also present in the Jiufotang & Huajiying Formations

A paddlefish

Sinamia

S. liaoningensis

Jianshangou Bed

A bowfin

Yanosteus

Y. longidorsalis

Also present in the Huajiying Formation

A peipiaosteid

Invertebrate fauna of the Yixian Formation

[edit]

The invertebrate species listed below follow a survey by Chen and colleagues published in 2005, unless otherwise noted.[77]

Arachnids

[edit]

The arachnid species listed below follow a catalogue compiled by Dunlop, Penney and Jekel in 2010, unless otherwise noted.[89]

Genus Species County Member Abundance Notes Images

?Araneus

A.? aethus

Orb-weaver spiders

A.? beipiaoensis

A.? liaoxiensis

A.? reheensis

Cretaraneus

C. liaoningensis

A nephiline spider

?Gnaphosa

G.? liaoningensis

A ground spider

?Theridion

T.? atalus

A tangle web spider

Beetles (Coleoptera)

[edit]
Genus Species County Member Abundance Notes Images

Anoixis

A. complanus

A desmatin click beetle[90]

Apoclion

A. antennatus

Desmatin click beetles[90]

A. clavatus

A. dolini

Coptoclava

C. longipoda

A beetle[91]

Desmatinus

D. cognatus

A desmatin click beetle[90]

Fortiseode

F. pervalimand

An eodromeine trachypachid beetle[92]

Glaresis

G. orthochilus

A laresid scarab beetle[93]

Hydrophilopsia

H. gracilis

Hydrophiloid water scavenger beetles[94]

H. hydraenoides

H. shatrovskiyi

Lasiosyne L. integera Huangbanjigou Village, Liaoning Province, northeastern China.[95] A lasiosynid beetle.[95]
L. parva Huangbanjigou Village, Liaoning Province, northeastern China.[95] A lasiosynid beetle.[95]

Liaoximordella

L. hongi

A mordellid beetle[96]

Mirimordella

M. gracilicruralis

A mordellid beetle[97]

Oxyporus

O. yixianus

An oxyporine rove beetle[98]

Paradesmatus

P. dilatatus

A desmati click beetle[90]

Parageotrupes

P. incanus

A Geotrupine scarab beetle[99]

Sinocupes

S. validus

A beetle[91]

Sinosperchopsis

S. silinae

A hydrophiloid water scavenger beetle[94]

Sinoxytelus

S. breviventer

Oxyteline rove beetles[100]

S. euglypheus

S. longisetosus

Crustaceans

[edit]
Genus Species County Member Abundance Notes Images
Chenops[101] C. oblongus Notostracans
C. yixianensis
Cricoidoscelosus C. aethus Junior synonym of Palaeocambarus[102]
Cypridea C. (Cypridea) liaoningensis Jianshangou Bed Ostracods
C. (C.) sihetunensis Jianshangou Bed
C. (C.) veridica arguta Jingangshan Bed
C. (C.) sp. Jianshangou Bed
C. (Ulwellia) beipiaoensis Beipiao Jianshangou Bed
Damonella D. formosa Jianshangou Bed An ostracod
Darwinula D. leguminella Jianshangou Bed An ostracod
Diestheria D. jeholensis Jianshangou Bed Clam shrimps
D. yixianensis Dawangzhangzi Bed
D.? sp. Jianshangou Bed
Eosestheria E. jingangshanensis Jingangshan Bed Clam shrimps
E. lingyuanensis Dawangzhangzi Bed
E. aff. middendorfii Jingangshan Bed
E. ovata Jianshangou Bed
E. sihetunensis Jianshangou Bed
Eosestheriopsis E. gujialingensis Jianshangou Bed A clam shrimp
Jeholops[101] J. hongi A notostracan
Liaoningogriphus L. quadripartitus Jianshangou and Dawangzhangzi beds A shrimp
Lycopterocypris L. infantilis Jianshangou Bed An ostracod
Mantelliana M. sp. Jianshangou Bed An ostracod
"Mongolarachne" "M." chaoyangensis Dawangzhangzi Beds Junior synonym of Palaeocambarus[103]
Mongolianella M. cf. palmosa Jianshangou Bed An ostracod
Palaeocambarus P. licenti A crayfish[104]
Rhinocyrpis R. jurassica Jianshangou Bed An ostracod
Timiriasevia T. jianshangouensis Jianshangou Bed An ostracod
?Yanjiestheria Y.? beipiaoensis Beipiao Jianshangou Bed A clam shrimp
Yanshania Y. dabeigouensis Jianshangou Bed An ostracod


Dragonflies (Odonata)

[edit]
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.

The dragonfly species listed below follow a review compiled by Zhang, Ren and Panf in 2008, unless otherwise noted.[105]

Genus Species County Member Abundance Notes Images
Abrohemeroscopus A. mengi A dragonfly
Sinaeschnidia sp.
Aeschnidium A. heishankowense Reclassified as Sinaeschnidia heishankowense[106]
A. sp.
Archaeogomphus A. labius Nymphs[106] Reclassified as Yixiangomphus labius,[107] which is a junior synonym of Sinaeschnidia cancellosa[106]
Bellabrunetia B. catherinae A dragonfly
Chrysogomphus C. beipiaoensis A dragonfly
Dissurus D. liaoyuanensis[107] Poorly preserved nymph[106] Reclassified as Sinaeschnidia cf. cancellosa[106]
Dracontaeschnidium D. orientale A dragonfly
Hebeiaeschnidia H. fengningensis A dragonfly
Huaxiagomphus H. taushanensis A dragonfly[107]
Liogomphus L. yixianensis A dragonfly
Mesocordulia

M. boreala

A corduliid dragonfly[96]
Palaeogomphus P. labius A dragonfly, reclassified as of Yixiangomphus labius,[107] which is a junior synonym of Sinaeschnidia cancellosa[106]
Parapetala P. liaoningensis A dragonfly
P. yixianensis Jianshangou Unit[108]
Protoliupanshania P. wangi Jianshangou Unit[108] A liupanshaniid dragonfly
Pseudocymatophlebia P. bisecta Ningcheng County[109] A well-preserved specimen.[109] An aktassiid dragonfly
Pseudoliupanshania P. magnanicellula Jianshangou Unit[108]
Pseudosamarura P. largina[91] One specimen, nymph[106] Reclassified as Sinaeschnidia cf. cancellosa[106]
Rudiaeschna R. limnobia An aeshnid dragonfly[96]
Sinaeschnidia S. heishankowense Jianshangou & Dawangzhangzi Beds Adults and nymphs[106]
S. cancellosa[106] Junior synonym of S. heishankowense[110]
S. sp. Indeterminate nymph specimens[106] Some nymph specimens have been given their own genera and species though probably belong so S. heishankowense but too poorly preserved to be certain[106]
Sinogomphus S. taoshanensis A dragonfly
Sinojagoria S. imperfecta A dragonfly
Sopholibellula S. amoena Dragonflies
S. eleganti
Stylaeschnidium S. rarum A dragonfly
Telmaeshna T. paradoxica Beipiao A dragonfly
Turanophlebia T. liaoningensis Beipiao[111] Jianshangou Unit[111] A tarsophlebiid dragonfly.
Yixiangomphus Y. labius Nymphs[106] Junior synonym of Sinaeschnidia cancellosa[106]

Flies (Diptera)

[edit]
Genus Species County Member Abundance Notes Images
Alleremonomus A. xingi Jianshangou Bed A brachyceran fly
Atalosciophila A. yanensis A mycetophilid gnat[112]
Chironomaptera C. gregaria A phantom midge
Gigantoberis G. liaoningensis A soldier fly[113]
Lichnoplecia L. kovalevi A protopleciid fly[114]
Palaepangonius P. eupterus A horse-fly[96]
Protapiocera P. megista An asiloid fly[96]
Protempis P. minuta A brachyceran fly[96]
Protonemestrius P. jurassicus A tangle-veined fly[96]

Wasps (Hymenoptera)

[edit]
Genus Species County Member Abundance Notes Images
Abropelecinus A. annulatus Beipiao An iscopinine pelecinid[115]
Angaridyela A. robusta Jianshangou Bed A sawfly
Azygopelecinus A. clavatus Beipiao

An iscopinine pelecinid[115]

Chengdeserphus C. petidatus A proctotrupid wasp[96]
Cretevania C. exquisita Beipiao Hatchet wasps[116]
C. pristine Jianshangou bed
C. vesca
Hoplitolyda H. duolunica The largest known sawfly ever[117]
Liaotoma L. linearis A xyelotomid woodwasp[96]
Manlaya M. flexuosus An aulacid woodwasp[96]
Procretevania P. exquisita

Reclassified as Cretevania spp.[118]

P. pristine
P. vesca
Rudisiricius R. belli

Praesiricid sawflies[119]

R. celsus
R. crassinodus
Scolichneumon S. rectivenius An ichneumonoid wasp[96]
Sinopelecinus S. viriosus A pelecinid[115]
Tanychora T. exquisita Jianshangou Bed An ichneumon wasp

Other insects

[edit]
Genus Species County Member Abundance Notes Images
Abrigramma A. calophleba A kalligrammatid lacewing[120]
Alloraphidia A. bliquivenatica[91] Snakeflies
A. longistigmasa Jianshangou Bed
Archeraphidia A. shangyuanensis A snakefly[91]
Baissopterus B. liaoningensis A snakefly[91]
Caenoephemera C. shangyuanensis Jianshangou Bed A mayfly[91]
Caloraphidia C. glossophylla A snakefly[96]
Choromyrmeleon C. othneius An antlion[96]
Circulaboilus C. aureus A grig[121]
Cretophasmomima C. melanogramma A stick insect[122]
Cretophasmomima, a stick insect
Cretophasmomima, a stick insect
?Ephemeropsis ?E. trisetalis Jianshangou and Jingangshan beds A mayfly. Chinese specimens probably represent species of Epicharmeropsis[123]
Epicharmeropsis E. hexavenulosus[123] Pingquan Mayflies
E. quadrivenulosus[123] Beipiao Jianshangou Bed
Hagiphasma H. paradoxa Jianshangou Bed A stick insect
Ilerdocossus I. beipiaoensis Palaeontinid hemipterans[124][125]
I. exiguus
I. exquisitus
I. fengningensis
I. hui
I. pingquanensis
Ithigramma I. multinervia Kalligrammatid lacewings[120]
Kalligramma K. liaoningensis Jianshangou Bed
Karatavoblatta K. formosa A cockroach[91]
Liaoaphidia L. fornicata A snakefly[91]
Liaonemobius L. tanae A sword-tail cricket[96]
Longicellochrysa L. yixiana A mesochrysopid lacewing[126]
Mesascalaphus M. yangi An owlfly[96]
Mesolygaeus M. laiyangensis A water bug[91]
Metabittacus M. colosseus Jianshangou Bed A scorpionfly[127]
Miracossus M. gongi Palaeontinid hemipterans[124]
M. ingentius
Nipponoblatta N. acerba Jianshangou Bed A cockroach
Oloberotha O. sinica A beaded lacewing[96]
Oregramma O. aureolosa Kalligrammatid lacewings[120]
O. illecebrosa
Pseudopulex P. magnus Stem-fleas[128][129]
P. tanlan
Renphasma R. sinica A stick insect[130]
Saurophthirus S. exquisitus Stem-fleas[131]
S. laevigatus
Sinochresmoda S. magnicornia A chresmodid water-skiing insect[132]
Sinosharaperla S. zhaoi Beipiao A stonefly[133]
Sinoviparosiphum S. lini An aphid[96]
Sophogramma S. lii Jianshangou Bed Kalligrammatid lacewings[134][120]
S. pingquanica
Tetraphelerus T. laetus A moth[91]
Tyrannopsylla T. beipiaoensis A stem-flea[135]
Varicapitatus V. sinuolatus "Huangbanjigou, Chaomidian Village, Beipiao City."[136] A hemipteran
Unnamed Vitimotauliidae Unnamed A caddisfly pupa, likely belonging to the Vitimotauliidae[137]

Molluscs

[edit]
Genus Species County Member Abundance Notes Images
Amplovalvata A. sp. Jianshangou Bed A gastropod
Arguniella A. lingyuanensis Jianshangou Bed A bivalve, also present in the Jiufotang and Fuxin Formations
A. yanshanensis Jianshangou Bed A bivalve, also present in the Jiufotang Formation
Bithynia B. haizhouensis Jingangshan Bed A gastropod
Corbicula C. anderssoni Jianshangou Bed Reclassified as Sphaerium anderssoni
C. jeholense Reclassified as Sphaerium jeholense
Gyraulus G. sp. Jianshangou Bed A ramshorn snail
Probaicalia P. vitimensis Jianshangou Bed A gastropod
Ptychastylus P. harpaeformis Jianshangou Bed Gastropods
P. cf. phillipi Jianshangou Bed
Sphaerium S. anderssoni Jianshangou Bed A fingernail clam, also present in the Jiufotang Formation
S. jeholense Junior synonym of S. anderssoni
S. selenginense Junior synonym of S. anderssoni
?Viviparus V.? cf. matsumotoi Jingangshan Bed A snail


Flora of the Yixian Formation

[edit]

Conifers

[edit]

The coniferous trees and shrubs listed below were compiled in a survey by Wang and colleagues in 2006, unless otherwise noted.[138]

Genus Species County Member Abundance Notes Images
Brachyphyllum A cypress-like conifer
Cephalotaxopsis A yew
Cunninghamia A cypress
Elatides A cypress
Elatocladus E. leptophyllus Conifers of uncertain classification
E. manchurica[16]
Liaoningocladus L. boii Jingangshan Bed A conifer of uncertain classification
Pagiophyllum A cypress-like conifer
Pityocladus A pine
Pityolepis P. sp.[16] A pine
Pityophyllum P. cf. lindstroemi[16] A pine
Pityostrobus A pine
Podozamites P. lanceolatus[16] A conifer of uncertain classification
Schizolepis S. jeholensis[16]
?Sequoia S.? jeholensis[16]
Taxus A yew
Torreya A yew

Yanliaoia

Y. sinensis[16]

Ferns

[edit]
Genus Species County Member Abundance Notes Images
Cladophlebis C. spp.[16]
Coniopteris C. sp.[16]
Onychiopsis O. elongata[16] Junior synonym of O. psilotoides
O. psilotoides[16]
?Ruffordia R.? sp.[16]

Flowering plants

[edit]
Genus Species County Member Abundance Notes Images
Archaefructus A. eoflora Jianshangou Bed A primitive, aquatic angiosperm
Archaefructus
A. liaoningensis Jianshangou Bed
A. sinensisis Dawangzhangzi Bed
Callianthus C. dilae[139] Liaoning A plant with true flowers
?Davallia D.? niehhutzuensis[16]
Erenia E. stenoptera[139] Liaoning Actually a specimen of Callianthus
Leefructus L. mirus[140] A eudicot
Liaoxia L. chenii[16]
?Potamogeton P.? jeholensis[16]
Ranunculus R. jeholensis[16] Junior synonym of Potamogeton jeholensis
Sinocarpus S. decussatus A primitive eudicot

Ginkgos

[edit]
Genus Species County Member Abundance Notes Images
Baiera B. cf. gracilis[16]
B. orientalis[16]
Czekanowskia C. rigida[16]
Ginkgo G. apodes[141]
Ginkgoites G. sibiricus[16]
Leptostrobus L. sp.[16]
?Nageiopsis N.? sp.[16]
Phoenicopsis P. cf. augustifolia[16]
Solenites S. sp.[16]
Sphenobaiera S. longifola[16]

Other plants

[edit]
Genus Species County Member Abundance Notes Images
Archaeamphora A. longicervia Originally described as pitcher plant, later study confirmed that it is likely to be an insect galls on Liaoningocladus.
Archaeamphora
Chaoyangia C. liangii[142] A gnetale gymnosperm
Chaoyangicarpus C. liangii[16] Junior synonym of Chaoyangia liangii
Ephedra E. sp. A gymnosperm
Equisetum E. sp. A horsetail
Eragrosites E. changii[142] A grass-like gnetale gymnosperm
Siphonospermum S. simplex[143] Beipiao Jianshangou Bed A gnetale gymnosperm
Tyrmia T. sp.[16] A bennettite

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