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 5 million years during the early Cretaceous period (early Aptian stage). It is known for its fossils, listed below.

The Yixian Formation is divided into the following subunits, from most recent to oldest: (ages when available from Chang et al., 2009[1])

  • Jingangshan Bed (youngest, 122.1 Ma)
  • Dawangzhangzi Bed
  • Lujiatun Bed (123.2 Ma)
  • Jianshangou Bed (124.1 Ma)
  • Basalt base (oldest, 129.7 Ma)

Amphibians[edit]

Genus Species County Member Abundance Notes

Callobatrachus

C. sanyanensis

Lujiatun Bed

A disc-tongued frog

Liaobatrachus

L. grabaui

Jianshangou Bed

A spadefoot toad

Liaoxitriton

L. zhongjiani

A salamander related to the earlier L. daohugouensis of the Daohugou Beds.

Mesophryne

M. beipiaoensis

Lujiatun Bed

A frog with no known living relatives.

Fish[edit]

Genus Species County Member Abundance Notes Images

Hybodontoidea indet.

Family, genus and species indeterminate

A shark known from a single dorsal fin spine[2]

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. zdanskyi

Jianshangou Bed

A Bowfin

Yanosteus

Y. longidorsalis

Also present in the Huajiying Formation

A peipiaosteid

Mammals[edit]

The mammals listed below were compiled in a survey by Meng and colleagues in 2006, unless otherwise noted.[3]

Genus Species County Member Abundance Notes Images

Acristatherium

A. yanensis[4]

Beipiao

Lujiatun Bed

A eutherian

Akidolestes

A. cifellii

Lingyuan

Dawangzhangzi Bed

A symmetrodont

Chaoyangodens[5]

C. lii

Lingyuan

Dawangzhangzi Bed

A eutriconodont

Eomaia

E. scansoria

Lingyuan

Dawangzhangzi Bed

A stem therian

Gobiconodon

G. zofiae

Beipiao

Lujiatun Bed

A gobiconodont

G. luoianus [6]

A gobiconodont

Jeholodens

J. jenkinsi

Chaoyang

Jianshangou Bed

A eutriconodont

Juchilestes[7]

J. liaoningensis

Beipiao

Lujiatun Bed

An amphidontid

Maotherium

M. sinensis

Chaoyang

Jianshangou Bed

A symmetrodont

M. asiaticus[8]

Beipiao

Lujiatun Bed

A symmetrodont

Meemannodon

M. lujiatunensis

Beipiao

Lujiatun Bed

A gobiconodont

Repenomamus

R. giganticus

Beipiao

Lujiatun Bed

A gobiconodont

R. robustus

Beipiao

Lujiatun Bed

A gobiconodont, which ate small dinosaurs

Sinobaatar

S. lingyuanensis

Lingyuan

Jianshangou and Dawangzhangzi Beds

A multituberculate

Sinodelphys

S. szalayi

Lingyuan

Dawangzhangzi Bed

A metatherian

Yanoconodon[9]

Y. allini[9]

Fengning[9]

A eutricondodont

Zhangheotherium

Z. quinquecuspidens

Chaoyang

Jianshangou Bed

A symmetrodont

Pterosaurs[edit]

The pterosaurs listed below were compiled in a survey by Wang and Zhou in 2006, unless otherwise noted.[10]

Genus Species County Member Abundance Notes Images

Beipiaopterus

B. chenianus

Jianshangou Bed

A ctenochasmatid

Boreopterus

B. cuiae

Jianshangou Bed

A boreopterid

Cathayopterus

C. grabaui

Dawangzhangzi Bed

A ctenochasmatid

Elanodactylus[11]

E. prolatus

Jianshangou Bed[12]

A ctenochasmatid

Eosipterus

E. yangi

Jianshangou Bed

A ctenochasmatid

Feilongus

F. youngi

Jianshangou Bed

A boreopterid

Gegepterus[13]

G. changae[12]

Jianshangou Bed[12]

A ctenochasmatid

Gladocephaloideus[14]

G. jingangshanensis

Jingangshan Bed

A gallodactylid

Haopterus

H. gracilis

Jianshangou Bed

An ornithocheirid

Moganopterus

M. zhuiana

Lujiatun bed?

An ornithocheiroid

Ningchengopterus

N. liuae

Jianshangou or Dawangzhangzi bed

A pterodactyloid[15]

Pterofiltrus

P. qiui

Jianshangou Bed

A ctenochasmatid[12]

Zhenyuanopterus

Z. longirostris

Jianshangou Bed

A boreopterid[16]

Misc reptiles[edit]

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 Abundance Notes Images

Hyphalosaurus

H. lingyuanensis

Dawangzhangzi Bed

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 Abundance Notes Images

Dalinghosaurus

D. longidigitus

Jianshangou Bed

A lizard related to shinisaurs

Liushusaurus

L. acanthocaudata[17]

Jianshangou or Dawangzhangzi bed

A lizard related to geckos and skinks

Xianglong

X. zhaoi

Dawangzhangzi Bed

A gliding lizard

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 Abundance Notes Images

Manchurochelys

M. liaoxiensis[18]

Jianshangou Bed

Reclassified as Ordosemys liaoxiensis[19]

M. manchoukuoensis[20]

Jingangshan Bed

Ordosemys

O. liaoxiensis[19]

Jianshangou Bed

Ornithischians[edit]

The dinosaurs listed below were compiled in a survey by Xu and Norell in 2006, unless otherwise noted.[21]

Genus Species County Member Material Notes Images

Bolong

B. yixianensis

An ornithopod

Jeholosaurus

J. shangyuanensis

Beipiao

Lujiatun Bed

"Skull with associated vertebrae and hindlimb, skull with cervical vertebrae."[22]

An ornithopod

Jinzhousaurus

J. yangi

Yixian

Dawangzhangzi Bed

"Nearly complete skull and skeleton."[23]

An ornithopod

Liaoceratops

L. yanzigouensis

Beipiao

Lujiatun Bed

"[Two] nearly complete skulls, juvenile to adult."[24]

A ceratopsian

Liaoningosaurus

L. paradoxus

Yixian

Dawangzhangzi Bed

"Partial skull [and] skeleton."[25]

A small ankylosaur

Psittacosaurus

P. lujiatunensis

Beipiao

Lujiatun & Jianshangou Beds

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

A ceratopsian

Saurischians[edit]

The dinosaurs listed below were compiled in a survey by Xu and Norell in 2006, unless otherwise noted.[21]

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

Archaeorhynchus

A. spathula

Dawangzhangzi Bed

An early, advanced short-tailed bird (ornithuran)[26]

Bohaiornis[27]

B. guoi

Jianshangou Bed?

Well-preserved skeleton

An enantiornithine

Changchengornis[28]

C. hengdaoziensis[28]

Jianshangou Bed

"Nearly complete skeleton."[28]

A confuciusornithid

Confuciusornis

C. chuonzhous

Junior synonym of Confuciusornis sanctus[20]

C. dui[29]

Jianshangou Bed

"[Two] skeletons."[29]

C. feducciai[30]

Jianshangou Bed

C. sanctus[29]

Jianshangou and Dawangzhangzi Beds

"Hundreds of complete skeletons."[29]

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

C. suniae

Junior synonym of Confuciusornis sanctus[20]

Eoenantiornis

E. buhleri

Jianshangou Bed

"Nearly complete skeleton."[31]

An entantiornithine

Dalingheornis

D. liweii

Dawangzhangzi Bed

An entantiornithine[32]

Jinzhouornis

J. yixianensis

Jianshangou Bed

A pygostylian, possibly a junior synonym of Confuciusornis[33]

J. zhangjiyingia

Jianshangou Bed

Possible synonym of J. yixianensis

Jixiangornis

J. orientalis

Jianshangou Bed

A basal avialan, also present in the Jiufotang Formation[34]

Hongshanornis

H. longicresta

4 specimens[35]

Jianshangou and Dawangzhangzi Beds[35]

An ornithuromorph[36]

Jibeinia

J. luahera

Previously regarded as Yixian, horizon reassigned to the Huajiying Formation by Jin et al., 2008.[37]

Liaoningornis

L. longidigitris

Jianshangou Bed

"Partial skeleton."[38]

An enantiornithine.[39]

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.[40]

Lingyuanornis

L. parvus

Dawangzhangzi Bed

Junior objective synonym of Liaoxiornis delicatus, a nomen dubium.[40]

Longicrusavis

L. houi

Dawangzhangzi Bed

An ornithuromorph[36]

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.[37]

Shanweiniao

S. cooperorum

Dawangzhangzi Bed

An enantiornithine, the only one known with a fan tail as in modern birds.[41]

Vescornis

V. hebeiensis

Previously regarded as Yixian, horizon reassigned to the Huajiying Formation by Jin et al., 2008.[37]

Zhongornis

Z. haoae

Dawangzhangzi Bed

A primitive avialan with a short but non-pygostylian tail[42]

Deinonychosaurs[edit]

Genus Species County Member Material Notes Images

Changyuraptor[43]

C. yangi

Jianchang

A complete articulated specimen with extensive feathering.[43]

A dromaeosaurid

Graciliraptor

G. lujiatunensis

Beipiao

Lujiatun Bed

Rare

A dromaeosaurid

Mei

M. long

Beipiao

Lujiatun Bed

A small troodontid fossilized in a sleeping position

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.[44] Feathered integument is also preserved.[44][45]

A dromaeosaurid, the first ever found with feather impressions.

Sinovenator

S. changii

Beipiao

Lujiatun Bed

Three specimens with partial skulls and postcrania.[46][47] Hundreds of additional specimens known but undescribed.

A troodontid

Sinusonasus

S. magnodens

Beipiao

Lujiatun Bed

A troodontid

Tianyuraptor

T. ostromi

Lingyuan

Dawangzhangzi Bed

A dromaeosaurid

Zhenyuanlong[48]

Z. suni

Jianchang

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

A dromaeosaurid

Zhongjianosaurus[49]

Z. yangi

Lingyuan

A partial skeleton including an arm, both legs, vertebrae, ribs and most of the tail.[50]

A microraptorine dromaeosaurid with very long legs.[51]

Undescribed deinonychosaur/possible troodontid

Dawangzhangzi Bed

One specimen with feather imprints

A possible troodontid.[52]

Undescribed troodontid

Specimen CAGS-IG01-004

Lujiatun Bed

Skull and partial skeleton

A troodontid similar to Mei.[53]

Unnamed dromaeosaurid

Specimen NGMC 91

Lingyuan

Dawangzhangzi Bed

Complete specimen with feather imprints

A juvenile dromaeosaurid that may be a specimen of Sinornithosaurus millenii.[54][55] Informally nicknamed "Dave".

Other theropods[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

Beipiaosaurus

B. inexpectus

Beipiao

Jianshangou Bed

"Incomplete skeleton with integument."[56]

A therizinosaur

Caudipteryx

C. dongi

Beipiao

Jianshangou Bed

"[One] partial postcranium with feather imprints."[57]

Oviraptorosaurs

C. zoui

Beipiao

Jianshangou Bed

Common, only over a small region

C. sp.[58]

Dilong

D. paradoxus

Beipiao

Lujiatun & Jianshangou Bed

Rare

A tyrannosauroid

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.[59]

Hexing

H. qingyi

A primitive ornithomimosaur[60]

Huaxiagnathus

H. orientalis

Beipiao

Jianshangou Bed

A compsognathid

Incisivosaurus

I. gauthieri

Beipiao

Lujiatun Bed

"[One] skull [and a] partial cervical."[57]

An oviraptorosaur

Ningyuansaurus

N. wangi

Complete skeleton with feather impressions.[61]

An oviraptorosaur

Protarchaeopteryx

P. robusta

Beipiao

Jianshangou Bed

"Nearly complete skull and skeleton, at least [two] individuals."[62]

A possible relative of Incisivosaurus

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.[63]

Shenzhousaurus

S. orientalis

Beipiao

Lujiatun Bed

"Articulated skull and incomplete skeleton."[64]

A primitive ornithomimosaur

Similicaudipteryx

S. yixianensis

Two specimens

An oviraptorosaur, also present in the Jiufotang Formation[65]

Sinocalliopteryx

S. gigas

Jianshangou Bed

A large compsognathid

Sinosauropteryx

S. prima

Beipiao & Lingyuan

Jianshangou & Dawangzhangzi Beds

Four nearly complete specimens[66][67]

The first non-avian dinosaur found with feather impressions, a compsognathid

S.? sp.

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.[68]

Yixianosaurus

Y. longimanus

Yixian

Dawangzhangzi Bed

A basal maniraptoran

Yutyrannus

Y. huali

Unknown

Three nearly complete skeletons of various ages

A large tyrannosauroid[69]

Unnamed theropod

Not yet described

Yixian

Lujiatun Bed

Partial post-cranial skeleton[70]

A possible carnosaur[71]

Sauropods[edit]

Genus Species County Member Material Notes Images

Dongbeititan

D. dongi

Jianshangou Bed

Partial postcrania[72]

A basal titanosauriform

cf. Euhelopus

cf. E. sp.

Lujiatun Bed

Isolated teeth[73]

A basal titanosauriform

Sauropoda spp.

Lujiatun & Jianshangou Beds

Isolated vertebrae and teeth[21]

One or more species of sauropods, possibly titanosaurs, have been recovered from the Yixian Formation, though most remain undescribed[73]

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.[18]

Arachnids[edit]

The arachnid species listed below follow a catalogue compiled by Dunlop, Penney and Jekel in 2010, unless otherwise noted.[74]

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[75]

Apoclion

A. antennatus

Desmatin click beetles[75]

A. clavatus

A. dolini

Coptoclava

C. longipoda

A beetle[76]

Desmatinus

D. cognatus

A desmatin click beetle[75]

Fortiseode

F. pervalimand

An eodromeine trachypachid beetle[77]

Glaresis

G. orthochilus

A laresid scarab beetle[78]

Hydrophilopsia

H. gracilis

Hydrophiloid water scavenger beetles[79]

H. hydraenoides

H. shatrovskiyi

Liaoximordella

L. hongi

A mordellid beetle[80]

Mirimordella

M. gracilicruralis

A mordellid beetle[81]

Oxyporus

O. yixianus

An oxyporine rove beetle[82]

Paradesmatus

P. dilatatus

A desmati click beetle[75]

Parageotrupes

P. incanus

A Geotrupine scarab beetle[83]

Sinocupes

S. validus

A beetle[76]

Sinosperchopsis

S. silinae

A hydrophiloid water scavenger beetle[79]

Sinoxytelus

S. breviventer

Oxyteline rove beetles[84]

S. euglypheus

S. longisetosus

Crustaceans[edit]

Genus Species County Member Abundance Notes Images

Chenops[85]

C. oblongus

A notostracans

Chenops yixianensis
Chenops oblongus (right) and Jeholops hongi (left)

C. yixianensis

Cricoidoscelosus

C. aethus

A crayfish[86]

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[85]

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

Mongolianella

M. cf. palmosa

Jianshangou Bed

An ostracod

Palaeocambarus

P. licenti

A crayfish[86]

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.[87]

Genus Species County Member Abundance Notes Images

Abrohemeroscopus

A. mengi

A dragonfly

Aeschnidium

A. heishankowense

Reclassified as Sinaeschnidia heishankowense[88]

A. sp.

Archaeogomphus

A. labius

Nymphs[88]

Reclassified as Yixiangomphus labius,[89] which is a junior synonym of Sinaeschnidia cancellosa[88]

Bellabrunetia

B. catherinae

A dragonfly

Chrysogomphus

C. beipiaoensis

A dragonfly

Dissurus

D. liaoyuanensis[89]

Poorly preserved nymph[88]

Reclassified as Sinaeschnidia cf. cancellosa[88]

Dracontaeschnidium

D. orientale

A dragonfly

Hebeiaeschnidia

H. fengningensis

A dragonfly

Huaxiagomphus

H. taushanensis

A dragonfly[89]

Liogomphus

L. yixianensis

A dragonfly

Mesocordulia

M. boreala

A corduliid dragonfly[80]

Palaeogomphus

P. labius

A dragonfly, reclassified as of Yixiangomphus labius,[89] which is a junior synonym of Sinaeschnidia cancellosa[88]

Parapetala

P. liaoningensis

A dragonfly

Pseudosamarura

P. largina[76]

One specimen, nymph[88]

Reclassified as Sinaeschnidia cf. cancellosa[88]

Rudiaeschna

R. limnobia

An aeshnid dragonfly[80]

Sinaeschnidia

S. heishankowense

Jianshangou & Dawangzhangzi Beds

Adults and nymphs[88]

S. cancellosa[88]

Junior synonym of S. heishankowense[90]

S. sp.

Indeterminate nymph specimens[88]

Some nymph specimens have been given their own genera and species though probably belong so S. heishankowense but too poorly preserved to be certain[88]

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

Yixiangomphus

Y. labius

Nymphs[88]

Junior synonym of Sinaeschnidia cancellosa[88]

Flies (Diptera)[edit]

Genus Species County Member Abundance Notes Images

Alleremonmus

A. xingi

Jianshangou Bed

A brachyceran fly

Atalosciophila

A. yanensis

A mycetophilid gnat[91]

Chironomaptera

C. gregaria

A phantom midge

Gigantoberis

G. liaoningensis

A soldier fly[92]

Lichnoplecia

L. kovalevi

A protopleciid fly[93]

Palaepangonius

P. eupterus

A horse-fly[80]

Protapiocera

P. megista

An asiloid fly[80]

Protempis

P. minuta

A brachyceran fly[80]

Protonemestrius

P. jurassicus

A tangle-veined fly[80]

Wasps (Hymenoptera)[edit]

Genus Species County Member Abundance Notes Images

Abropelecinus

A. annulatus

Beipiao

An iscopinine pelecinid[94]

Angaridyela

A. robusta

Jianshangou Bed

A sawfly

Azygopelecinus

A. clavatus

Beipiao

An iscopinine pelecinid[94]

Chengdeserphus

C. petidatus

A proctotrupid wasp[80]

Cretevania

C. exquisita

Beipiao

Hatchet wasps[95]

C. pristine

Jianshangou bed

C. vesca

Liaotoma

L. linearis

A xyelotomid woodwasp[80]

Manlaya

M. flexuosus

An aulacid woodwasp[80]

Procretevania

P. exquisita

Reclassified as Cretevania spp.[96]

P. pristine

P. vesca

Rudisiricius

R. belli

Praesiricid sawflies[97]

R. celsus

R. crassinodus

Scolichneumon

S. rectivenius

An ichneumonoid wasp[80]

Sinopelecinus

S. viriosus

A pelecinid[94]

Tanychora

T. exquisita

Jianshangou Bed

An ichneumon wasp

Other insects[edit]

Genus Species County Member Abundance Notes Images

Alloraphidia

A. bliquivenatica[76]

Snakeflies

Epicharmeropsis, a mayfly

A. longistigmasa

Jianshangou Bed

Archeraphidia

A. shangyuanensis

A snakefly[76]

Baissopterus

B. liaoningensis

A snakefly[76]

Caenoephemera

C. shangyuanensis

Jianshangou Bed

A mayfly[76]

Caloraphidia

C. glossophylla

A snakefly[80]

Choromyrmeleon

C. othneius

An antlion[80]

?Ephemeropsis

?E. trisetalis

Jianshangou and Jingangshan beds

A mayfly. Chinese specimens probably represent species of Epicharmeropsis[98]

Epicharmeropsis

E. hexavenulosus[98]

Pingquan

Mayflies

E. quadrivenulosus[98]

Beipiao

Jianshangou Bed

Hagiphasma

H. paradoxa

Jianshangou Bed

A stick insect

Ilerdocossus

I. beipiaoensis

Cicadas[99]

I. exiguus

I. fengningensis

I. hui

I. pingquanensis

Kalligramma

K. liaoningensis

Jianshangou Bed

A lacewing

Karatavoblatta

K. formosa

A cockroach[76]

Liaoaphidia

L. fornicata

A snakefly[76]

Liaonemobius

L. tanae

A sword-tail cricket[80]

Longicellochrysa

L. yixiana

A mesochrysopid lacewing[100]

Mesascalaphus

M. yangi

An owlfly[80]

Mesolygaeus

M. laiyangensis

A water bug[76]

Metabittacus

M. colosseus

Jianshangou Bed

Miracossus

M. ingentius

A cicada[99]

Nipponoblatta

N. acerba

Jianshangou Bed

A cockroach

Oloberotha

O. sinica

A beaded lacewing[80]

Sinochresmoda

S. magnicornia

A chresmodid water-skiing insect[101]

Sinosharaperla

S. zhaoi

Beipiao

A stonefly[102]

Sinoviparosiphum

S. lini

An aphid[80]

Sophogramma

S. lii

Jianshangou Bed

A lacewing[103]

Tetraphelerus

T. laetus

A moth[76]

Unnamed Vitimotauliidae

Unnamed

A caddisfly pupa, likely belonging to the Vitimotauliidae.[104]

Unnamed Siphonaptera

Unnamed

A giant, non-jumping flea[105]

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.[106]

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[20]

Liaoningocladus

L. boii

Jingangshan Bed

A conifer of uncertain classification

Pagiophyllum

A cypress-like conifer

Pityocladus

A pine

Pityolepis

P. sp.[20]

A pine

Pityophyllum

P. cf. lindstroemi[20]

A pine

Pityostrobus

A pine

Podozamites

P. lanceolatus[20]

A conifer of uncertain classification

Schizolepis

S. jeholensis[20]

?Sequoia

S.? jeholensis[20]

Taxus

A yew

Torreya

A yew

Yanliaoia

Y. sinensis[20]

Ferns[edit]

Genus Species County Member Abundance Notes Images

Cladophlebis

C. spp.[20]

Coniopteris

C. sp.[20]

Onychiopsis

O. elongata[20]

Junior synonym of O. psilotoides

O. psilotoides[20]

?Ruffordia

R.? sp.[20]

Flowering plants[edit]

Genus Species County Member Abundance Notes Images

Archaeamphora

A. longicervia

Jianshangou Bed

A pitcher plant, and the earliest known carnivorous plant

Archaefructus

A. eoflora

Jianshangou Bed

A primitive, aquatic angiosperm

A. liaoningensis

Jianshangou Bed

A. sinensisis

Dawangzhangzi Bed

Callianthus

C. dilae[107]

Liaoning

A plant with true flowers

?Davallia

D.? niehhutzuensis[20]

Erenia

E. stenoptera[107]

Liaoning

Actually a specimen of Callianthus

Leefructus

L. mirus[108]

A eudicot

Liaoxia

L. chenii[20]

?Potamogeton

P.? jeholensis[20]

Ranunculus

R. jeholensis[20]

Junior synonym of Potamogeton jeholensis

Sinocarpus

S. decussatus

A primitive eudicot

Ginkgos[edit]

Genus Species County Member Abundance Notes Images

Baiera

B. cf. gracilis[20]

B. orientalis[20]

Czekanowskia

C. rigida[20]

Ginkgo

G. apodes[109]

Ginkgoites

G. sibiricus[20]

Leptostrobus

L. sp.[20]

?Nageiopsis

N.? sp.[20]

Phoenicopsis

P. cf. augustifolia[20]

Solenites

S. sp.[20]

Sphenobaiera

S. longifola[20]

Other plants[edit]

Genus Species County Member Abundance Notes Images

Chaoyangia

C. liangii[110]

A gnetale gymnosperm

Chaoyangicarpus

C. liangii[20]

Junior synonym of Chaoyangia liangii

Ephedra

E. sp.

A gymnosperm

Equisetum

E. sp.

A horsetail

Eragrosites

E. changii[110]

A grass-like gnetale gymnosperm

Siphonospermum

S. simplex[111]

Beipiao

Jianshangou Bed

A gnetale gymnosperm

Tyrmia

T. sp.[20]

A cycad

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