Paleobiota of the Yixian Formation
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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
[edit]| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
D. dongi |
Beipiao | Jianshangou Bed |
Partial postcrania[2] |
A basal titanosauriform |
 | |
|
cf. Euhelopus |
cf. E. sp. |
Lujiatun Bed |
A basal titanosauriform | |||
|
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 |
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
[edit]| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
L. paradoxus |
Yixian |
Dawangzhangzi Bed |
"Partial skull [and] skeleton."[8] |
A small ankylosaur |  |
Ceratopsians
[edit]| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
L. yanzigouensis |
Beipiao |
Lujiatun Bed |
"[Two] nearly complete skulls, juvenile to adult."[9] |
  | ||
|
Beipiao |
Lujiatun & Jianshangou Beds |
Thousands of specimens in the Lujiatun Bed, somewhat less common in the Jianshangou Bed |
A ceratopsian |
Ornithopods
[edit]| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
B. yixianensis |
Jianshangou Bed |
A nearly complete skeleton. |
   | |||
| Changmiania[10] | C. liaoningensis | Lujiatun Bed | Two nearly complete skeletons | A basal ornithopod that may have dug burrows | ||
|
J. shangyuanensis |
Lujiatun Bed |
"Skull with associated vertebrae and hindlimb, skull with cervical vertebrae."[11] |
An ornithopod | |||
|
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
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
| Abitusavis[13] | A. lii | Ningcheng | "Nearly complete skeleton."[13] |         | ||
|
A. spathula |
Dawangzhangzi Bed |
An early, advanced short-tailed bird (ornithuran)[14] | ||||
|
C. hengdaoziensis[15] |
Jianshangou Bed |
"Nearly complete skeleton."[15] |
||||
|
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] | |||||
|
D. liweii |
Dawangzhangzi Bed |
|||||
|
E. buhleri |
Jianshangou Bed |
"Nearly complete skeleton."[20] |
||||
|
E. edentulata |
Dawangzhangzi Bed |
"nearly complete skeleton with skull"[21] |
An ornithuromorph | |||
|
I. huchzermeyeri |
An ornithuromorph | |||||
|
J. yixianensis |
Jianshangou Bed |
A pygostylian, a junior synonym of Confuciusornis sanctus[22] | ||||
|
J. zhangjiyingia |
Jianshangou Bed |
Junior synonym of C. sanctus | ||||
|
J. orientalis |
Jianshangou Bed |
A basal avialan, also present in the Jiufotang Formation[23] | ||||
|
H. longicresta |
Jianshangou and Dawangzhangzi Beds[24] |
4 specimens[24] |
An ornithuromorph[25] | |||
|
J. luahera |
Previously regarded as Yixian, horizon reassigned to the Huajiying Formation by Jin et al., 2008.[26] | |||||
|
L. longidigitris |
Lujiatun Bed |
"Partial skeleton."[27] |
An enantiornithine.[28] | |||
|
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] | ||||
|
L. parvus |
Dawangzhangzi Bed |
Junior objective synonym of Liaoxiornis delicatus, a nomen dubium.[29] | ||||
|
L. houi |
Dawangzhangzi Bed |
An ornithuromorph[25] | ||||
|
L. hani |
bed unknown |
An enantiornithine | ||||
|
P. fengningensis |
Previously regarded as Yixian, horizon reassigned to the Huajiying Formation by Jin et al., 2008.[26] | |||||
|
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] | |||
|
S. cooperorum |
Dawangzhangzi Bed |
An enantiornithine.[32] | ||||
|
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 | |||
|
Z. haoae |
Dawangzhangzi Bed |
A primitive avialan with a short but non-pygostylian tail[33] |
Compsognathids
[edit]Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
H. orientalis |
Beipiao |
Jianshangou Bed |
  | |||
|
S. gigas |
Beipiao |
Jianshangou Bed |
A large compsognathid | |||
|
S. prima |
Beipiao & Lingyuan |
Jianshangou & Dawangzhangzi Beds |
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
[edit]| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
C. yangi |
Jianchang |
A complete articulated specimen with extensive feathering.[37] |
      | |||
|
G. lujiatunensis |
Beipiao |
Lujiatun Bed |
An incomplete specimen. |
A dromaeosaurid | ||
|
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. | ||
|
T. ostromi |
Lingyuan |
Dawangzhangzi Bed |
A dromaeosaurid | |||
|
Z. suni |
Jianchang |
One nearly complete skeleton with preserved feathers, including long tail feathers and large wing feathers. |
A dromaeosaurid | |||
|
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
[edit]Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
H. qingyi |
Beipiao |
Lujiatun |
A primitive ornithomimosaur[44] |
 | ||
|
S. orientalis |
Beipiao |
Lujiatun Bed |
"Articulated skull and incomplete skeleton."[45] |
A primitive ornithomimosaur |
Oviraptorosaurs
[edit]Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
C. dongi |
Beipiao |
Jianshangou Bed |
"[One] partial postcranium with feather imprints."[46] |
 .png)  | ||
|
C. zoui |
Beipiao |
Jianshangou Bed |
Common, only over a small region | |||
|
C. sp.[47] |
||||||
|
I. gauthieri |
Beipiao |
Lujiatun Bed |
"[One] skull [and a] partial cervical."[46] |
An oviraptorosaur | ||
|
N. wangi |
Complete skeleton with feather impressions.[48] |
An oviraptorosaur | ||||
|
P. robusta |
Beipiao |
Jianshangou Bed |
"Nearly complete skull and skeleton, at least [two] individuals."[49] |
A possible relative of Incisivosaurus | ||
|
S. yixianensis |
Two specimens |
An oviraptorosaur, also present in the Jiufotang Formation[50] | ||||
|
X. ganqi |
Liaoning |
Dakangpu Bed |
One nearly complete specimen (IVPP V13390), minus the head |
Therizinosaurs
[edit]Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
B. inexpectus |
Jianshangou Bed |
"Incomplete skeleton with integument."[52] |
  | |||
| Jianchangosaurus | J. yixianensis | Jianchang | A nearly complete juvenile specimen with feather impressions.[53] | A therizinosaur |
Troodontids
[edit]| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
| Daliansaurus | D. liaoningensis | Beipiao | Lujiatun Bed | A very complete specimen.[54] | A troodontid |     .png)  |
|
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 | ||
|
M. long |
Beipiao |
Lujiatun Bed |
A small troodontid fossilized in a sleeping position | |||
|
S. changii |
Beipiao |
Lujiatun Bed |
Three specimens with partial skulls and postcrania.[56][57] Hundreds of additional specimens known but undescribed. |
A troodontid | ||
|
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
[edit]Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
D. paradoxus |
Beipiao |
Lujiatun & Jianshangou Bed |
Rare |
  | ||
|
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] | |||||
|
R. kriegsteini |
A tyrannosaurid, probably from higher Mongolian strata. | |||||
|
S. heilmanni |
Initially reported from the Dawangzhangzi Bed, probably from the Daohugou Beds instead.[61] | |||||
|
Y. huali |
Unknown |
Three nearly complete skeletons of various ages |
A large tyrannosauroid[62] | |||
|
Unnamed theropod |
Not yet described |
Lujiatun Bed |
Partial post-cranial skeleton[63] |
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 |
|---|---|---|---|---|---|---|
|
B. chenianus |
Jianshangou Bed |
A ctenochasmatid |
     | |||
|
B. cuiae |
Jianshangou Bed |
A boreopterid | ||||
|
C. grabaui |
Dawangzhangzi Bed |
A ctenochasmatid | ||||
|
E. prolatus |
Jianshangou Bed[67] |
A ctenochasmatid | ||||
| Eopteranodon | E. lii[68] | Beipiao | A tapejarid | |||
| E. yixianensis[69] | Beipiao | Jianshangou Bed | A tapejarid | |||
|
E. yangi |
Jianshangou Bed |
A ctenochasmatid | ||||
|
G. changae[67] |
Jianshangou Bed[67] |
A ctenochasmatid | ||||
|
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 | |||
|
M. zhuiana |
Lujiatun bed? |
A ctenochasmatid | ||||
|
N. liuae |
Jianshangou or Dawangzhangzi bed |
A pterodactyloid[72] | ||||
| Nurhachius | N. sp. | Jingangshan Member[73] | An incomplete specimen[73] | An istiodactylid | ||
|
P. qiui |
Jianshangou Bed |
A ctenochasmatid[67] | ||||
| Yixianopterus | Y. jingangshanensis | An istiodactyliform | ||||
|
Z. longirostris |
Jianshangou Bed |
A boreopterid.[74] May be adult form of Boreopterus.[75] |
Misc. reptiles
[edit]Choristoderans
[edit]Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
H. lingyuanensis |
Dawangzhangzi Bed |
  | ||||
|
H. baitaigouensis |
Jingangshan Bed |
Thousands of specimens |
||||
|
M. splendens |
Dawangzhangzi Bed |
|||||
|
S. lingyuanensis |
Dawangzhangzi Bed |
Junior synonym of Hyphalosaurus lingyuanensis |
Lizards
[edit]| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
D. longidigitus |
Jianshangou Bed |
A lizard related to shinisaurs |
  | |||
|
L. acanthocaudata[76] |
Jianshangou or Dawangzhangzi bed |
A lizard related to geckos and skinks | ||||
|
X. zhaoi |
Jianshangou Bed |
A gliding lizard related to chameleons and dragon lizards | ||||
|
Y. tenuis |
Dawangzhangzi & Jingangshan Beds |
A lizard related to geckos and skinks |
Turtles
[edit]Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
M. liaoxiensis[77] |
Jianshangou Bed |
Reclassified as Ordosemys liaoxiensis[78] |
 | |||
|
M. manchoukuoensis[16] |
Jingangshan Bed |
|||||
|
O. liaoxiensis[78] |
Jianshangou Bed |
Mammals
[edit]Color key
|
Notes Uncertain or tentative taxa are in small text; |
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 |
|---|---|---|---|---|---|---|
|
A. yanensis[80] |
Beipiao |
Lujiatun Bed |
A eutherian |
     | ||
|
A. cifellii |
Lingyuan |
Dawangzhangzi Bed |
A spalacotheriid "symmetrodont" | |||
| Ambolestes[81] | A. zhoui | Nincheng | Jianshangou Bed | A eutherian | ||
|
C. lii |
Lingyuan |
Dawangzhangzi Bed |
A eutriconodont | |||
|
E. scansoria |
Lingyuan |
Dawangzhangzi Bed |
A tribosphenidan, possibly a eutherian or stem-therian | |||
|
G. zofiae |
Beipiao |
Lujiatun Bed |
A gobiconodont | |||
|
G. luoianus[83] |
Lujiatun Bed |
A gobiconodont; junior synonym of G. zofiae | ||||
|
J. jenkinsi |
Chaoyang |
Jianshangou Bed |
A eutriconodont | |||
|
J. liaoningensis |
Beipiao |
Lujiatun Bed |
An amphidontid | |||
|
J. cheni |
A eutriconodont | |||||
|
M. sinensis |
Chaoyang |
Jianshangou Bed |
A zhangeotheriid "symmetrodont" | |||
|
M. asiaticus[86] |
Beipiao |
Lujiatun Bed |
A zhangeotheriid "symmetrodont" | |||
|
M. lujiatunensis |
Beipiao |
Lujiatun Bed |
A gobiconodont | |||
| Origolestes | O. lii | A zhangeotheriid "symmetrodont" | ||||
| Ningchengodon | N. foxi[87] | Ningcheng | Ningcheng Basin | A partial right dentary. | A symmetrodont. | |
|
R. giganticus |
Beipiao |
Lujiatun Bed |
A gobiconodont | |||
|
R. robustus |
Beipiao |
Lujiatun Bed |
A large gobiconodont, which ate small dinosaurs | |||
|
S. lingyuanensis |
Lingyuan |
Jianshangou and Dawangzhangzi Beds |
A multituberculate | |||
|
S. szalayi |
Lingyuan |
Dawangzhangzi Bed |
A therian, initially interpreted as a metatherian and later reinterpreted as a eutherian | |||
|
Y. allini[88] |
Fengning[88] |
A eutricondodont | ||||
|
Z. quinquecuspidens |
Chaoyang |
Jianshangou Bed |
A zhangeotheriid "symmetrodont" |
Amphibians
[edit]Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Genus | Species | County | Member | Material | Notes | Images |
|---|---|---|---|---|---|---|
|
C. sanyanensis |
Sihetun |
Jianshangou Bed |
Junior synonym of Liaobatrachus |
 | ||
|
L. subsolanus |
Paozhanggou |
Fengshan Bed |
A urodelan | |||
|
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 | ||
|
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[89] |
    | |||
|
L. davidi |
Jianshangou & Dawangzhangzi Beds |
Also present in the Jiufotang Formation & Huajiying Formation |
||||
|
L. muroii |
Jingangshan Bed |
|||||
|
L. sinensis |
Jianshangou Bed |
|||||
|
M. mengae |
A lamprey | |||||
|
P. pani |
Jianshangou & Dawangzhangzi Bed |
Also present in the Jiufotang Formation |
A peipiaosteid fish related to sturgeons | |||
|
P. liui |
Dawangzhangzi Bed |
Also present in the Jiufotang & Huajiying Formations |
||||
|
S. liaoningensis |
Jianshangou Bed |
A bowfin | ||||
|
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.[90]
| Genus | Species | County | Member | Abundance | Notes | Images |
|---|---|---|---|---|---|---|
|
A.? aethus |
||||||
|
A.? beipiaoensis |
||||||
|
A.? liaoxiensis |
||||||
|
A.? reheensis |
||||||
|
C. liaoningensis |
||||||
|
G.? liaoningensis |
||||||
|
T.? atalus |
Beetles (Coleoptera)
[edit]| Genus | Species | County | Member | Abundance | Notes | Images |
|---|---|---|---|---|---|---|
|
A. complanus |
||||||
|
A. antennatus |
Desmatin click beetles[91] | |||||
|
A. clavatus |
||||||
|
A. dolini |
||||||
|
C. longipoda |
A beetle[92] | |||||
|
D. cognatus |
A desmatin click beetle[91] | |||||
|
F. pervalimand |
An eodromeine trachypachid beetle[93] | |||||
|
G. orthochilus |
||||||
|
H. gracilis |
Hydrophiloid water scavenger beetles[95] | |||||
|
H. hydraenoides |
||||||
|
H. shatrovskiyi |
||||||
| Lasiosyne | L. integera | Huangbanjigou Village, Liaoning Province, northeastern China.[96] | A lasiosynid beetle.[96] | |||
| L. parva | Huangbanjigou Village, Liaoning Province, northeastern China.[96] | A lasiosynid beetle.[96] | ||||
|
L. hongi |
||||||
|
M. gracilicruralis |
A mordellid beetle[98] | |||||
|
O. yixianus |
||||||
|
P. dilatatus |
A desmati click beetle[91] | |||||
|
P. incanus |
||||||
|
S. validus |
A beetle[92] | |||||
|
S. silinae |
A hydrophiloid water scavenger beetle[95] | |||||
|
S. breviventer |
||||||
|
S. euglypheus |
||||||
|
S. longisetosus |
Crustaceans
[edit]| Genus | Species | County | Member | Abundance | Notes | Images |
|---|---|---|---|---|---|---|
| Chenops[102] | C. oblongus | Notostracans | 
| |||
| C. yixianensis | ||||||
| Cricoidoscelosus | C. aethus | Junior synonym of Palaeocambarus[103] | ||||
| 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[102] | 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[104] | |||
| Mongolianella | M. cf. palmosa | Jianshangou Bed | An ostracod | |||
| Palaeocambarus | P. licenti | A crayfish[105] |  | |||
| 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
|
Notes Uncertain or tentative taxa are in small text; |
The dragonfly species listed below follow a review compiled by Zhang, Ren and Panf in 2008, unless otherwise noted.[106]
| Genus | Species | County | Member | Abundance | Notes | Images |
|---|---|---|---|---|---|---|
| Abrohemeroscopus | A. mengi | A dragonfly |  | |||
| Aeschnidium | A. heishankowense | Reclassified as Sinaeschnidia heishankowense[107] | ||||
| A. sp. | ||||||
| Archaeogomphus | A. labius | Nymphs[107] | Reclassified as Yixiangomphus labius,[108] which is a junior synonym of Sinaeschnidia cancellosa[107] | |||
| Bellabrunetia | B. catherinae | A dragonfly | ||||
| Chrysogomphus | C. beipiaoensis | A dragonfly | ||||
| Dissurus | D. liaoyuanensis[108] | Poorly preserved nymph[107] | Reclassified as Sinaeschnidia cf. cancellosa[107] | |||
| Dracontaeschnidium | D. orientale | A dragonfly | ||||
| Hebeiaeschnidia | H. fengningensis | A dragonfly | ||||
| Huaxiagomphus | H. taushanensis | A dragonfly[108] | ||||
| Liogomphus | L. yixianensis | A dragonfly | ||||
| Mesocordulia |
M. boreala |
A corduliid dragonfly[97] | ||||
| Palaeogomphus | P. labius | A dragonfly, reclassified as of Yixiangomphus labius,[108] which is a junior synonym of Sinaeschnidia cancellosa[107] | ||||
| Parapetala | P. liaoningensis | A dragonfly | ||||
| P. yixianensis | Jianshangou Unit[109] | |||||
| Protoliupanshania | P. wangi | Jianshangou Unit[109] | A liupanshaniid dragonfly | |||
| Pseudocymatophlebia | P. bisecta | Ningcheng County[110] | A well-preserved specimen.[110] | An aktassiid dragonfly | ||
| Pseudoliupanshania | P. magnanicellula | Jianshangou Unit[109] | ||||
| Pseudosamarura | P. largina[92] | One specimen, nymph[107] | Reclassified as Sinaeschnidia cf. cancellosa[107] | |||
| Rudiaeschna | R. limnobia | An aeshnid dragonfly[97] | ||||
| Sinaeschnidia | S. heishankowense | Jianshangou & Dawangzhangzi Beds | Adults and nymphs[107] | |||
| S. cancellosa[107] | Junior synonym of S. heishankowense[111] | |||||
| S. sp. | Indeterminate nymph specimens[107] | Some nymph specimens have been given their own genera and species though probably belong so S. heishankowense but too poorly preserved to be certain[107] | ||||
| 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[112] | Jianshangou Unit[112] | A tarsophlebiid dragonfly. | ||
| Yixiangomphus | Y. labius | Nymphs[107] | Junior synonym of Sinaeschnidia cancellosa[107] |
Flies (Diptera)
[edit]| Genus | Species | County | Member | Abundance | Notes | Images |
|---|---|---|---|---|---|---|
| Alleremonomus | A. xingi | Jianshangou Bed | A brachyceran fly | |||
| Atalosciophila | A. yanensis | A mycetophilid gnat[113] | ||||
| Chironomaptera | C. gregaria | A phantom midge | ||||
| Gigantoberis | G. liaoningensis | A soldier fly[114] | ||||
| Lichnoplecia | L. kovalevi | A protopleciid fly[115] | ||||
| Palaepangonius | P. eupterus | A horse-fly[97] | ||||
| Protapiocera | P. megista | An asiloid fly[97] | ||||
| Protempis | P. minuta | A brachyceran fly[97] | ||||
| Protonemestrius | P. jurassicus | A tangle-veined fly[97] |
Wasps (Hymenoptera)
[edit]| Genus | Species | County | Member | Abundance | Notes | Images |
|---|---|---|---|---|---|---|
| Abropelecinus | A. annulatus | Beipiao | An iscopinine pelecinid[116] | |||
| Angaridyela | A. robusta | Jianshangou Bed | A sawfly | |||
| Azygopelecinus | A. clavatus | Beipiao |
An iscopinine pelecinid[116] | |||
| Chengdeserphus | C. petidatus | A proctotrupid wasp[97] | ||||
| Cretevania | C. exquisita | Beipiao | Hatchet wasps[117] | |||
| C. pristine | Jianshangou bed | |||||
| C. vesca | ||||||
| Hoplitolyda | H. duolunica | The largest known sawfly ever[118] | ||||
| Liaotoma | L. linearis | A xyelotomid woodwasp[97] | ||||
| Manlaya | M. flexuosus | An aulacid woodwasp[97] | ||||
| Procretevania | P. exquisita |
Reclassified as Cretevania spp.[119] | ||||
| P. pristine | ||||||
| P. vesca | ||||||
| Rudisiricius | R. belli |
Praesiricid sawflies[120] | ||||
| R. celsus | ||||||
| R. crassinodus | ||||||
| Scolichneumon | S. rectivenius | An ichneumonoid wasp[97] | ||||
| Sinopelecinus | S. viriosus | A pelecinid[116] | ||||
| Tanychora | T. exquisita | Jianshangou Bed | An ichneumon wasp |
Other insects
[edit]| Genus | Species | County | Member | Abundance | Notes | Images |
|---|---|---|---|---|---|---|
| Abrigramma | A. calophleba | A kalligrammatid lacewing[121] |  | |||
| Alloraphidia | A. bliquivenatica[92] | Snakeflies | ||||
| A. longistigmasa | Jianshangou Bed | |||||
| Archeraphidia | A. shangyuanensis | A snakefly[92] | ||||
| Baissopterus | B. liaoningensis | A snakefly[92] | ||||
| Caenoephemera | C. shangyuanensis | Jianshangou Bed | A mayfly[92] | |||
| Caloraphidia | C. glossophylla | A snakefly[97] | ||||
| Choromyrmeleon | C. othneius | An antlion[97] | ||||
| Circulaboilus | C. aureus | A grig[122] | ||||
| Cretophasmomima | C. melanogramma | A stick insect[123] |  | |||
| ?Ephemeropsis | ?E. trisetalis | Jianshangou and Jingangshan beds | A mayfly. Chinese specimens probably represent species of Epicharmeropsis[124] | |||
| Epicharmeropsis | E. hexavenulosus[124] | Pingquan | Mayflies | |||
| E. quadrivenulosus[124] | Beipiao | Jianshangou Bed |  | |||
| Hagiphasma | H. paradoxa | Jianshangou Bed | A stick insect | |||
| Ilerdocossus | I. beipiaoensis | Palaeontinid hemipterans[125][126] | ||||
| I. exiguus | ||||||
| I. exquisitus | ||||||
| I. fengningensis | ||||||
| I. hui | ||||||
| I. pingquanensis | ||||||
| Ithigramma | I. multinervia | Kalligrammatid lacewings[121] |  | |||
| Kalligramma | K. liaoningensis | Jianshangou Bed | ||||
| Karatavoblatta | K. formosa | A cockroach[92] | ||||
| Liaoaphidia | L. fornicata | A snakefly[92] | ||||
| Liaonemobius | L. tanae | A sword-tail cricket[97] | ||||
| Longicellochrysa | L. yixiana | A mesochrysopid lacewing[127] | ||||
| Mesascalaphus | M. yangi | An owlfly[97] | ||||
| Mesolygaeus | M. laiyangensis | A water bug[92] | ||||
| Metabittacus | M. colosseus | Jianshangou Bed | A scorpionfly[128] | |||
| Miracossus | M. gongi | Palaeontinid hemipterans[125] | ||||
| M. ingentius | ||||||
| Nipponoblatta | N. acerba | Jianshangou Bed | A cockroach | |||
| Oloberotha | O. sinica | A beaded lacewing[97] | ||||
| Oregramma | O. aureolosa | Kalligrammatid lacewings[121] |  | |||
| O. illecebrosa |  | |||||
| Pseudopulex | P. magnus | Stem-fleas[129][130] | ||||
| P. tanlan | .jpg) | |||||
| Renphasma | R. sinica | A stick insect[131] |  | |||
| Saurophthirus | S. exquisitus | Stem-fleas[132] | ||||
| S. laevigatus |  | |||||
| Sinochresmoda | S. magnicornia | A chresmodid water-skiing insect[133] | ||||
| Sinosharaperla | S. zhaoi | Beipiao | A stonefly[134] | |||
| Sinoviparosiphum | S. lini | An aphid[97] | ||||
| Sophogramma | S. lii | Jianshangou Bed | Kalligrammatid lacewings[135][121] |  | ||
| S. pingquanica | ||||||
| Tetraphelerus | T. laetus | A moth[92] | ||||
| Tyrannopsylla | T. beipiaoensis | A stem-flea[136] | ||||
| Varicapitatus | V. sinuolatus | "Huangbanjigou, Chaomidian Village, Beipiao City."[137] | A hemipteran | |||
| Unnamed Vitimotauliidae | Unnamed | A caddisfly pupa, likely belonging to the Vitimotauliidae[138] |
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.[139]
| 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 | |||||
|
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 |  | ||
| A. liaoningensis | Jianshangou Bed | |||||
| A. sinensisis | Dawangzhangzi Bed | |||||
| Callianthus | C. dilae[140] | Liaoning | A plant with true flowers | |||
| ?Davallia | D.? niehhutzuensis[16] | |||||
| Erenia | E. stenoptera[140] | Liaoning | Actually a specimen of Callianthus | |||
| Leefructus | L. mirus[141] | 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[142] | |||||
| 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. |  | |||
| Chaoyangia | C. liangii[143] | 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[143] | A grass-like gnetale gymnosperm | ||||
| Siphonospermum | S. simplex[144] | Beipiao | Jianshangou Bed | A gnetale gymnosperm | ||
| Tyrmia | T. sp.[16] | A bennettite |
References
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