Paleobiota of the Yixian Formation

The

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

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

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

Ankylosaurs

Genus	Species	County	Member	Material	Notes	Images
Liaoningosaurus	L. paradoxus	Yixian	Dawangzhangzi Bed	"Partial skull [and] skeleton."^[8]	A small ankylosaur	Liaoningosaurus

Ceratopsians

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	Liaoceratops Psittacosaurus

Ornithopods

Genus	Species	County	Member	Material	Notes	Images
Bolong	B. yixianensis		Jianshangou Bed	A nearly complete skeleton.	An 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

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

Avialans

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]
Jinzhouornis	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]
Shanweiniao	S. cooperorum		Dawangzhangzi Bed		An enantiornithine.^[31]
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^[32]

Compsognathids

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^[33]^[34]	A compsognathid. The first non-avian dinosaur found with feather impressions.
Sinosauropteryx	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.^[35]

Dromaeosaurs

Genus	Species	County	Member	Material	Notes	Images
Changyuraptor^[36]	C. yangi	Jianchang		A complete articulated specimen with extensive feathering.^[36]	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.
Sinornithosaurus	S. millenii	Beipiao	Jianshangou Bed	Three specimens with skulls and postcrania, two with preserved feathers.^[37] Feathered integument is also preserved.^[37]^[38]	A dromaeosaurid, the first ever found with feather impressions.
Tianyuraptor	T. ostromi	Lingyuan	Dawangzhangzi Bed		A dromaeosaurid
Zhenyuanlong^[39]	Z. suni	Jianchang		One nearly complete skeleton with preserved feathers, including long tail feathers and large wing feathers.	A dromaeosaurid
Zhongjianosaurus^[40]	Z. yangi	Lingyuan		A partial skeleton includes an arm, legs, vertebrae, ribs, and most of the tail.^[40]	A microraptorian dromaeosaurid with very long legs.^[40]
Unnamed dromaeosaurid	Specimen NGMC 91	Lingyuan	Dawangzhangzi Bed	Complete specimen with feather imprints	A juvenile dromaeosaurid that may be a specimen of Sinornithosaurus millenii.^[41]^[42] Informally nicknamed "Dave".

Ornithomimosaurs

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^[43]	Shenzhousaurus
Shenzhousaurus	S. orientalis	Beipiao	Lujiatun Bed	"Articulated skull and incomplete skeleton."^[44]	A primitive ornithomimosaur

Oviraptorosaurs

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."^[45]	Oviraptorosaurs	Caudipteryx zoui Incisivosaurus Protarchaeopteryx
	C. zoui	Beipiao	Jianshangou Bed	Common, only over a small region
	C. sp.^[46]
Incisivosaurus	I. gauthieri	Beipiao	Lujiatun Bed	"[One] skull [and a] partial cervical."^[45]	An oviraptorosaur
Ningyuansaurus	N. wangi			Complete skeleton with feather impressions.^[47]	An oviraptorosaur
Protarchaeopteryx	P. robusta	Beipiao	Jianshangou Bed	"Nearly complete skull and skeleton, at least [two] individuals."^[48]	A possible relative of Incisivosaurus
Similicaudipteryx	S. yixianensis			Two specimens	An oviraptorosaur, also present in the Jiufotang Formation^[49]
Xingtianosaurus	X. ganqi	Liaoning	Dakangpu Bed	One nearly complete specimen (IVPP V13390), minus the head	An oviraptorosaur^[50]

Therizinosaurs

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."^[51]	A therizinosaur	Beipiaosaurus Jianchangosaurus
Jianchangosaurus	J. yixianensis	Jianchang		A nearly complete juvenile specimen with feather impressions.^[52]	A therizinosaur

Troodontids

Genus	Species	County	Member	Material	Notes	Images
Daliansaurus	D. liaoningensis	Beipiao	Lujiatun Bed	A very complete specimen.^[53]	A troodontid	Daliansaurus Jianianhualong Liaoningvenator Mei Sinovenator Sinusonasus
Jianianhualong	J. tengi	Beipiao	Dawangzhangzi Bed	A very complete specimen with feather impressions.^[54]	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.^[55]^[56] 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.^[57]
Undescribed troodontid	Specimen CAGS-IG01-004		Lujiatun Bed	Skull and partial skeleton	A troodontid similar to Mei.^[58]

Tyrannosaurs and other theropods

Color key

Taxon	Reclassified taxon	Taxon falsely reported as present	Dubious taxon or junior synonym	Ichnotaxon	Ootaxon	Morphotaxon

Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.

Genus	Species	County	Member	Material	Notes	Images
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.^[59]
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.^[60]
Yutyrannus	Y. huali		Unknown	Three nearly complete skeletons of various ages	A large tyrannosauroid^[61]
Unnamed theropod	Not yet described	Yixian	Lujiatun Bed	Partial post-cranial skeleton^[62]	A possible carnosaur^[63]

Pterosaurs

The pterosaurs listed below were compiled in a survey by Wang and Zhou in 2006, unless otherwise noted.^[64]

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^[65]	E. prolatus		Jianshangou Bed^[66]		A ctenochasmatid
Eopteranodon	E. lii^[67]	Beipiao			A tapejarid
Eopteranodon	E. yixianensis^[68]	Beipiao	Jianshangou Bed		A tapejarid
Eosipterus	E. yangi		Jianshangou Bed		A ctenochasmatid
Gegepterus^[69]	G. changae^[66]		Jianshangou Bed^[66]		A ctenochasmatid
Gladocephaloideus	G. jingangshanensis		Jianshangou Bed		A ctenochasmatid
Haopterus	H. gracilis		Jianshangou Bed		An ornithocheirid
Luchibang^[70]	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^[71]
Nurhachius	N. sp.		Jingangshan Member^[72]	An incomplete specimen^[72]	An istiodactylid
Pterofiltrus	P. qiui		Jianshangou Bed		A ctenochasmatid^[66]
Yixianopterus	Y. jingangshanensis				An istiodactyliform
Zhenyuanopterus	Z. longirostris		Jianshangou Bed		A boreopterid.^[73] May be adult form of Boreopterus.^[74]

Misc. reptiles

Choristoderans

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	Member	Material	Notes	Images
Hyphalosaurus	H. lingyuanensis	Dawangzhangzi Bed			Hyphalosaurus baitaigouensis Monjurosuchus splendens
Hyphalosaurus	H. baitaigouensis	Jingangshan Bed	Thousands of specimens
Monjurosuchus	M. splendens	Dawangzhangzi Bed
Sinohydrosaurus	S. lingyuanensis	Dawangzhangzi Bed		Junior synonym of Hyphalosaurus lingyuanensis

Lizards

Genus	Species	Member	Notes	Images
Dalinghosaurus	D. longidigitus	Jianshangou Bed	A lizard related to shinisaurs	Dalinghosaurus Xianglong
Liushusaurus	L. acanthocaudata^[75]	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

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	Member	Notes	Images
Manchurochelys	M. liaoxiensis^[76]	Jianshangou Bed	Reclassified as Ordosemys liaoxiensis^[77]	Ordosemys
Manchurochelys	M. manchoukuoensis^[16]	Jingangshan Bed
Ordosemys	O. liaoxiensis^[77]	Jianshangou Bed

Mammals

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.^[78]

Genus	Species	County	Member	Notes	Images
Acristatherium	A. yanensis^[79]	Beipiao	Lujiatun Bed	A eutherian	Eomaia Gobiconodon zofiae Jeholodens Repenomamus robustus Yanoconodon
Akidolestes	A. cifellii	Lingyuan	Dawangzhangzi Bed	A symmetrodont "
Ambolestes^[80]	A. zhoui	Nincheng	Jianshangou Bed	A eutherian
Chaoyangodens^[81]	C. lii	Lingyuan	Dawangzhangzi Bed	A eutriconodont
Eomaia	E. scansoria	Lingyuan	Dawangzhangzi Bed	A stem-therian
Gobiconodon	G. zofiae	Beipiao	Lujiatun Bed	A gobiconodont
Gobiconodon	G. luoianus^[82]		Lujiatun Bed	A gobiconodont; junior synonym of G. zofiae
Jeholodens	J. jenkinsi	Chaoyang	Jianshangou Bed	A eutriconodont
Juchilestes^[83]	J. liaoningensis	Beipiao	Lujiatun Bed	An amphidontid
Jueconodon^[84]	J. cheni			A eutriconodont
Maotherium	M. sinensis	Chaoyang	Jianshangou Bed	A zhangeotheriid "symmetrodont"
Maotherium	M. asiaticus^[85]	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
Repenomamus	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^[86]	Y. allini^[86]	Fengning^[86]		A eutricondodont
Zhangheotherium	Z. quinquecuspidens	Chaoyang	Jianshangou Bed	A zhangeotheriid "symmetrodont"

Amphibians

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	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
Liaobatrachus	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

Genus	Species	Member	Abundance	Notes	Images
Hybodontoidea indet.	Family, genus and species indeterminate			An elasmobranch known from a single dorsal fin spine^[87]	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

The invertebrate species listed below follow a survey by Chen and colleagues published in 2005, unless otherwise noted.^[76]

Arachnids

The arachnid species listed below follow a catalogue compiled by Dunlop, Penney and Jekel in 2010, unless otherwise noted.^[88]

Genus	Species	Notes
?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)

Genus	Species	County	Notes
Anoixis	A. complanus		A desmatin click beetle^[89]
Apoclion	A. antennatus		Desmatin click beetles^[89]
	A. clavatus
	A. dolini
Coptoclava	C. longipoda		A beetle^[90]
Desmatinus	D. cognatus		A desmatin click beetle^[89]
Fortiseode	F. pervalimand		An eodromeine trachypachid beetle^[91]
Glaresis	G. orthochilus		A laresid scarab beetle^[92]
Hydrophilopsia	H. gracilis		Hydrophiloid water scavenger beetles^[93]
	H. hydraenoides
	H. shatrovskiyi
Lasiosyne	L. integera	Huangbanjigou Village, Liaoning Province, northeastern China.^[94]	A lasiosynid beetle.^[94]
Lasiosyne	L. parva	Huangbanjigou Village, Liaoning Province, northeastern China.^[94]	A lasiosynid beetle.^[94]
Liaoximordella	L. hongi		A mordellid beetle^[95]
Mirimordella	M. gracilicruralis		A mordellid beetle^[96]
Oxyporus	O. yixianus		An oxyporine rove beetle^[97]
Paradesmatus	P. dilatatus		A desmati click beetle^[89]
Parageotrupes	P. incanus		A Geotrupine scarab beetle^[98]
Sinocupes	S. validus		A beetle^[90]
Sinosperchopsis	S. silinae		A hydrophiloid water scavenger beetle^[93]
Sinoxytelus	S. breviventer		Oxyteline rove beetles^[99]
	S. euglypheus
	S. longisetosus

Crustaceans

Genus	Species	County	Member	Notes
Chenops^[100]	C. oblongus			Notostracans
Chenops^[100]	C. yixianensis			Notostracans
Cricoidoscelosus	C. aethus			Junior synonym of Palaeocambarus^[101]
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^[100]	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^[102]
Mongolianella	M. cf. palmosa		Jianshangou Bed	An ostracod
Palaeocambarus	P. licenti			A crayfish^[103]
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)

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.^[104]

Genus	Species	County	Member	Abundance	Notes	Images
Abrohemeroscopus	A. mengi				A dragonfly	Sinaeschnidia sp.
Aeschnidium	A. heishankowense				Reclassified as Sinaeschnidia heishankowense^[105]
Aeschnidium	A. sp.				Reclassified as Sinaeschnidia heishankowense^[105]
Archaeogomphus	A. labius			Nymphs^[105]	Reclassified as Yixiangomphus labius,^[106] which is a junior synonym of Sinaeschnidia cancellosa^[105]
Bellabrunetia	B. catherinae				A dragonfly
Chrysogomphus	C. beipiaoensis				A dragonfly
Dissurus	D. liaoyuanensis^[106]			Poorly preserved nymph^[105]	Reclassified as Sinaeschnidia cf. cancellosa^[105]
Dracontaeschnidium	D. orientale				A dragonfly
Hebeiaeschnidia	H. fengningensis				A dragonfly
Huaxiagomphus	H. taushanensis				A dragonfly^[106]
Liogomphus	L. yixianensis				A dragonfly
Mesocordulia	M. boreala				A corduliid dragonfly^[95]
Palaeogomphus	P. labius				A dragonfly, reclassified as of Yixiangomphus labius,^[106] which is a junior synonym of Sinaeschnidia cancellosa^[105]
Parapetala	P. liaoningensis				A dragonfly
Parapetala	P. yixianensis		Jianshangou Unit^[107]
Protoliupanshania	P. wangi		Jianshangou Unit^[107]		A liupanshaniid dragonfly
Pseudocymatophlebia	P. bisecta	Ningcheng County^[108]		A well-preserved specimen.^[108]	An aktassiid dragonfly
Pseudoliupanshania	P. magnanicellula		Jianshangou Unit^[107]
Pseudosamarura	P. largina^[90]			One specimen, nymph^[105]	Reclassified as Sinaeschnidia cf. cancellosa^[105]
Rudiaeschna	R. limnobia				An aeshnid dragonfly^[95]
Sinaeschnidia	S. heishankowense		Jianshangou & Dawangzhangzi Beds	Adults and nymphs^[105]
	S. cancellosa^[105]				Junior synonym of S. heishankowense^[109]
	S. sp.			Indeterminate nymph specimens^[105]	Some nymph specimens have been given their own genera and species though probably belong so S. heishankowense but too poorly preserved to be certain^[105]
Sinogomphus	S. taoshanensis				A dragonfly
Sinojagoria	S. imperfecta				A dragonfly
Sopholibellula	S. amoena				Dragonflies
Sopholibellula	S. eleganti				Dragonflies
Stylaeschnidium	S. rarum				A dragonfly
Telmaeshna	T. paradoxica	Beipiao			A dragonfly
Turanophlebia	T. liaoningensis	Beipiao^[110]	Jianshangou Unit^[110]		A tarsophlebiid dragonfly.
Yixiangomphus	Y. labius			Nymphs^[105]	Junior synonym of Sinaeschnidia cancellosa^[105]

Flies (Diptera)

Genus	Species	Member	Notes
Alleremonomus	A. xingi	Jianshangou Bed	A brachyceran fly
Atalosciophila	A. yanensis		A mycetophilid gnat^[111]
Chironomaptera	C. gregaria		A phantom midge
Gigantoberis	G. liaoningensis		A soldier fly^[112]
Lichnoplecia	L. kovalevi		A protopleciid fly^[113]
Palaepangonius	P. eupterus		A horse-fly^[95]
Protapiocera	P. megista		An asiloid fly^[95]
Protempis	P. minuta		A brachyceran fly^[95]
Protonemestrius	P. jurassicus		A tangle-veined fly^[95]

Wasps (Hymenoptera)

Genus	Species	County	Member	Notes
Abropelecinus	A. annulatus	Beipiao		An iscopinine pelecinid^[114]
Angaridyela	A. robusta		Jianshangou Bed	A sawfly
Azygopelecinus	A. clavatus	Beipiao		An iscopinine pelecinid^[114]
Chengdeserphus	C. petidatus			A proctotrupid wasp^[95]
Cretevania	C. exquisita	Beipiao		Hatchet wasps^[115]
	C. pristine		Jianshangou bed
	C. vesca
Hoplitolyda	H. duolunica			The largest known sawfly ever^[116]
Liaotoma	L. linearis			A xyelotomid woodwasp^[95]
Manlaya	M. flexuosus			An aulacid woodwasp^[95]
Procretevania	P. exquisita			Reclassified as Cretevania spp.^[117]
	P. pristine
	P. vesca
Rudisiricius	R. belli			Praesiricid sawflies^[118]
	R. celsus
	R. crassinodus
Scolichneumon	S. rectivenius			An ichneumonoid wasp^[95]
Sinopelecinus	S. viriosus			A pelecinid^[114]
Tanychora	T. exquisita		Jianshangou Bed	An ichneumon wasp

Other insects

Genus	Species	County	Member	Notes	Images
Abrigramma	A. calophleba			A kalligrammatid lacewing^[119]
Alloraphidia	A. bliquivenatica^[90]			Snakeflies
Alloraphidia	A. longistigmasa		Jianshangou Bed	Snakeflies
Archeraphidia	A. shangyuanensis			A snakefly^[90]
Baissopterus	B. liaoningensis			A snakefly^[90]
Caenoephemera	C. shangyuanensis		Jianshangou Bed	A mayfly^[90]
Caloraphidia	C. glossophylla			A snakefly^[95]
Choromyrmeleon	C. othneius			An antlion^[95]
Circulaboilus	C. aureus			A grig^[120]
Cretophasmomima	C. melanogramma			A stick insect^[121]	Cretophasmomima, a stick insect
?Ephemeropsis	?E. trisetalis		Jianshangou and Jingangshan beds	A mayfly. Chinese specimens probably represent species of Epicharmeropsis^[122]
Epicharmeropsis	E. hexavenulosus^[122]	Pingquan		Mayflies
Epicharmeropsis	E. quadrivenulosus^[122]	Beipiao	Jianshangou Bed	Mayflies
Hagiphasma	H. paradoxa		Jianshangou Bed	A stick insect
Ilerdocossus	I. beipiaoensis			Palaeontinid hemipterans^[123]^[124]
	I. exiguus
	I. exquisitus
	I. fengningensis
	I. hui
	I. pingquanensis
Ithigramma	I. multinervia			Kalligrammatid lacewings^[119]
Kalligramma	K. liaoningensis		Jianshangou Bed	Kalligrammatid lacewings^[119]
Karatavoblatta	K. formosa			A cockroach^[90]
Liaoaphidia	L. fornicata			A snakefly^[90]
Liaonemobius	L. tanae			A sword-tail cricket^[95]
Longicellochrysa	L. yixiana			A mesochrysopid lacewing^[125]
Mesascalaphus	M. yangi			An owlfly^[95]
Mesolygaeus	M. laiyangensis			A water bug^[90]
Metabittacus	M. colosseus		Jianshangou Bed	A scorpionfly^[126]
Miracossus	M. gongi			Palaeontinid hemipterans^[123]
Miracossus	M. ingentius			Palaeontinid hemipterans^[123]
Nipponoblatta	N. acerba		Jianshangou Bed	A cockroach
Oloberotha	O. sinica			A beaded lacewing^[95]
Oregramma	O. aureolosa			Kalligrammatid lacewings^[119]
Oregramma	O. illecebrosa			Kalligrammatid lacewings^[119]
Pseudopulex	P. magnus			Stem-fleas^[127]^[128]
Pseudopulex	P. tanlan			Stem-fleas^[127]^[128]
Renphasma	R. sinica			A stick insect^[129]
Saurophthirus	S. exquisitus			Stem-fleas^[130]
Saurophthirus	S. laevigatus			Stem-fleas^[130]
Sinochresmoda	S. magnicornia			A chresmodid water-skiing insect^[131]
Sinosharaperla	S. zhaoi	Beipiao		A stonefly^[132]
Sinoviparosiphum	S. lini			An aphid^[95]
Sophogramma	S. lii		Jianshangou Bed	Kalligrammatid lacewings^[133]^[119]
Sophogramma	S. pingquanica			Kalligrammatid lacewings^[133]^[119]
Tetraphelerus	T. laetus			A moth^[90]
Tyrannopsylla	T. beipiaoensis			A stem-flea^[134]
Varicapitatus	V. sinuolatus	"Huangbanjigou, Chaomidian Village, Beipiao City."^[135]		A hemipteran
Unnamed Vitimotauliidae	Unnamed			A caddisfly pupa, likely belonging to the Vitimotauliidae^[136]

Molluscs

Genus	Species	Member	Notes
Amplovalvata	A. sp.	Jianshangou Bed	A gastropod
Arguniella	A. lingyuanensis	Jianshangou Bed	A bivalve, also present in the Jiufotang and Fuxin Formations
Arguniella	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
Corbicula	C. jeholense	Jianshangou Bed	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
Ptychastylus	P. cf. phillipi	Jianshangou Bed	Gastropods
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

Conifers

The coniferous trees and shrubs listed below were compiled in a survey by Wang and colleagues in 2006, unless otherwise noted.^[137]

Genus	Species	Member	Notes
Brachyphyllum			A cypress-like conifer
Cephalotaxopsis			A yew
Cunninghamia			A cypress
Elatides			A cypress
Elatocladus	E. leptophyllus		Conifers of uncertain classification
Elatocladus	E. manchurica^[16]		Conifers of uncertain classification
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

Genus	Species	Notes
Cladophlebis	C. spp.^[16]
Coniopteris	C. sp.^[16]
Onychiopsis	O. elongata^[16]	Junior synonym of O. psilotoides
Onychiopsis	O. psilotoides^[16]
?Ruffordia	R.? sp.^[16]

Flowering plants

Genus	Species	County	Member	Notes	Images
Archaefructus	A. eoflora		Jianshangou Bed	A primitive, aquatic angiosperm	Archaefructus
	A. liaoningensis		Jianshangou Bed
	A. sinensisis		Dawangzhangzi Bed
Callianthus	C. dilae^[138]	Liaoning		A plant with true flowers
?Davallia	D.? niehhutzuensis^[16]
Erenia	E. stenoptera^[138]	Liaoning		Actually a specimen of Callianthus
Leefructus	L. mirus^[139]			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

Genus	Species	County	Member	Abundance	Notes	Images
Baiera	B. cf. gracilis^[16]
Baiera	B. orientalis^[16]
Czekanowskia	C. rigida^[16]
Ginkgo	G. apodes^[140]
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

Genus	Species	County	Member	Notes
Chaoyangia	C. liangii^[141]			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^[141]			A grass-like gnetale gymnosperm
Siphonospermum	S. simplex^[142]	Beipiao	Jianshangou Bed	A gnetale gymnosperm
Tyrmia	T. sp.^[16]			A bennettite

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[1] PMID 34691675
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[Wangetal07-2] S2CID 128812879
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[titanosaurteeth-3] 
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