Paleobiota of the Yixian Formation

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

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

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	Lycoptera davidi Protosephurus Sinamia
	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

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	Repenomamus robustus Sinodelphys Zhangheotherium
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

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	Haopterus Ningchengopterus
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?		A ctenochasmatoid
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

Choristoderans

Genus	Species	County	Member	Abundance	Notes	Images
Hyphalosaurus	H. lingyuanensis		Dawangzhangzi Bed			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	County	Member	Abundance	Notes	Images
Dalinghosaurus	D. longidigitus		Jianshangou Bed		A lizard related to shinisaurs	Dalinghosaurus
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

Genus	Species	County	Member	Abundance	Notes	Images
Manchurochelys	M. liaoxiensis[18]		Jianshangou Bed		Reclassified as Ordosemys liaoxiensis[19]	Ordosemys
	M. manchoukuoensis[20]		Jingangshan Bed
Ordosemys	O. liaoxiensis[19]		Jianshangou Bed

Ornithischians

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 Liaoceratops Psittacosaurus
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

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

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
Archaeorhynchus	A. spathula		Dawangzhangzi Bed		An early, advanced short-tailed bird (ornithuran)[26]	Changchengornis Confuciusornis Eoenantiornis Liaoxiornis Shanweiniao
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 entantiornithine, 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

Genus	Species	County	Member	Material	Notes	Images
Graciliraptor	G. lujiatunensis	Beipiao	Lujiatun Bed	Rare	A dromaeosaurid	Graciliraptor Mei Sinornithosaurus millenii Sinusonasus Tianyuraptor
Mei	M. long	Beipiao	Lujiatun Bed		A small troodontid fossilized in a sleeping position
Sinornithosaurus	S. haoiana	Yixian	Dawangzhangzi Bed		A dromaeosaurid, the first ever found with feather impressions
	S. millenii	Beipiao	Jianshangou Bed	Three specimens with skulls and postcrania, two with preserved feathers.[43] Feathered integument is also preserved.[43][44]
Sinovenator	S. changii	Beipiao	Lujiatun Bed	Three specimens with partial skulls and postcrania.[45][46] 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	Z. suni			One nearly complete skeleton with traces of feathers, including long tail feathers and large wing feathers.	A dromaeosaurid
Undescribed deinonychosaur/possible troodontid			Dawangzhangzi Bed	One specimen with feather imprints	A possible troodontid.[47]
Undescribed troodontid	Specimen CAGS-IG01-004		Lujiatun Bed	Skull and partial skeleton	A troodontid similar to Mei.[48]
Unnamed dromaeosaurid	Specimen NGMC 91	Lingyuan	Dawangzhangzi Bed	Complete specimen with feather imprints	Once tentatively referred to both Sinornithosaurus and Cryptovolans, it is probably a new genus closely related to Microraptor.[49] Informally nicknamed "Dave"

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
Beipiaosaurus	B. inexpectus	Beipiao	Jianshangou Bed	"Incomplete skeleton with integument."[50]	A therizinosaur	Beipiaosaurus Caudipteryx zoui Dilong Huaxiagnathus Incisivosaurus Protarchaeopteryx Shenzhousaurus Sinocalliopteryx Sinosauropteryx prima Yutyrannus
Caudipteryx	C. dongi	Beipiao	Jianshangou Bed	"[One] partial postcranium with feather imprints."[51]	Oviraptorosaurs
	C. zoui	Beipiao	Jianshangou Bed	Common, only over a small region
	C. sp.[52]
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.[53]
Hexing	H. qingyi				A primitive ornithomimosaur[54]
Huaxiagnathus	H. orientalis	Beipiao	Jianshangou Bed		A compsognathid
Incisivosaurus	I. gauthieri	Beipiao	Lujiatun Bed	"[One] skull [and a] partial cervical."[51]	An oviraptorosaur
Ningyuansaurus	N. wangi			Complete skeleton with feather impressions.[55]	An oviraptorosaur
Protarchaeopteryx	P. robusta	Beipiao	Jianshangou Bed	"Nearly complete skull and skeleton, at least [two] individuals."[56]	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.[57]
Shenzhousaurus	S. orientalis	Beipiao	Lujiatun Bed	"Articulated skull and incomplete skeleton."[58]	A primitive ornithomimosaur
Similicaudipteryx	S. yixianensis			Two specimens	An oviraptorosaur, also present in the Jiufotang Formation[59]
Sinocalliopteryx	S. gigas		Jianshangou Bed		A large compsognathid
Sinosauropteryx	S. prima	Beipiao & Lingyuan	Jianshangou & Dawangzhangzi Beds	Four nearly complete specimens[60][61]	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.[62]
Yixianosaurus	Y. longimanus	Yixian	Dawangzhangzi Bed		A basal maniraptoran
Yutyrannus	Y. huali		Unknown	Three nearly complete skeletons of various ages	A large tyrannosauroid[63]
Unnamed theropod	Not yet described	Yixian	Lujiatun Bed	Partial post-cranial skeleton[64]	A possible carnosaur[65]

Sauropods

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
Dongbeititan	D. dongi	Jianshangou Bed		Partial postcrania[66]	A basal titanosauriform
cf. Euhelopus	cf. E. sp.		Lujiatun Bed	Isolated teeth[67]	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[67]

Invertebrate fauna of the Yixian Formation

The invertebrate species listed below follow a survey by Chen and colleagues published in 2005, unless otherwise noted.^[18]

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.

Arachnids

The arachnid species listed below follow a catalogue compiled by Dunlop, Penney and Jekel in 2010, unless otherwise noted.^[68]

Genus	Species	County	Member	Abundance	Notes	Images
?Araneus	A.? aethus				Orb-weaver spiders
	A.? beipiaoensis
	A.? liaoxiensis
	A.? reheensis
Cretaraneus	C. liaoningensis				A nephilid spider
?Gnaphosa	G.? liaoningensis				A ground spider
?Theridion	T.? atalus				A tangle web spider

Beetles (Coleoptera)

Genus	Species	County	Member	Abundance	Notes	Images
Anoixis	A. complanus				A desmatin click beetle[69]
Apoclion	A. antennatus				Desmatin click beetles[69]
	A. clavatus
	A. dolini
Coptoclava	C. longipoda				A beetle[70]
Desmatinus	D. cognatus				A desmatin click beetle[69]
Fortiseode	F. pervalimand				An eodromeine trachypachid beetle[71]
Glaresis	G. orthochilus				A laresid scarab beetle[72]
Hydrophilopsia	H. gracilis				Hydrophiloid water scavenger beetles[73]
	H. hydraenoides
	H. shatrovskiyi
Liaoximordella	L. hongi				A mordellid beetle[74]
Mirimordella	M. gracilicruralis				A mordellid beetle[75]
Oxyporus	O. yixianus				An oxyporine rove beetle[76]
Paradesmatus	P. dilatatus				A desmati click beetle[69]
Parageotrupes	P. incanus				A Geotrupine scarab beetle[77]
Sinocupes	S. validus				A beetle[70]
Sinosperchopsis	S. silinae				A hydrophiloid water scavenger beetle[73]
Sinoxytelus	S. breviventer				Oxyteline rove beetles[78]
	S. euglypheus
	S. longisetosus

Crustaceans

Genus	Species	County	Member	Abundance	Notes	Images
Chenops[79]	C. oblongus				A notostracans	Chenops yixianensis Chenops oblongus (right) and Jeholops hongi (left)
	C. yixianensis
Cricoidoscelosus	C. aethus				A crayfish[80]
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[79]	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[80]
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.^[81]

Genus	Species	County	Member	Abundance	Notes	Images
Abrohemeroscopus	A. mengi				A dragonfly
Aeschnidium	A. heishankowense				Reclassified as Sinaeschnidia heishankowense[82]
	A. sp.
Archaeogomphus	A. labius			Nymphs[82]	Reclassified as Yixiangomphus labius,[83] which is a junior synonym of Sinaeschnidia cancellosa[82]
Bellabrunetia	B. catherinae				A dragonfly
Chrysogomphus	C. beipiaoensis				A dragonfly
Dissurus	D. liaoyuanensis[83]			Poorly preserved nymph[82]	Reclassified as Sinaeschnidia cf. cancellosa[82]
Dracontaeschnidium	D. orientale				A dragonfly
Hebeiaeschnidia	H. fengningensis				A dragonfly
Huaxiagomphus	H. taushanensis				A dragonfly[83]
Liogomphus	L. yixianensis				A dragonfly
Mesocordulia	M. boreala				A corduliid dragonfly[74]
Palaeogomphus	P. labius				A dragonfly, reclassified as of Yixiangomphus labius,[83] which is a junior synonym of Sinaeschnidia cancellosa[82]
Parapetala	P. liaoningensis				A dragonfly
Pseudosamarura	P. largina[70]			One specimen, nymph[82]	Reclassified as Sinaeschnidia cf. cancellosa[82]
Rudiaeschna	R. limnobia				An aeshnid dragonfly[74]
Sinaeschnidia	S. heishankowense		Jianshangou & Dawangzhangzi Beds	Adults and nymphs[82]
	S. cancellosa[82]				Junior synonym of S. heishankowense[84]
	S. sp.			Indeterminate nymph specimens[82]	Some nymph specimens have been given their own genera and species though probably belong so S. heishankowense but too poorly preserved to be certain[82]
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[82]	Junior synonym of Sinaeschnidia cancellosa[82]

Flies (Diptera)

Genus	Species	County	Member	Abundance	Notes	Images
Alleremonmus	A. xingi		Jianshangou Bed		A brachyceran fly
Atalosciophila	A. yanensis				A mycetophilid gnat[85]
Chironomaptera	C. gregaria				A phantom midge
Gigantoberis	G. liaoningensis				A soldier fly[86]
Lichnoplecia	L. kovalevi				A protopleciid fly[87]
Palaepangonius	P. eupterus				A horse-fly[74]
Protapiocera	P. megista				An asiloid fly[74]
Protempis	P. minuta				A brachyceran fly[74]
Protonemestrius	P. jurassicus				A tangle-veined fly[74]

Wasps (Hymenoptera)

Genus	Species	County	Member	Abundance	Notes	Images
Abropelecinus	A. annulatus	Beipiao			An iscopinine pelecinid[88]
Angaridyela	A. robusta		Jianshangou Bed		A sawfly
Azygopelecinus	A. clavatus	Beipiao			An iscopinine pelecinid[88]
Chengdeserphus	C. petidatus				A proctotrupid wasp[74]
Cretevania	C. exquisita	Beipiao			Hatchet wasps[89]
	C. pristine		Jianshangou bed
	C. vesca
Liaotoma	L. linearis				A xyelotomid woodwasp[74]
Manlaya	M. flexuosus				An aulacid woodwasp[74]
Procretevania	P. exquisita				Reclassified as Cretevania spp.[90]
	P. pristine
	P. vesca
Rudisiricius	R. belli				Praesiricid sawflies[91]
	R. celsus
	R. crassinodus
Scolichneumon	S. rectivenius				An ichneumonoid wasp[74]
Sinopelecinus	S. viriosus				A pelecinid[88]
Tanychora	T. exquisita		Jianshangou Bed		An ichneumon wasp

Other insects

Genus	Species	County	Member	Abundance	Notes	Images
Alloraphidia	A. bliquivenatica[70]				Snakeflies	Epicharmeropsis, a mayfly
	A. longistigmasa		Jianshangou Bed
Archeraphidia	A. shangyuanensis				A snakefly[70]
Baissopterus	B. liaoningensis				A snakefly[70]
Caenoephemera	C. shangyuanensis		Jianshangou Bed		A mayfly[70]
Caloraphidia	C. glossophylla				A snakefly[74]
Choromyrmeleon	C. othneius				An antlion[74]
?Ephemeropsis	?E. trisetalis		Jianshangou and Jingangshan beds		A mayfly. Chinese specimens probably represent species of Epicharmeropsis[92]
Epicharmeropsis	E. hexavenulosus[92]	Pingquan			Mayflies
	E. quadrivenulosus[92]	Beipiao	Jianshangou Bed
Hagiphasma	H. paradoxa		Jianshangou Bed		A stick insect
Ilerdocossus	I. beipiaoensis				Cicadas[93]
	I. exiguus
	I. fengningensis
	I. hui
	I. pingquanensis
Kalligramma	K. liaoningensis		Jianshangou Bed		A lacewing
Karatavoblatta	K. formosa				A cockroach[70]
Liaoaphidia	L. fornicata				A snakefly[70]
Liaonemobius	L. tanae				A sword-tail cricket[74]
Longicellochrysa	L. yixiana				A mesochrysopid lacewing[94]
Mesascalaphus	M. yangi				An owlfly[74]
Mesolygaeus	M. laiyangensis				A water bug[70]
Metabittacus	M. colosseus		Jianshangou Bed
Miracossus	M. ingentius				A cicada[93]
Nipponoblatta	N. acerba		Jianshangou Bed		A cockroach
Oloberotha	O. sinica				A beaded lacewing[74]
Sinochresmoda	S. magnicornia				A chresmodid water-skiing insect[95]
Sinosharaperla	S. zhaoi	Beipiao			A stonefly[96]
Sinoviparosiphum	S. lini				An aphid[74]
Sophogramma	S. lii		Jianshangou Bed		A lacewing[97]
Tetraphelerus	T. laetus				A moth[70]
Unnamed Vitimotauliidae	Unnamed				A caddisfly pupa, likely belonging to the Vitimotauliidae.[98]
Unnamed Siphonaptera	Unnamed				A giant, non-jumping flea[99]

Molluscs

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

Conifers

The coniferous trees and shrubs listed below were compiled in a survey by Wang and colleagues in 2006, unless otherwise noted.^[100]

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

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

Genus	Species	County	Member	Abundance	Notes	Images
Archaeamphora	A. longicervia		Jianshangou Bed		A pitcher plant, and the earliest known carnivorous plant	Archaefructus
Archaefructus	A. eoflora		Jianshangou Bed		A primitive, aquatic angiosperm
	A. liaoningensis		Jianshangou Bed
	A. sinensisis		Dawangzhangzi Bed
Callianthus	C. dilae[101]	Liaoning			A plant with true flowers
?Davallia	D.? niehhutzuensis[20]
Erenia	E. stenoptera[101]	Liaoning			Actually a specimen of Callianthus
Leefructus	L. mirus[102]				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

Genus	Species	County	Member	Abundance	Notes	Images
Baiera	B. cf. gracilis[20]
	B. orientalis[20]
Czekanowskia	C. rigida[20]
Ginkgo	G. apodes[103]
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

Genus	Species	County	Member	Abundance	Notes	Images
Chaoyangia	C. liangii[104]				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[104]				A grass-like gnetale gymnosperm
Siphonospermum	S. simplex[105]	Beipiao	Jianshangou Bed		A gnetale gymnosperm
Tyrmia	T. sp.[20]				A cycad

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