Tiaojishan Formation

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Tiaojishan Formation
Stratigraphic range: Bathonian-Oxfordian
~165–153  Ma
Tiaojishan Formation.png
Exposure of the Tiaojishan Formation at Nanshimenzi Village, Qinglong Manchu Autonomous County, Hebei Province, with red arrow pointing to fossiliferous beds
Type Geological formation
Underlies Tuchengzi Formation, Houcheng Formation
Overlies Haifanggou Formation
Thickness2,420 m (7,940 ft)
Primary Andesite
Other Sandstone, shale, tuff, coal
Coordinates 41°18′N119°12′E / 41.3°N 119.2°E / 41.3; 119.2
Approximate paleocoordinates 43°00′N123°06′E / 43.0°N 123.1°E / 43.0; 123.1
Region Hebei, Inner Mongolia, & Liaoning
CountryFlag of the People's Republic of China.svg  China
Extent Yanshan Belt
China edcp relief location map.jpg
Blue pog.svg
Tiaojishan Formation (China)
China Liaoning relief location map.png
Blue pog.svg
Tiaojishan Formation (Liaoning)

The Tiaojishan Formation is a geological formation in Hebei and Liaoning, People's Republic of China, dating to the middle-late Jurassic period (Bathonian-Oxfordian stages). It is known for its exceptionally preserved fossils, including those of plants, insects and vertebrates. It is made up mainly of pyroclastic rock interspersed with basic volcanic and sedimentary rocks. Previously, the Tiaojishan Formation was grouped together with the underlying Haifanggou Formation (also known as the Jiulongshan Formation) as a single "Lanqi Formation." [1] The Tiaojishan Formation forms a key part of the Yanliao Biota assemblage, alongside the Haifanggou Formation.



Using Argon–argon dating, Wang and colleagues in 2005 dated part of the Tiaojishan Formation to about 160 million years ago, the beginning of the Oxfordian stage, the first stage of the Upper Jurassic epoch. [2] In 2006, a study by Liu and colleagues used U-Pb zircon dating to conclude that the Tiaojishan Formation correlates with the Daohugou Beds, and the complete chronological range of this shared biota dates to between 168 and 164/152 Ma ago. [3] A subsequent study, published in 2008, refined the age range of the formation further, finding that the lower boundary of the Tiaojishan was formed 165 Ma ago, and the upper boundary somewhere between 156 and 153 Ma ago. [4]


Based on the plant life present in the Tiaojishan Formation, Wang Yongdong and colleagues determined that the climate in Liaoning during the mid Jurassic would have been subtropical to temperate, warm and humid. [1]


Beautifully preserved fossils of dinosaurs, pterosaurs, salamanders, insects, arachnids [5] and other invertebrates, conifers, ginkgoes, cycads, horsetails, and ferns, and even the earliest known gliding mammal ( Volaticotherium ) have been discovered in these rocks.The tuffaceous composition of some rock layers show that this was a volcanic area, occasionally experiencing heavy ashfalls from eruptions. The landscape then was dominated by mountain streams and deep lakes surrounded by forests of gymnosperm trees. [6]

The forests of the Yanliao biota grew in a humid, warm - temperate climate and were dominated by gymnosperm trees. There were ginkgopsids like Ginkoites, Ginkgo , Baiera, Czekanowskia, and Phoenicopsis. There were also conifers like Pityophyllum, Rhipidiocladus, Elatocladus, Schizolepis, and Podozamites. Also, Lycopsids like Lycopodites and Sellaginellities, horsetails (Sphenopsida) like Equisetum, cycads like Anomozamites, and ferns (Filicopsida) like Todites and Coniopteris. [7]


Salamanders of the Tiaojishan Formation
GenusSpeciesLocationStratigraphic positionAbundanceNotesImages


B. jianpingensis [8]


A salamandroid known from an almost complete and articulated skeleton exposed in ventral view.

Chunerpeton Chunerpeton BW.jpg


C. tianyiensis

A cryptobranchoid measuring 18 centimeters in length.


J. paradoxus

A cryptobranchoid with a strange skull morphology, at first believed to come from the Early Cretaceous.


L. daohugouensis

A little-known cryptobranchoid.


P. sinensis

A cryptobranchoid characterized by its short trunk (only 14 presacrals) and short and wide head, giving a fat body shape, from which the genus name was derived ("Pang" means fat in Chinese).


Pterosaurs of the Tiaojishan Formation
GenusSpeciesLocationStratigraphic positionAbundanceNotesImages


A. linglongtaensis [9]


A monofenestratan known from an incomplete skeleton with a partial skull and lower jaw.

Darwinopterus modularis Darwinopterus NT.jpg
Darwinopterus modularis
Jeholopterus Jeholopterus BW.jpg
Kunpengopterus antipollicatus Kunpengopterus antipollicatus and Allaboilus gigantus.jpg
Kunpengopterus antipollicatus
Sinomacrops Life reconstruction of Sinomacrops bondei.png
Wukongopterus Wukongopterus NT.jpg


C. rong [10]


Daohugou bed [11]

One specimen

A long-tailed batrachognathine anurognathid known from a complete skeleton of a juvenile with extensive preservation of pycnofibres and wing membranes.


C. pani [12]


A pterodactyliform known only from a single specimen of a young juvenile, measuring 475 millimeters (18.7 in) in wingspan.


D. delicatus [13]

Inner Mongolia

One specimen

A relatively basal pterosaur known from a partial skeleton with soft tissue impressions.


D. modularis [14]


A wukongopterid named after Charles Darwin. The type species, D. modularis was the first known pterosaur to display features of both long-tailed rhamphorhynchoids and short-tailed pterodactyloids, and was described as a transitional fossil between the two groups. Darwinopterus specimens have also been reported to show several differences between males and females, with the males having distinctive crests on their heads. They are known to have laid their eggs on the ground, and may have also not shown that much for parental care.

D. linglongtaensis

D. robustodens

Dendrorhynchoides ?

D. curvidentatus?

One specimen

An anurognathid pterosaur of uncertain age. Originally reported from the Cretaceous aged Yixian Formation, it may be instead be from the mid-Jurassic Daohugou beds. [15]


D. zhengi [16]


One specimen

A non-pterodactyloid monofenestratan with a proportionately short tail.


F. lii [17]


A scaphognathine rhamphorhynchid similar to other scaphognathines in its short, blunt skull with a large antorbital fenestra, and widely spaced, vertically oriented teeth (as opposed to the horizontally-oriented teeth of other rhamphorhynchids).


J. ninchengensis

Inner Mongolia

Several specimens [18] [19]

A batrachognathine anurognathid preserved with pycnofibres and skin remains.


J. robustus [20]


A scaphognathine rhamphorhynchid known from a single fossil skeleton.


J. zhaoianus [21]


A scaphognathine rhamphorhynchid known from a nearly complete skeleton with the skull preserved.


K. sinensis [22]


Daohugou bed

Three specimens

A wukongopterid with an elongated head, 106.9 millimeters long, and an opposed thumb.

K. antipollicatus [23]


Two nearly complete specimens


L. primus [24]

Daxishan (Linglongta)

One specimen

A ctenochasmatid pterosaur with elongated, comb-like teeth.


L. mutoudengensis [15] [25]


One specimen

A batrachognathine anurognathid, originally thought to be from the Early Cretaceous, with a wingspan that is about 40 centimeters, making it one of the smallest known pterosaurs. Originally classed as a species of Dendrorhynchoides .


P. wellnhoferi

Inner Mongolia

Daohugou bed

One specimen [18]

A darwinopteran with a tall crest on its head and an elongated skull 11.8 centimeters (4.6 in) long, a long tail and a wingspan of about 85 centimeters (33 in). It was originally believed to be a rhamphorhynchid. The only known specimen consists of an articulated, nearly complete skeleton with remains of the integument. These included the wing membrane, hair-like pycnofibers, a long version of the vane found at the end of "rhamphorhynchoid" tails, and a head crest with both a low bony base and a large keratin extension.


Q. guoi [26]


A rhamphorhynchine rhamphorhynchid known from only one specimen that includes a skeleton with a skull.


S. bondei [11]


Daohugou bed

One specimen

A long-tailed batrachognathine anurognathid known from a relatively complete skull and skeleton with soft tissue patches.


W. lii


Daohugou bed

One specimen [27]

A wukongopterid unusual for having both an elongate neck and a long tail. Its wingspan is estimated at 730 millimeters (29 in).


Dinosaurs of the Tiaojishan Formation
GenusSpeciesLocationStratigraphic positionAbundanceNotesImages


A. huxleyi [28]


Several specimens [29]

An anchiornithid at first believed to be a troodont. Given the exquisite preservation of one of the first specimen's fossils, Anchiornis became the first dinosaur species for which almost the entire life coloration could be determined. Most of the body feathers of Anchiornis were gray and black. The crown of head feathers was mainly rufous with a gray base and front, and the face had rufous speckles among predominantly black head feathers. The wing and hind leg feathers were white with black tips. The coverts were gray, contrasting the mainly white main wings. The larger coverts of the wing were also white with gray or black tips, forming rows of darker dots along mid-wing. These took the form of dark stripes or even rows of dots on the outer wing (primary feather coverts) but a more uneven array of speckles on the inner wing (secondary coverts). The shanks of the legs were gray other than the long leg feathers, and the feet and toes were black. It was 13 inches long and weighed only 110 grams (3.9 ounces).

Anchiornis Anchiornis BW.jpg
Aurornis Aurornis.jpg
Caihong Caihong , life restoration.jpg
Eosinopteryx Eosinopteryx.jpg
Epidexipteryx Epidexipteryx NT.jpg
Scansoriopteryx Epidendrosaurus ningchingensis.png
Serikornis Serikornis.jpg
Tianyulong Tianyulong BW.jpg
Xiaotingia Xiaotingia .jpg
Yi Yi qi restoration.jpg


A. xui


One specimen

An anchiornithid roughly the size of a modern pheasant, with a length of 20 inches. Its leg bones were similar to those of Archaeopteryx , but overall its anatomy was more primitive.


C. juji


Yanliao Biota

One specimen

An anchiornithid known from an adult specimen measuring 400 mm in body length. Its fossilized feathers possess nanostructures which were analyzed and interpreted as melanosomes, showing similarity to organelles that produce a black iridescent color in certain species of extant birds. Other feathers found on the head, chest, and the base of the tail preserve flattened sheets of platelet-like melanosomes very similar in shape to those which create brightly colored iridescent hues in the feathers of modern hummingbirds. However, these structures are seemingly solid and lack air bubbles, and thus are internally more akin to the melanosomes in trumpeters than hummingbirds. Caihong represents the oldest known evidence of platelet-like melanosomes. It is named for the large crest on the lacrimal bone of the skull. [30]


E. brevipenna


One specimen

An anchiornithid at first believed to be a troodont, known from a single fossil specimen representing the nearly complete skeleton of a subadult or adult individual. The specimen is very small, measuring about 12 inches long.


E. hui

Inner Mongolia

Daohugou beds

One specimen

A scansoriopterygid known from a well-preserved partial skeleton, measuring 10 inches in length (17.5 inches including the incomplete tail feathers), that includes four long feathers on the tail, composed of a central rachis and vanes. However, unlike in modern-style rectrices, the vanes were not branched into individual filaments but made up of a single ribbon-like sheet. Epidexipteryx also preserved a covering of simpler body feathers, composed of parallel barbs as in more primitive feathered dinosaurs. However, the body feathers of Epidexipteryx are unique in that some appear to arise from a "membranous structure" at the base of each feather. It has been suggested that this may represent a stage in the evolution of the feather. Epidexipteryx and its kin represent the earliest known examples of ornamental feathers in the fossil record.


P. daohugouensis

Inner Mongolia

Daohugou beds

One specimen

An anchiornithid that probably measured 1 meter (3.3 feet) or less in length, but since this species is only known from the hind legs, the actual length is difficult to estimate. Apart from having a very birdlike skeletal structure in its legs, Pedopenna was remarkable due to the presence of long pennaceous feathers on the metatarsus (foot). Some other paravians are also known to have these 'hind wings', but those of Pedopenna differ from those of animals like Microraptor . Pedopenna hind wings were smaller and more rounded in shape. The longest feathers were slightly shorter than the metatarsus, at about 55 mm (2.2 in) long. Additionally, the feathers of Pedopenna were symmetrical, unlike the asymmetrical feathers of some other non-avian paravians and birds. Since asymmetrical feathers are typical of animals adapted to flying, it is likely that Pedopenna represents an early stage in the development of these structures. While many of the feather impressions in the fossil are weak, it is clear that each possessed a rachis and barbs, and while the exact number of foot feathers is uncertain, they are more numerous than in the hind-wings of Microraptor. Pedopenna also shows evidence of shorter feathers overlying the long foot feathers, evidence for the presence of coverts as seen in modern birds. Since the feathers show fewer aerodynamic adaptations than the similar hind wings of Microraptor, and appear to be less stiff, suggests that if they did have some kind of aerodynamic function, it was much weaker than in other non-avian paravians and birds.


S. heilmanni


Exact provenance of type specimen unknown, most likely from the Daohugou Beds [31]

One or two specimens

A sparrow-sized scansoriopterygid known from one or two juvenile specimens.


S. sungei


One specimen [32]

An anchiornithid with plumulaceous-like feathers. Feather imprints include wispy bundles along the neck, short and symmetrical vaned feathers on the arms, and both fuzz and long pennaceous feathers on its hind limbs (bearing a striking resemblance of the delicate hind limb filaments to the modern Silkie breed of domestic chicken). While its anatomy and integument share features with birds as well as derived dromaeosaurs such as Microraptor, cladistic analysis places the genus within the cluster of feathered non-avian dinosaurs near the origin of avialans. It was unlikely to be a flier.


T. confuciusi


A heterodontosaur that was initially reported as being from the Early Cretaceous Jehol group. The fossil was collected at a locality transliterated as Linglengta or Linglongta. Lu et al., 2010, reported that these beds were actually part of the Tiaojishan Formation, dating from the Late Jurassic period. Tianyulong has a row of long, filamentous integumentary structures on the back, tail and neck of the specimen, similar to the feathers found in certain theropods (this suggests that all heterodontosaurs may have had these filaments). The holotype is from a subadult individual that probably measured 70 centimeters in length based on the proportions of the related Southern African species Heterodontosaurus .


X. zhengi [33]


One specimen

An anchiornithid originally thought to be either a dromaeosaur or a troodont. It was morphologically similar to Archaeopteryx and was the size of a domestic chicken hen. It was about 60 cm long and weighed an estimated 0.82 kg. Like Archaeopteryx it had long forelimbs. Its femur was longer than its humerus, 84 mm as against 71 mm.


Y. qi [34]


Daohugou beds

One specimen

A gliding scansoriopterygid, weighing about 380 grams (0.84 pounds), that, like other scansoriopterygids, possessed an unusual, elongated third finger, that (in the case of Yi) helped to support a membranous gliding plane made of skin. The planes of Yi were also supported by a long, bony strut attached to the wrist. This modified wrist bone and membrane-based plane is unique among all known dinosaurs, and might have resulted in wings similar in appearance to those of bats. This also leads to the hypothesis that the other two genera of scansoriopterygids also had gliding membranes, but this is yet to be proven official.


Color key
Taxon Reclassified taxonTaxon falsely reported as presentDubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.
Lepidosaurs (lizards and relatives) of the Daohugou Beds

Unnamed lizard [35]

Inner Mongolia

One specimen

A new lizard with relatively short forelimbs

Unnamed lizard [35]

Inner Mongolia

One specimen

A lizard with long hind limbs and a narrow body


Cynodonts of the Tiaojishan Formation
GenusSpeciesLocationStratigraphic positionAbundanceNotesImages


A. scansorius

Inner Mongolia

Daohugou bed

A shrew-sized, arboreal docodontid that is known to be one of the earliest tree-climbing mammaliaforms. [36] It measured approximately 13 centimeters from head to tail and weighed about 27 grams. Its appearance was similar to a squirrel, with a long snout, curved, horny claws and flexible ankle and wrist joints typical of modern arboreal mammals. The front teeth were spade-like, indicating that Agilodocodon could gnaw tree bark and consume gum or sap.

Agilodocodon Agilodocodon scansorius by Kaek.jpg
Docofossor Docofossor NT flipped.jpg
Juramaia Juramaia NT.jpg
Maiopatagium Maiopatagium NT.jpg
Volaticotherium Volaticotherium.jpg


A. jenkinsi


One specimen

An arboreal, prehensile-tailed euharamiyid haramiyidan that was the largest known haramiyidan, estimated to have weighed about 354 grams. [37] Arboroharamiya is unlike any modern mammal in having a lower jaw that can move up, down, and backward, but not forward. It has a rodent-like dentition with enlarged incisors and molars and no canines.


D. brachydactylus


One specimen [38]

A docodontid specialized for a subterranean burrowing lifestyle. The skeletal structure and body proportions are strikingly similar to the golden mole. It was at least 9 centimeters long, exempting the tail, and weighed at least 9 grams, or perhaps 16 grams.


J. sinensis [39]


One specimen

A small, shrew-like very basal eutherian with a body length approximately 70-100 millimeters. The discovery of Juramaia provides new insight into the evolution of placental mammals by showing that their lineage diverged from that of the metatheres 35 million years earlier than previously thought. Furthermore, its discovery fills gaps in the fossil record and helps to calibrate modern, DNA-based methods of dating the evolution. Based on climbing adaptations found in the forelimb bones, it has been suggested that the basal stock of eutherians was arboreal.

Maiopatagium M. furculiferumA gliding euharamiyidan similar in lifestyle to a colugo.


M. mammaliaformis

Inner Mongolia

Daohugou bed

One specimen

An eleutherodontid haramiyidan thought to have been a herbivore that lived on the ground, having a similar posture to modern-day armadillos and rock hyraxes. [40] Megaconus is estimated to have weighed about 250 grams (8.8 ounces).

Microdocodon M. gracilisDaohugou bedA tegotheriid docodontan known from a specimen with a preserved hyoid bone, which is almost unknown in the early mammal fossil record.
Qishou Q. jizantang Euharamiyida


R. eurasiaticus


Daxishan site

One specimen

An omnivorous paulchoffatoid multituberculate that is the oldest so far described in the multituberculates. It strongly resembled a small rodent (like a rat or a chipmunk). [41] It is estimated to have weighed between 65 and 80 grams, about that of an average chipmunk.


V. antiquum

Inner Mongolia

Daohugou bed

One specimen

A gliding, flying squirrel-like volaticotherian eutriconodont with a specialized gliding membrane. The teeth of Volaticotherium were highly specialized for eating insects, and its limbs were adapted to living in trees. The gliding membrane was insulated by a thick covering of fur, and was supported by the limbs as well as the tail. The discovery of Volaticotherium provided the earliest-known record of a gliding mammal, and provided further evidence of mammalian diversity during the Mesozoic Era.

Vilevolodon V. diplomylosA gliding eleutherodontid haramiyidan with a herbivorous diet.
Xianshou X. linglong, X. songaeA gliding euharamiyidan


The following orders are represented in the formation; Ephemeroptera, Odonata, Plecoptera, Blattodea, Orthoptera, Hemiptera, Neuroptera, Coleoptera, Hymenoptera, and Diptera.

An indeterminate aeshnoid (insect) species is known from Liaoning. [28]

Color key
Taxon Reclassified taxonTaxon falsely reported as presentDubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.
Arthropods of the Daohugou Beds


A. neimengguensis [42]

Inner Mongolia

A tangle-veined fly

Mongolarachne Mongolarachne.jpg


A. striatus [43]

Archisargid flies

A. zhangi [44]

Inner Mongolia


A. spurivenius [43]

Archisargid flies

A. strigatus [43]


C. (Calosargus) antiquus [43]

Archisargid flies

C. (C.) bellus [43]

C. (C.) daohugouensis [43]

C. (C.) hani [43]

C. (C.) tenuicellulatus [43]

C. (C.) validus [43]

C. (Pterosargus) sinicus [43]

Inner Mongolia


D. impudica [28]


An ostracod

D. magna [28]


An ostracod

D. sarytirmenensis [28]


An ostracod


D. vulcanica [43]

A water boatman


F. gregarious [43]

A mayfly


E. gertschi [45]

1 Specimen

A plectreurid spider


H. liui [46]

A schizophorid flying water beetle


J. orientalis [43]

Inner Mongolia

A Nemestrinoid fly


M. explanatus [46]

Schizophorid flying water beetles

M. oxycerus [46]

M. grammicus [46]


M. daohugouensis [43]

An archisargid fly


M. portentosus [43]

Inner Mongolia

Archisargid flies

M. signatus [43]

Inner Mongolia


M. jurassica [47]

2 Specimens

An araneomorph spider originally thought to be a species of golden silk orb-weaver. From a female specimen, the carapace is 9.31 by 6.83 millimeters (0.367 by 0.269 inches) and the opisthosoma is 15.36 by 9.5 millimeters (0.605 by 0.374 inches). The total body length is approximately 24.6 millimeters (0.97 in) while the front legs reach about 56.5 millimeters (2.22 in) in length. A male specimen has a body length of 16.54 millimeters (0.651 in) with elongated pedipalps.


S. lacustris [43]

A mayfly


S. darani [46]

A schizophorid flying water beetle

Other invertebrates

GenusSpeciesProvinceStratigraphic PositionAbundanceNotes


S. cliovata [28]


A bivalve


Survey based on Wang et al. 2006 unless otherwise noted. [1]


Cycad-like plants, the most abundant plant group in the formation. 27 species in 11 genera.

Bennettitales of the Tiaojishan Formation
GenusSpeciesLocationStratigraphic positionAbundanceNotesImages












Prehistoric ginkgo trees, common, with 11 species present in 6 genera.

Ginkoales of the Tiaojishan Formation
GenusSpeciesLocationStratigraphic positionAbundanceNotesImages







Conifers, 5 species present in 4 genera.

Pinophytans of the Tiaojishan Formation
GenusSpeciesLocationStratigraphic positionAbundanceNotesImages







Leptosporangiate ferns, represented by 17 species in 8 genera, are the second most abundant plant type in the formation.

Pteridophytans of the Tiaojishan Formation
GenusSpeciesLocationStratigraphic positionAbundanceNotesImages
AshicaulisBeipiaoOne specimenA whole-plant osmundacean. Preserved stem 50 cm high and 35–41 cm across. Sterile fronds of Cladophlebis-type, fertile fronds of Todites-type with in-situ spores of Osmundacidites-type. [48]


Form genus of sterile fern fronds, typically assigned to Osmundaceae. A whole-plant osmundacean tree-fern with Cladophlebis fronds attached is known from this formation.

Claytosmunda C. chengii, C. liaoningensis, C. plumites, C. preosmunda, C. sinica, C. wangiiBeipiao, LiaoningNumerous specimensInterrupted ferns. Numerous fossil rhizomes previously assigned to Millerocaulis or Ashicaulis were interpreted to be close relatives and possible precursors of Claytosmunda claytoniana , the only extant representative of the genus. [49]


Dicksoniaceae Tree ferns.

Dicksonia D. changeyingziensis Dicksoniaceae Tree ferns.
D. charielsa
Eboracia Dicksoniaceae Tree ferns.


H. shebudaiensis


A dipterid fern.


M. hoerenensis


A marattiopsid fern.


R. stricta

A fern.


T. denticulataFertile fronds of osmundacean ferns that resemble Todea. Known to attach to "Ashicaulis"-type stems with sterile Cladophlebis-type fronds in this formation.
T. williamsoniiIsolated fertile fronds of osmundacean ferns resembling Todea.

Other plants

Cycads, fairly diverse, with 10 species present in 2 genera.

Cycads of the Tiaojishan Formation
GenusSpeciesLocationStratigraphic positionAbundanceNotesImages










H. shebudaiensis


A bryophyte.

Taeniopteris sp.


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The Haifanggou Formation, also known as the Jiulongshan Formation, is a fossil-bearing rock deposit located near Daohugou village of Ningcheng County, in Inner Mongolia, northeastern China.

<span class="mw-page-title-main">Rhamphorhynchidae</span> Family of breviquartossan pterosaurs from the Jurassic period

Rhamphorhynchidae is a group of early pterosaurs named after Rhamphorhynchus, that lived in the Late Jurassic. The family Rhamphorhynchidae was named in 1870 by Harry Govier Seeley. Members of the group possess no more than 11 pairs of teeth in the rostrum, a deltopectoral crest that is constricted at the base but expanded at the distal end, and a bent phalange on the fifth toe.

<i>Kunpengopterus</i> Genus of wukongopterid pterosaur from the Jurassic period

Kunpengopterus is a genus of wukongopterid pterosaur from the middle-late Jurassic Tiaojishan Formation of northeastern China. The genus contains two species, the type species K. sinensis and K. antipollicatus.

<span class="mw-page-title-main">Wukongopteridae</span> Family of darwinopteran pterosaurs from the Jurassic period

Wukongopteridae is a group of basal pterosaurs, found in China and the UK. It contains eight species in five genera, all dated to the Middle to Late Jurassic period,

Archaeoistiodactylus is an extinct genus of wukongopterid pterosaur from the Middle Jurassic of China.

<i>Chuanqilong</i> Extinct genus of dinosaurs

Chuanqilong is a monospecific genus of basal ankylosaurid dinosaur from the Liaoning Province, China that lived during the Early Cretaceous in what is now the Jiufotang Formation. The type and only species, Chuanqilong chaoyangensis, is known from a nearly complete skeleton with a skull of a juvenile individual. It was described in 2014 by Fenglu Han, Wenjie Zheng, Dongyu Hu, Xing Xu, and Paul M. Barrett. Chuanqilong shows many similarities with Liaoningosaurus and may represent a later ontogenetic stage of the taxon.

Daohugoupterus is a genus of pterosaur from the Middle to Late Jurassic Daohugou Beds of the Tiaojishan Formation in Inner Mongolia, China.

<i>Liaodactylus</i> Genus of ctenochasmatid pterosaur from the Late Jurassic

Liaodactylus is a genus of filter-feeding ctenochasmatid pterosaur from the Jurassic of China. The genus contains one species, L. primus, described by Zhou et al. in 2017. As an adaptation to filter-feeding, Liaodactylus had approximately 150 long, comb-like teeth packed closely together. It is both the earliest known ctenochasmatid and the first filter-feeding pterosaur from the Jurassic Tiaojishan Formation. Later and more specialized ctenochasmatids differ from Liaodactylus in having longer snouts, smaller openings in the skull, and more teeth. Within the Ctenochasmatidae, Liaodactylus was most closely related to the European Ctenochasma.

<i>Douzhanopterus</i> Genus of monofenestratan pterosaur from the Late Jurassic

Douzhanopterus is an extinct genus of monofenestratan pterosaur from the Late Jurassic of Liaoning, China. It contains a single species, D. zhengi, named by Wang et al. in 2017. In many respects, it represents a transitional form between basal pterosaurs and the more specialized pterodactyloids; for instance, its tail is intermediate in length, still being about twice the length of the femur but relatively shorter compared to that of the more basal Wukongopteridae. Other intermediate traits include the relative lengths of the neck vertebrae and the retention of two, albeit reduced, phalanx bones in the fifth digit of the foot. Phylogenetically, Douzhanopterus is nested between the wukongopterids and a juvenile pterosaur specimen from Germany known as the "Painten pro-pterodactyloid", which is similar to Douzhanopterus in many respects but approaches pterodactyloids more closely elsewhere.


  1. 1 2 3 Wang, Y.; Ken, S.; Zhang, W.; Zheng, S. (2006). "Biodiversity and palaeoclimate of the Middle Jurassic floras from the Tiaojishan Formation in western Liaoning, China". Progress in Natural Science. 16 (1): 222–230. doi:10.1080/10020070612330087.
  2. Xiaolin Wang; Zhonghe Zhou; Huaiyu He; Fan Jin; Yuanqing Wang; Jiangyong Zhang; Yuan Wang; Xing Xu; Fucheng Zhang (2005). "Stratigraphy and age of the Daohugou Bed in Ningcheng, Inner Mongolia". Chinese Science Bulletin. 50 (20): 2369–2376. Bibcode:2005ChSBu..50.2369W. doi:10.1007/BF03183749. S2CID   198142479.
  3. Liu, Y.; Ji, S.; Yang, Z. (2006). "U-Pb zircon age for the Daohugou Biota at Ningcheng of Inner Mongolia and comments on related issues". Chinese Science Bulletin. 51 (21): 2634–2644. Bibcode:2006ChSBu..51.2634L. doi:10.1007/s11434-006-2165-2. S2CID   96442710.
  4. Zhang, H.; Wang, M.; Liu, X. (2008). "Constraints on the upper boundary age of the Tiaojishan Formation volcanic rocks in West Liaoning-North Hebei by LA-ICP-MS dating". Chinese Science Bulletin. 53 (22): 3574–3584. Bibcode:2008SciBu..53.3574Z. doi: 10.1007/s11434-008-0287-4 .
  5. Notably Mongolarachne jurassica .
  6. Tan, Jingjing; Ren, Dong; Shih, Chungkun (2006). "New Cupedids from the Middle Jurassic of Inner Mongolia, China (Coleoptera: Archostemata)". Annales Zoologici. 56 (1): 1–6.
  7. Zhang, Kuiyan; Yang, Ding; Ren, Dong (2006). "The first snipe fly (Diptera: Rhagionidae) from the Middle Jurassic of Inner Mongolia, China". Zootaxa. 1134: 51–57. doi:10.11646/zootaxa.1134.1.3. S2CID   83904231.
  8. Gao, K. -Q.; Shubin, N. H. (2012). "Late Jurassic salamandroid from western Liaoning, China". Proceedings of the National Academy of Sciences. 109 (15): 5767–72. Bibcode:2012PNAS..109.5767G. doi: 10.1073/pnas.1009828109 . PMC   3326464 . PMID   22411790.
  9. Lü Junchang and Fucha Xiaohui (2011). "A new pterosaur (Pterosauria) from Middle Jurassic Tiaojishan Formation of western Liaoning, China". Global Geology Z1: 113–118.
  10. Yang, Z.; Benton, M. J.; Hone, D. W.; Xu, X.; McNamara, M.E.; Jiang, B. (2022). "Allometric analysis sheds light on the systematics and ontogeny of anurognathid pterosaurs". Journal of Vertebrate Paleontology. 41 (5): e2028796. doi:10.1080/02724634.2021.2028796. hdl: 10468/12968 . S2CID   247262846.
  11. 1 2 Wei, X.; Pêgas, R. V.; Shen, C.; Guo, Y.; Ma, W.; Sun, D.; Zhou, X. (2021). "Sinomacrops bondei, a new anurognathid pterosaur from the Jurassic of China and comments on the group". PeerJ. 9: e11161. doi: 10.7717/peerj.11161 . PMC   8019321 . PMID   33850665.
  12. Lü, J. (2009). "A new non-pterodactyloid pterosaur from Qinglong County, Hebei Province of China". Acta Geologica Sinica (English Edition). 83 (2): 189–199. doi:10.1111/j.1755-6724.2009.00062.x. S2CID   247739978.
  13. Cheng, Xin; Wang, Xiaolin; Jiang, Shunxing; Kellner, Alexander W.A. (2014). "Short note on a non-pterodactyloid pterosaur from Upper Jurassic deposits of Inner Mongolia, China". Historical Biology: An International Journal of Paleobiology. 27 (6): 1–6. doi:10.1080/08912963.2014.974038. S2CID   128454476.
  14. Lü, J.; Unwin, D.M.; Jin, X.; Liu, Y.; Ji, Q. (2010). "Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull". Proceedings of the Royal Society B. 277 (1680): 383–389. doi:10.1098/rspb.2009.1603. PMC   2842655 . PMID   19828548.
  15. 1 2 Lü Junchang & David W.E. Hone (2012). "A New Chinese Anurognathid Pterosaur and the Evolution of Pterosaurian Tail Lengths". Acta Geologica Sinica. 86 (6): 1317–1325. doi:10.1111/1755-6724.12002. S2CID   140741278.
  16. Wang, X.; Jiang, S.; Zhang, J.; Cheng, X.; Yu, X.; Li, Y.; Wei, G.; Wang, X. (2017). "New evidence from China for the nature of the pterosaur evolutionary transition". Scientific Reports. 7: 42763. Bibcode:2017NatSR...742763W. doi:10.1038/srep42763. PMC   5311862 . PMID   28202936.
  17. Lu, J.; Fucha, X.; Chen, J. (2010). "A new scaphognathine pterosaur from the Middle Jurassic of western Liaoning, China". Acta Geoscientica Sinica. 31 (2): 263–266.
  18. 1 2 Wang, X.; Zhou, Z. (2006). "Pterosaur assemblages of the Jehol Biota and their implication for the Early Cretaceous pterosaur radiation". Geological Journal. 41 (3–4): 405–418. doi:10.1002/gj.1046. S2CID   131602205.
  19. Lü, J.; Ji, Q. (2006). "Preliminary results of a phylogenetic analysis of the pterosaurs from western Liaoning and surrounding areas". J. Paleont. Soc. Korea. 22 (1): 239–261.
  20. Cheng, Xin; Wang, Xiaolin; Jiang, Shunxing; Kellner, Alexander W.A. (2012). "A new scaphognathid pterosaur from western Liaoning, China". Historical Biology. 24: 101–111. doi:10.1080/08912963.2011.635423. S2CID   128539358.
  21. Lü Junchang & Bo Xue (2011). "A New Rhamphorhynchid Pterosaur (Pterosauria) from the Middle Jurassic Tiaojishan Formation of Western Liaoning, China". Acta Geologica Sinica. 85 (5): 977–983. doi:10.1111/j.1755-6724.2011.00531.x. S2CID   140196879.
  22. Wang, Xiaolin; Kellner, Alexander W.A.; Jiang, Shunxing; Cheng, Xin; Meng, Xi & Rodrigues, Taissa (2010). "New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China" (PDF). Anais da Academia Brasileira de Ciências. 82 (4): 1045–1062. doi:10.1590/S0001-37652010000400024. PMID   21152776.
  23. Zhou, X.; Pêgas, R. V.; Ma, W.; Han, G.; Jin, X.; Leal, M. E. C.; Bonde, N.; Kobayashi, Y.; Lautenschlager, S.; Wei, X.; Shen, C.; Ji, S. (2021). "A new darwinopteran pterosaur reveals arborealism and an opposed thumb". Current Biology . 31 (11): 2429–2436.e7. doi: 10.1016/j.cub.2021.03.030 . PMID   33848460. S2CID   233215450.
  24. Zhou, C.-F.; Gao, K.-Q.; Yi, H.; Xue, J.; Li, Q.; Fox, R.C. (2017). "Earliest filter-feeding pterosaur from the Jurassic of China and ecological evolution of Pterodactyloidea". Royal Society Open Science. 4 (2): 160672. Bibcode:2017RSOS....460672Z. doi:10.1098/rsos.160672. PMC   5367317 . PMID   28386425.
  25. Hone, D.W.E. (2020). "A review of the taxonomy and palaeoecology of the Anurognathidae (Reptilia, Pterosauria)". Acta Geologica Sinica - English Edition. 94 (5): 1676–1692. doi:10.1111/1755-6724.14585. S2CID   225169094.
  26. Lü J, Unwin DM, Zhao B, Gao C, Shen C (2012). "A new rhamphorhynchid (Pterosauria: Rhamphorhynchidae) from the Middle/Upper Jurassic of Qinglong, Hebei Province, China" (PDF). Zootaxa. 3158: 1–19. doi:10.11646/zootaxa.3158.1.1.
  27. Wang, X.; Kellner, A.W.A.; Jiang, S.; Meng, X. (2009). "An unusual long-tailed pterosaur with elongated neck from western Liaoning of China". Annals of the Brazilian Academy of Sciences. 81 (4): 793–812. doi: 10.1590/s0001-37652009000400016 . PMID   19893903.
  28. 1 2 3 4 5 6 Hu, D.; Hou, L.; Zhang, L. & Xu, X. (2009). "A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus". Nature. 461 (7264): 640–643. Bibcode:2009Natur.461..640H. doi:10.1038/nature08322. PMID   19794491. S2CID   205218015.
  29. Li Q, Gao KQ, Vinther J, et al. (March 2010). "Plumage color patterns of an extinct dinosaur" (PDF). Science. 327 (5971): 1369–72. Bibcode:2010Sci...327.1369L. doi:10.1126/science.1186290. PMID   20133521. S2CID   206525132.
  30. Hu, Dongyu; Clarke, Julia A.; Eliason, Chad M.; Qiu, Rui; Li, Quanguo; Shawkey, Matthew D.; Zhao, Cuilin; D'Alba, Liliana; Jiang, Jinkai; Xu, Xing (2018). "A bony-crested Jurassic dinosaur with evidence of iridescent plumage highlights complexity in early paravian evolution". Nature Communications. 9 (1): 217. Bibcode:2018NatCo...9..217H. doi:10.1038/s41467-017-02515-y. PMC   5768872 . PMID   29335537.
  31. Wang, Y; et al. (2006). "Biodiversity and palaeoclimate of the Middle Jurassic floras from the Tiaojishan Formation in western Liaoning, China". Progress in Natural Science. 16 (S1): 222–30. doi:10.1080/10020070612330087A (inactive 31 January 2024).{{cite journal}}: CS1 maint: DOI inactive as of January 2024 (link)
  32. Lefèvre, Ulysse; Cau, Andrea; Cincotta, Aude; Hu, Dongyu; Chinsamy, Anusuya; Escuillié, François; Godefroit, Pascal (2017). "A new Jurassic theropod from China documents a transitional step in the macrostructure of feathers". The Science of Nature. 104 (9–10): 74. Bibcode:2017SciNa.104...74L. doi:10.1007/s00114-017-1496-y. PMID   28831510. S2CID   32780661.
  33. Xu, Xing; You, Hailu; Du, Kai; Han, Fenglu (28 July 2011). "An Archaeopteryx-like theropod from China and the origin of Avialae". Nature. 475 (7357): 465–470. doi:10.1038/nature10288. PMID   21796204. S2CID   205225790.
  34. Xu, X.; Zheng, X.; Sullivan, C.; Wang, X.; Xing, L.; Wang, Y.; Zhang, X.; o’Connor, J. K.; Zhang, F.; Pan, Y. (2015). "A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings". Nature. 521 (7550): 70–3. Bibcode:2015Natur.521...70X. doi:10.1038/nature14423. PMID   25924069. S2CID   205243599.
  35. 1 2 Evans, S.E.; Wang, Y. (2010). "A new lizard (Reptilia: Squamata) with exquisite preservation of soft tissue from the Lower Cretaceous of Inner Mongolia, China". Journal of Systematic Palaeontology. 8 (1): 81–95. doi:10.1080/14772010903537773. S2CID   56046960.
  36. Meng, Qing-Jin; Ji, Qiang; Zhang, Yu-Guang; Liu, Di; Grossnickle, David M.; Luo, Zhe-Xi (2015). "An arboreal docodont from the Jurassic and mammaliaform ecological diversification". Science. 347 (6223): 764–768. Bibcode:2015Sci...347..764M. doi:10.1126/science.1260879. PMID   25678661. S2CID   206562565.
  37. Zheng, X.; Bi, S.; Wang, X.; Meng, J. (2013). "A new arboreal haramiyid shows the diversity of crown mammals in the Jurassic period". Nature. 500 (7461): 199–202. Bibcode:2013Natur.500..199Z. doi:10.1038/nature12353. PMID   23925244. S2CID   2164378.
  38. Zhe-Xi Luo; Qing-Jin Meng; Qiang Ji; Di Liu; Yu-Guang Zhang; April I. Neander (2015). "Evolutionary development in basal mammaliaforms as revealed by a docodontan". Science. 347 (6223): 760–764. Bibcode:2015Sci...347..760L. doi:10.1126/science.1260880. PMID   25678660. S2CID   206562572.
  39. Luo, Z.; Yuan, C.; Meng, Q.; Ji, Q. (2011). "Jurassic eutherian mammal and divergence of marsupials and placentals". Nature. 476 (7361): 442–445. Bibcode:2011Natur.476..442L. doi:10.1038/nature10291. PMID   21866158. S2CID   205225806.
  40. Zhou, C. F.; Wu, S.; Martin, T.; Luo, Z. X. (2013). "A Jurassic mammaliaform and the earliest mammalian evolutionary adaptations". Nature. 500 (7461): 163–167. Bibcode:2013Natur.500..163Z. doi:10.1038/nature12429. PMID   23925238. S2CID   4346751.
  41. Yuan, C. -X.; Ji, Q.; Meng, Q. -J.; Tabrum, A. R.; Luo, Z. -X. (2013). "Earliest Evolution of Multituberculate Mammals Revealed by a New Jurassic Fossil". Science. 341 (6147): 779–783. Bibcode:2013Sci...341..779Y. doi:10.1126/science.1237970. PMID   23950536. S2CID   25885140.
  42. Zhang, K.; Yang, D.; Ren, D.; Ge, F. (2008). "New Middle Jurassic tangle−veined flies from Inner Mongolia, China". Acta Palaeontologica Polonica. 53 (1): 161–164. doi: 10.4202/app.2008.0112 .
  43. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Zhang, J.F. (2010). "Records of bizarre Jurassic brachycerans in the Daohugou biota, China (Diptera, Brachycera, Archisargidae and Rhagionemestriidae)". Palaeontology. 53 (2): 307–317. Bibcode:2010Palgy..53..307Z. doi: 10.1111/j.1475-4983.2010.00934.x .
  44. Zhang, K.; Li; Yang, D.; Ren, D. (2009). "A new species of Archirhagio Rohdendorf, 1938 from the Middle Jurassic of Inner Mongolia of China (Diptera: Archisargidae)". Zootaxa. 1984: 61–65. doi:10.11646/zootaxa.1984.1.4. S2CID   82881259.
  45. Selden, P.A.; Huang, D. (2010). "The oldest haplogyne spider (Araneae: Plectreuridae), from the Middle Jurassic of China". Naturwissenschaften. 97 (5): 449–59. Bibcode:2010NW.....97..449S. doi:10.1007/s00114-010-0649-z. PMID   20140419. S2CID   24576304.
  46. 1 2 3 4 5 Jarzembowski, E. A.; Yan, E. V.; Wang, B.; Zhang, H. C. (2012). "A new flying water beetle (Coleoptera: Schizophoridae) from the Jurassic Daohugou lagerstätte". Palaeoworld . 21 (3–4): 160–166. doi:10.1016/j.palwor.2012.09.002.
  47. Selden, P. A.; Shih, C.; Ren, D. (2011). "A golden orb-weaver spider (Araneae: Nephilidae: Nephila) from the Middle Jurassic of China". Biology Letters. 7 (5): 775–778. doi:10.1098/rsbl.2011.0228. PMC   3169061 . PMID   21508021.
  48. XiaoJu Yang; Wu Zhang; ShaoLin Zheng (2010). "An osmundaceous rhizome with sterile and fertile fronds and in situ spores from the Jurassic of western Liaoning". Chinese Science Bulletin. 55 (34): 3864–3867. Bibcode:2010ChSBu..55.3864Y. doi:10.1007/s11434-010-4191-3. S2CID   85810468.
  49. Benjamin Bomfleur; Guido W. Grimm; Stephen McLoughlin (2017). "The fossil Osmundales (Royal Ferns)—a phylogenetic network analysis, revised taxonomy, and evolutionary classification of anatomically preserved trunks and rhizomes". PeerJ. 5: e3433. doi: 10.7717/peerj.3433 . PMC   5508817 . PMID   28713650.