Tiaojishan Formation
| Tiaojishan Formation | ||
|---|---|---|
Ma | ||
Approximate paleocoordinates 43°00′N 123°06′E / 43.0°N 123.1°E | | |
| Region | Hebei, Inner Mongolia, & Liaoning | |
| Country |  China | |
| Extent | Yanshan Belt | |
  Tiaojishan Formation (China)  Tiaojishan Formation (Liaoning) | ||
The Tiaojishan Formation is a
Age
Using
Climate
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
Fauna
Beautifully preserved fossils of
The forests of the Yanliao biota grew in a humid, warm - temperate climate and were dominated by
Salamanders
| Salamanders of the Tiaojishan Formation | ||||||
|---|---|---|---|---|---|---|
| Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
|
B. jianpingensis[8] |
Liaoning |
A salamandroid known from an almost complete and articulated skeleton exposed in ventral view. |
 | |||
|
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
| Pterosaurs of the Tiaojishan Formation | ||||||
|---|---|---|---|---|---|---|
| Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
|
A. linglongtaensis[9] |
Liaoning |
A monofenestratan known from an incomplete skeleton with a partial skull and lower jaw. |
Darwinopterus modularis  Kunpengopterus antipollicatus   | |||
|
C. rong[10] |
Daohugou bed[11] |
One specimen |
A long-tailed anurognathid known from a complete skeleton of a juvenile with extensive preservation of pycnofibres and wing membranes.
| |||
|
C. pani[12] |
Hebei |
A pterodactyliform known only from a single specimen of a young juvenile, measuring 475 millimeters (18.7 in) in wingspan.
| ||||
|
D. delicatus[13] |
One specimen |
A relatively basal pterosaur known from a partial skeleton with soft tissue impressions. | ||||
|
D. modularis[14] |
Liaoning |
A 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 |
||||||
|
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] |
Linglongta |
One specimen |
A non-pterodactyloid monofenestratan with a proportionately short tail. | |||
|
F. lii[17] |
Liaoning |
A 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 |
A batrachognathine anurognathid preserved with pycnofibres and skin remains. | ||||
|
J. robustus[20] |
Liaoning |
A scaphognathine rhamphorhynchid known from a single fossil skeleton. | ||||
|
J. zhaoianus[21] |
Liaoning |
A scaphognathine rhamphorhynchid known from a nearly complete skeleton with the skull preserved. | ||||
|
K. sinensis[22] |
Liaoning |
Daohugou bed |
Three specimens |
A wukongopterid with an elongated head, 106.9 millimeters long, and an opposed thumb. | ||
|
K. antipollicatus[23] |
Liaoning |
Two nearly complete specimens | ||||
|
L. primus[24] |
Daxishan (Linglongta) |
One specimen |
A ctenochasmatid pterosaur with elongated, comb-like teeth. | |||
|
Hebei |
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] |
Liaoning |
A rhamphorhynchine rhamphorhynchid known from only one specimen that includes a skeleton with a skull.
| ||||
|
S. bondei[11] |
Hebei |
Daohugou bed |
One specimen |
A long-tailed batrachognathine anurognathid known from a relatively complete skull and skeleton with soft tissue patches. | ||
|
W. lii |
Liaoning |
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
| Dinosaurs of the Tiaojishan Formation | ||||||
|---|---|---|---|---|---|---|
| Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
|
A. huxleyi[28] |
Several specimens[29] |
An 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).
|
          | |||
|
A. xui |
Liaoning |
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 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 |
Liaoning |
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 |
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 |
Liaoning |
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 |
Liaoning |
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 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 |
Liaoning |
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] |
Liaoning |
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. | ||
Lizards
Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Lepidosaurs (lizards and relatives) of the Daohugou Beds | |||||
|---|---|---|---|---|---|
| Genus | Species | State | Abundance | Notes | Images |
|
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
| Cynodonts of the Tiaojishan Formation | ||||||
|---|---|---|---|---|---|---|
| Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
|
A. scansorius |
Daohugou bed |
A mammals . The front teeth were spade-like, indicating that Agilodocodon could gnaw tree bark and consume gum or sap.
|
     | |||
|
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] |
Liaoning |
One specimen |
A small, shrew-like very basal 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. furculiferum | A 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. gracilis | Daohugou bed | A 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 |
Liaoning |
Daxishan site |
One specimen |
An omnivorous 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, Mesozoic Era .
| ||
| Vilevolodon | V. diplomylos | A gliding eleutherodontid haramiyidan with a herbivorous diet. | ||||
| Xianshou | X. linglong, X. songae | A gliding euharamiyidan | ||||
Arthropods
The following orders are represented in the formation;
An indeterminate aeshnoid (insect) species is known from Liaoning.[28]
Color key
|
Notes Uncertain or tentative taxa are in small text; |
| Arthropods of the Daohugou Beds | ||||||
|---|---|---|---|---|---|---|
| Genus | Species | State | Abundance | Notes | Images | |
|
A. neimengguensis[42] |
 | |||||
|
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] |
Liaoning |
An ostracod |
||||
|
D. magna[28] |
Liaoning |
An ostracod | ||||
|
D. sarytirmenensis[28] |
Liaoning |
An ostracod | ||||
|
D. vulcanica[43] |
||||||
|
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 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
| Genus | Species | Province | Stratigraphic Position | Abundance | Notes |
|---|---|---|---|---|---|
|
S. cliovata[28] |
Liaoning |
A bivalve
|
Flora
Survey based on Wang et al. 2006 unless otherwise noted.[1]
Bennettitales
Cycad-like plants, the most abundant plant group in the formation. 27 species in 11 genera.
| Bennettitales of the Tiaojishan Formation | ||||||
|---|---|---|---|---|---|---|
| Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
Ginkgoales
Prehistoric ginkgo trees, common, with 11 species present in 6 genera.
| Ginkoales of the Tiaojishan Formation | ||||||
|---|---|---|---|---|---|---|
| Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
Pinophyta
Conifers, 5 species present in 4 genera.
Pinophytans of the Tiaojishan Formation
| ||||||
|---|---|---|---|---|---|---|
| Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
Pteridophyta
Pteridophytans of the Tiaojishan Formation
| ||||||
|---|---|---|---|---|---|---|
| Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
| Ashicaulis | Beipiao | One specimen | A 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. wangii | Beipiao, Liaoning | Numerous specimens | Interrupted 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]
| ||
| Dicksonia | D. changeyingziensis | Dicksoniaceae Tree ferns. | ||||
| D. charielsa | ||||||
| Eboracia | Dicksoniaceae Tree ferns. | |||||
|
H. shebudaiensis |
Uncommon. |
|||||
|
M. hoerenensis |
Uncommon. |
A marattiopsid fern.
| ||||
|
R. stricta |
A fern. | |||||
| T. denticulata | Fertile fronds of osmundacean ferns that resemble Todea. Known to attach to "Ashicaulis"-type stems with sterile Cladophlebis-type fronds in this formation. | |||||
| T. williamsonii | Isolated fertile fronds of osmundacean ferns resembling Todea. | |||||
Other plants
Cycads, fairly diverse, with 10 species present in 2 genera.
| Cycads of the Tiaojishan Formation | ||||||
|---|---|---|---|---|---|---|
| Genus | Species | Location | Stratigraphic position | Abundance | Notes | Images |
|
Horsetails .
| ||||||
|
Horsetails .
| ||||||
|
H. shebudaiensis |
Uncommon. |
A bryophyte. | ||||
|
Taeniopteris sp. |
Uncommon. |
|||||
References
- ^ .
- S2CID 198142479.
- S2CID 96442710.
- .
- ^ Notably Mongolarachne jurassica.
- ^ 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.
- S2CID 83904231.
- PMID 22411790.
- ^ Lü Junchang and Fucha Xiaohui (2011). "A new pterosaur (Pterosauria) from Middle Jurassic Tiaojishan Formation of western Liaoning, China". Global Geology Z1: 113–118.
- S2CID 247262846.
- ^ PMID 33850665.
- S2CID 247739978.
- S2CID 128454476.
- PMID 19828548.
- ^ S2CID 140741278.
- PMID 28202936.
- ^ 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.
- ^ S2CID 131602205.
- ^ 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.
- S2CID 128539358.
- S2CID 140196879.
- PMID 21152776.
- S2CID 233215450.
- PMID 28386425.
- S2CID 225169094.
- .
- PMID 19893903.
- ^ S2CID 205218015.
- S2CID 206525132.
- PMID 29335537.
- doi:10.1080/10020070612330087A (inactive 31 January 2024).)
{{cite journal}}: CS1 maint: DOI inactive as of January 2024 (link - S2CID 32780661.
- S2CID 205225790.
- S2CID 205243599.
- ^ S2CID 56046960.
- S2CID 206562565.
- S2CID 2164378.
- S2CID 206562572.
- S2CID 205225806.
- S2CID 4346751.
- S2CID 25885140.
- .
- ^ .
- S2CID 82881259.
- S2CID 24576304.
- ^ .
- PMID 21508021.
- S2CID 85810468.
- PMID 28713650.
