Tianyulong

Tianyulong (Chinese: 天宇龍; Pinyin: tiānyǔlóng; named for the Shandong Tianyu Museum of Nature where the holotype fossil is housed) was a genus of heterodontosaurid ornithischian dinosaur. The only species was T. confuciusi, whose remains were discovered in Jianchang County, Western Liaoning Province, China.[1]

Tianyulong
Temporal range: Late Jurassic, 158.5 Ma
Tianyulong
Specimen IVPP V17090, muzzle, hand, feet and tail framed in red
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Ornithischia
Family: Heterodontosauridae
Genus: Tianyulong
Zheng et al., 2009
Species:
T. confuciusi
Binomial name
Tianyulong confuciusi
Zheng et al., 2009

History

The holotype of Tianyulong, STMN 26-3 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 at least 158.5 million years ago.[2]

Another specimen, IVPP V17090, was described in 2012. At least four other specimens remain undescribed.[3]

Description

Tianyulong holotype
The holotype skeleton, STMN 26-3

STMN 26-3 consists of an incomplete skeleton preserving a partial skull and mandible, partial presacral vertebrae, proximal–middle caudal vertebrae, nearly complete right scapula, both humeri, the proximal end of the left ulna, partial pubes, both ischia, both femora, the right tibia and fibula and pes, as well as remains of long, singular and unbranched filamentous integumentary structures. The holotype is from a subadult individual that probably measured 70 cm in length based on the proportions of the related South African species Heterodontosaurus tucki. However, Tianyulong had unusual proportions compared to other heterodontosaurids. The head was large and the legs and tail were long, but the neck and forelimbs were short.[3]

Tianyulong has a row of long, filamentous integumentary structures on the back, tail and neck of the specimen. The similarity of these structures with those found on some theropods suggests their homology with feathers and raises the possibility that the earliest dinosaurs and their ancestors were covered with homologous dermal filamentous structures that can be considered primitive feathers ("proto-feathers").

Classification

Heterodontosaurus and Tianyulong
Skull diagrams of Heterodontosaurus and Tianyulong

Tianyulong is classified as a heterodontosaurid, a group of small ornithischian dinosaur characterized by a slender body, long tail and a pair of enlarged canine-like tusks. They were herbivorous or possibly omnivorous. Until the discovery of Tianyulong, known members of the group were restricted to the Early Jurassic of South Africa, with one genus (Fruitadens) from the Late Jurassic of the USA, and possibly one additional genus (Echinodon) from the Early Cretaceous of England.

The cladogram below follows the analysis by Butler et al., 2011:[4]

Heterodontosauridae 

Echinodon

Abrictosaurus

NHM RU A100

Heterodontosaurus

Lycorhinus

Fruitadens

Tianyulong

Paleobiology

Tianyulong BW
Restoration

The filamentous integumentary structures are preserved on three areas of the fossil: in one patch just below the neck, another one on the back, and the largest one above the tail. The hollow filaments are parallel to each other and are singular with no evidence of branching. They also appear to be relatively rigid, making them more analogous to the integumentary structures found on the tail of Psittacosaurus[5] than to the proto-feather structures found in avian and non-avian theropods. Among the theropods, the structures in Tianyulong are most similar to the singular unbranched proto-feathers of Sinosauropteryx[6] and Beipiaosaurus.[7] The estimated length of the integumentary structures on the tail is about 60 mm which is seven times the height of a caudal vertebra. Their length and hollow nature argue against of them being subdermal structures such as collagen fibers.

Tianyulong confuciusi
Restored skeleton

Such dermal structures have previously been reported only in derived theropods and ornithischians, and their discovery in Tianyulong extends the existence of such structures further down in the phylogenetic tree. However, the homology between the ornithischian filaments and the theropods' proto-feathers is not obvious. If the homology is supported, the consequence is that the common ancestor of both saurischians and ornithischians were covered by feather-like structures, and that groups for which skin impression are known such as the sauropods were only secondarily featherless. If the homology is not supported, it would indicate that these filamentous dermal structures evolved independently in saurischians and ornithischians, as well as in other archosaurs such as the pterosaurs. The authors (in supplementary information to their primary article) noted that discovery of similar filamentous structures in the theropod Beipiaosaurus bolstered the idea that the structures on Tianyulong are homologous with feathers. Both the filaments of Tianyulong and the filaments of Beipiaosaurus were long, singular, and unbranched. In Beipiaosaurus, however, the filaments were flattened. In Tianyulong, the filaments were round in cross section, and therefore closer in structure to the earliest forms of feathers predicted by developmental models.[1]

References

  1. ^ a b Zheng, Xiao-Ting; You, Hai-Lu; Xu, Xing; Dong, Zhi-Ming (19 March 2009). "An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures". Nature. 458 (7236): 333–336. doi:10.1038/nature07856. PMID 19295609.
  2. ^ Liu Y.-Q. Kuang H.-W., Jiang X.-J., Peng N., Xu H. & Sun H.-Y. (2012). "Timing of the earliest known feathered dinosaurs and transitional pterosaurs older than the Jehol Biota." Palaeogeography, Palaeoclimatology, Palaeoecology (advance online publication).
  3. ^ a b Sereno, Paul (2012-03-10). "Taxonomy, morphology, masticatory function and phylogeny of heterodontosaurid dinosaurs". ZooKeys. 226: 1–225. doi:10.3897/zookeys.226.2840. ISSN 1313-2970. PMC 3491919.
  4. ^ Richard J. Butler, Jin Liyong, Chen Jun, Pascal Godefroit (2011). "The postcranial osteology and phylogenetic position of the small ornithischian dinosaur Changchunsaurus parvus from the Quantou Formation (Cretaceous: Aptian–Cenomanian) of Jilin Province, north-eastern China". Palaeontology. 54 (3): 667–683. doi:10.1111/j.1475-4983.2011.01046.x.CS1 maint: Multiple names: authors list (link)
  5. ^ Mayr, Gerald; Peters, D. Stephan; Plodowski, Gerhard; Vogel, Olaf (August 2002). "Bristle-like integumentary structures at the tail of the horned dinosaur Psittacosaurus". Naturwissenschaften. Heidelberg: Springer Berlin. 89 (8): 361–365. doi:10.1007/s00114-002-0339-6. PMID 12435037.
  6. ^ Currie, Philip J.; Pei-ji Chen (2001). "Anatomy of Sinosauropteryx prima from Liaoning, northeastern China". Canadian Journal of Earth Sciences. NRC Canada. 38 (12): 1705–1727. doi:10.1139/cjes-38-12-1705.
  7. ^ Xu, Xing; Zheng Xiao-ting; You, Hai-lu (20 January 2009). "A new feather type in a nonavian theropod and the early evolution of feathers". Proceedings of the National Academy of Sciences. 106 (3): 832–4. doi:10.1073/pnas.0810055106. PMC 2630069. PMID 19139401.

External links

Averostra

Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs that have a promaxillary fenestra (fenestra promaxillaris), an extra opening in the front outer side of the maxilla, the bone that makes up the upper jaw. Two groups of averostrans, the Ceratosauria and the Orionides, survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians and two groups of orionideans within the clade Coelurosauria, the Tyrannosauroidea and Maniraptoriformes, were still extant. Only one subgroup of maniraptoriformes, Aves, survived the extinction event and persisted to the present day.

Avetheropoda

Avetheropoda, or "bird theropods", is a clade that includes carnosaurians and coelurosaurs to the exclusion of other dinosaurs.

Cerapoda

Cerapoda ("ceratopsians and ornithopods") is a clade of the dinosaur order Ornithischia.

Dinosauriformes

Dinosauriformes is a clade of archosaurian reptiles that include the dinosaurs and their most immediate relatives. All dinosauriformes are distinguished by several features, such as shortened forelimbs and a partially to fully perforated acetabulum, the hole in the hip socket traditionally used to define dinosaurs. The oldest known member is Asilisaurus, dating to about 245 million years ago in the Anisian age of the middle Triassic period.

Echinodon

Echinodon (pronounced eh-KY-no-don) meaning "hedgehog tooth" in reference to the spines on its teeth (Ancient Greek: εχινος, romanized: echinos, lit. 'hedgehog', + ὀδών, odṓn, 'tooth'), occasionally known as Saurechinodon, is a genus of small European dinosaur of the early Cretaceous Period (Berriasian age), 140 million years ago.

Fruitadens

Fruitadens is a genus of heterodontosaurid dinosaur. The name means "Fruita tooth", in reference to Fruita, Colorado (USA), where its fossils were first found. It is known from partial skulls and skeletons from at least four individuals of differing biological ages, found in Tithonian (Late Jurassic) rocks of the Morrison Formation in Colorado. Fruitadens is the smallest known ornithischian dinosaur, with young adults estimated at 65 to 75 cm (26 to 30 in) in length and 0.5 to 0.75 kg (1.1 to 1.7 lb) in weight. It is interpreted as an omnivore and represents one of the latest-surviving heterodontosaurids.

Haya griva

Haya is an extinct genus of basal neornithischian dinosaur known from Mongolia.

Heterodontosauridae

Heterodontosauridae is a family of early ornithischian dinosaurs that were likely among the most basal (primitive) members of the group. Although their fossils are relatively rare and their group small in numbers, they lived across all continents except Australia for approximately 100 million years, from the Late Triassic to the Early Cretaceous.

Heterodontosaurids were fox-sized dinosaurs less than 2 metres (6.6 feet) in length, including a long tail. They are known mainly for their characteristic teeth, including enlarged canine-like tusks and cheek teeth adapted for chewing, analogous to those of Cretaceous hadrosaurids. Their diet was herbivorous or possibly omnivorous.

Heterodontosaurinae

Heterodontosaurinae is an extinct subfamily of heterodontosaurid ornithischian dinosaurs from the earliest to the mid Middle Jurassic (Hettangian - Bajocian) of Africa and South America. Currently, the basalmost known heterodontosaurine is Lycorhinus angustidens from the Early Jurassic of Cape Province, South Africa. Heterodontosaurines are small-bodied ornithischians characterized by their cheek tooth crowns that are taller than wide, and jaw joint set below the axis of occlusion between maxillary and dentary teeth. Heterodontosaurinae was implicitly named in 1966 by Oskar Kuhn as he is the author of the family Heterodontosauridae. It is a stem-based taxon defined phylogenetically for the first time by Paul Sereno in 2012 as "the most inclusive clade containing Heterodontosaurus tucki Crompton and Charig 1962 but not Tianyulong confuciusi Zheng et al. 2009, Fruitadens haagarorum Butler et al. 2010, Echinodon becklesii Owen 1861."

Jeholosauridae

Jeholosaurids were herbivorous neornithischian dinosaurs from the Cretaceous Period (Aptian - Santonian, with a possible Campanian record) of Asia. The family was first proposed by Han et al. in 2012. The jeholosaurids were defined as those ornithischians more closely related to Jeholosaurus shangyuanensis than to Hypsilophodon foxii, Iguanodon bernissartensis, Protoceratops andrewsi, Pachycephalosaurus wyomingensis, or Thescelosaurus neglectus. The Jeholosauridae includes the type genus Jeholosaurus and Yueosaurus.

Jingshanosaurus

Jingshanosaurus (meaning "Jingshan lizard") is a genus of sauropodomorph dinosaurs from the early Jurassic period.

Kulindadromeus

Kulindadromeus was a herbivorous dinosaur, a basal neornithischian from the Jurassic. The first Kulindadromeus fossil was found in Russia. Its feather-like integument is evidence for protofeathers being basal to Dinosauria as a whole, rather than just to Coelurosauria, as previously suspected.

Manidens

Manidens is a genus of heterodontosaurid dinosaur from the Middle Jurassic of Patagonia. Fossils have been found in the Cañadón Asfalto Formation in Chubut Province, Argentina, dating to the Bajocian.

Neotheropoda

Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and the only group of theropods who survived the Triassic–Jurassic extinction event. Yet all of the neotheropods became extinct during the early Jurassic period except for Averostra.

Orionides

Orionides is a clade of tetanuran theropod dinosaurs from the Middle Jurassic to the Present. The clade includes most theropod dinosaurs, including birds.

Ornithischia

Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure similar to that of birds. The name Ornithischia, or "bird-hipped", reflects this similarity and is derived from the Greek stem ornith- (ὀρνιθ-), meaning "of a bird", and ischion (ἴσχιον), plural ischia, meaning "hip joint". However, birds are only distantly related to this group as birds are theropod dinosaurs.Ornithischians with well known anatomical adaptations include the ceratopsians or "horn-faced" dinosaurs (e.g. Triceratops), armored dinosaurs (Thyreophora) such as stegosaurs and ankylosaurs, pachycephalosaurids and the ornithopods. There is strong evidence that certain groups of ornithischians lived in herds, often segregated by age group, with juveniles forming their own flocks separate from adults. Some were at least partially covered in filamentous (hair- or feather- like) pelts, and there is much debate over whether these filaments found in specimens of Tianyulong, Psittacosaurus, and Kulindadromeus may have been primitive feathers.

Orodrominae

Orodrominae is a subfamily of parksosaurid dinosaurs from the Cretaceous of North America and Asia.

Riojasauridae

Riojasauridae is a family of sauropod-like dinosaurs from the Upper Triassic. It is known primarily from the genera Riojasaurus and Eucnemesaurus. Sites containing Riojasauridae include the Lower Elliot Formation of Orange Free State, South Africa (where fossils of Eucnemesaurus have been found), and Ischigualasto, in La Rioja Province, Argentina ( where fossils of Riojasaurus have been recovered).

Xixiposaurus

Xixiposaurus is a genus of prosauropod dinosaur which existed in what is now Lower Lufeng Formation, China during the lower Jurassic period. It was first named by Sekiya Toru in 2010 and the type species is Xixiposaurus suni.

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