Epidexipteryx

Epidexipteryx is a genus of small paravian dinosaurs, known from one fossil specimen in the collection of the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing. Epidexipteryx represents the earliest known example of ornamental feathers in the fossil record.[1] The type specimen is catalog number IVPP V 15471. It has been reported to be a maniraptoran dinosaur from the Middle Jurassic or Upper Jurassic age Daohugou Beds of Inner Mongolia, China (about 160 or 168 mya).[2]

The specific name, Epidexipteryx hui ("Hu's display feather"), and its Chinese name Hushi Yaolong ("Hu Yaoming's dragon") were coined in memory of paleomammologist Hu Yaoming.[3]

Epidexipteryx
Temporal range:
Oxfordian, 160 Ma
Holotype
Holotype fossil skeleton
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Family: Scansoriopterygidae
Genus: Epidexipteryx
Zhang et al., 2008
Species:
E. hui
Binomial name
Epidexipteryx hui
Zhang et al., 2008

Description

Epidexipteryx hui
Skeletal restoration

E. hui is known from a well-preserved partial skeleton that includes four long feathers on the tail, composed of a central rachis and vanes. However, unlike in modern-style rectrices (tail feathers), 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"[2] at the base of each feather. It has been suggested that this may represent a stage in the evolution of the feather.[4]

Epidexipteryx scale
Size of E. hui (holotype specimen) compared with a human.

In all, the skeleton of Epidexipteryx hui measures 25 centimeters (10 inches) in length (44.5 cm or 17.5 in including the incomplete tail feathers),[5] and the authors estimated a weight of 164 grams, smaller than most other basal avialans.[2]

The skull of Epidexipteryx is also unique in a number of features, and bears an overall similarity to the skull of Sapeornis, oviraptorosaurs and, to a lesser extent, therizinosauroids. It had teeth only in the front of the jaws, with unusually long front teeth angled forward, a feature only seen in Masiakasaurus among other theropods. The rest of the skeleton bore an overall similarity to the possibly closely related Scansoriopteryx, including a hip configuration unusual among other dinosaurs: the pubis was shorter than the ischium, and the ischium itself was expanded towards the tip. The tail of Epidexipteryx also bore unusual vertebrae towards the tip which resembled the feather-anchoring pygostyle of modern birds and some oviraptorosaurs.[2]

Epidexipteryx
Restored skull

Epidexipteryx appears to have lacked remiges (wing feathers), though based on the related Yi, it may have possessed some sort of membrane wing to allow gliding.[2][6]

Classification

The exact phylogenetic position of Epidexipteryx within Paraves is uncertain. The phylogenetic analysis conducted by the authors of its description recovered it as a member of the family Scansoriopterygidae and as a basal member of the clade Avialae;[2] this was confirmed by the subsequent analysis conducted by Hu et al. (2009).[7] A later analysis conducted by Agnolín and Novas (2011) confirmed it to be a scansoriopterygid, but recovered a different phylogenetic position of this family: Scansoriopterygidae was recovered in polytomy with the family Alvarezsauridae and the clade Eumaniraptora (containing the clades Avialae and Deinonychosauria).[8] Turner, Makovicky and Norell (2012) included Epidexipteryx but not Scansoriopteryx/Epidendrosaurus in their primary phylogenetic analysis, as a full-grown specimen is known only of the former taxon; regarding Scansoriopteryx/Epidendrosaurus, the authors were worried that including it in the primary analysis would be problematic, because it is only known from juvenile specimens, which "do not necessarily preserve all the adult morphology needed to accurately place a taxon phylogenetically" (Turner, Makovicky and Norell 2012, p. 89). Epidexipteryx was recovered as basal paravian that didn't belong to Eumaniraptora. The authors did note that its phylogenetic position is unstable; constraining Epidexipteryx hui as a basal avialan required two additional steps compared to the most parsimonious solution, while constraining it as a basal member of Oviraptorosauria required only one additional step.

Epidexipteryx NT
Life restoration

A separate exploratory analysis included Scansoriopteryx/Epidendrosaurus, which was recovered as a basal member of Avialae; the authors noted that it did not clade with Epidexipteryx, which stayed outside Eumaniraptora. Constraining the monophyly of Scansoriopterygidae required four additional steps and moved Epidexipteryx into Avialae.[9] A monophyletic Scansoriopterygidae was recovered by Godefroit et al. (2013); the authors found scansoriopterygids to be basalmost members of Paraves and the sister group to the clade containing Avialae and Deinonychosauria.[10] Agnolín and Novas (2013) recovered monophyletic Scansoriopterygidae as well, but found them to be non-paravian maniraptorans and the sister group to Oviraptorosauria.[11]

An abbreviated version of Zhang et al.'s 2008 cladogram is presented below.

Maniraptora

Therizinosauroidea

unnamed

Alvarezsauridae

unnamed

Oviraptorosauria

unnamed

Troodontidae

Dromaeosauridae

Avialae
Scansoriopterygidae

Scansoriopteryx (=Epidendrosaurus)

Epidexipteryx

Aves

Due to a pre-publication error,[12] a manuscript of the Epidexipteryx hui description first appeared on a preprint Web portal in late September 2008. The paper was officially published in the October 23, 2008 issue of the journal Nature.[2]

References

  1. ^ Morgan, James (2008-10-22). "New feathered dinosaur discovered". BBC. Retrieved 2009-07-02.
  2. ^ a b c d e f g Zhang, F.; Zhou, Z.; Xu, X.; Wang, X.; Sullivan, C. (October 2008). "A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers" (PDF). Nature. 455: 1105–1108. doi:10.1038/nature07447. PMID 18948955. Archived from the original (PDF) on 2015-05-24. Retrieved 2015-05-24.
  3. ^ "Chinese scientists discovers new dinosaur species". People's Daily Online. October 27, 2008. Retrieved November 4, 2008.
  4. ^ "Archived copy". Archived from the original on 2011-06-06. Retrieved 2010-12-10.CS1 maint: Archived copy as title (link)
  5. ^ Zhang, F., Zhou, Z., Xu, X., Wang, X. and Sullivan, C. (2008). "A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers", Supplementary Information. Nature, 455: 46pp. doi:10.1038/nature07447 PMID 18948955
  6. ^ Cau, A (2012), Il ritorno del paraviano pterosauro-mimo?, Theropoda, July 2012
  7. ^ Dongyu Hu; Lianhai Hou; Lijun Zhang; Xing Xu (2009). "A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus". Nature. 461 (7264): 640–643. doi:10.1038/nature08322. PMID 19794491.
  8. ^ Agnolín, Federico L.; Novas, Fernando E. (2011). "Unenlagiid theropods: are they members of the Dromaeosauridae (Theropoda, Maniraptora)?". Anais da Academia Brasileira de Ciências. 83 (1): 117–162. doi:10.1590/S0001-37652011000100008.
  9. ^ Alan Hamilton Turner; Peter J. Makovicky; Mark Norell (2012). "A review of dromaeosaurid systematics and paravian phylogeny". Bulletin of the American Museum of Natural History. 371: 1–206. doi:10.1206/748.1.
  10. ^ Pascal Godefroit; Helena Demuynck; Gareth Dyke; Dongyu Hu; François Escuillié & Philippe Claeys (2013). "Reduced plumage and flight ability of a new Jurassic paravian theropod from China". Nature Communications. 4: Article number 1394. doi:10.1038/ncomms2389. PMID 23340434.
  11. ^ Federico L. Agnolín; Fernando E. Novas (2013). "Avian ancestors. A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae". SpringerBriefs in Earth System Sciences: 1–96. doi:10.1007/978-94-007-5637-3.
  12. ^ Dr. Thomas Holtz, Jr. "The mistaken scansoripterygid". Message to the Dinosaur Mailing List <http://dml.cmnh.org/2008Oct/msg00008.html> (October 1, 2008)

External links

2008 in paleontology

Paleontology or palaeontology is the study of prehistoric life forms on Earth through the examination of plant and animal fossils. This includes the study of body fossils, tracks (ichnites), burrows, cast-off parts, fossilised feces (coprolites), palynomorphs and chemical residues. Because humans have encountered fossils for millennia, paleontology has a long history both before and after becoming formalized as a science. This article records significant discoveries and events related to paleontology that occurred or were published in the year 2008.

Anchiornithidae

Anchiornithidae ("near birds") is a family of eumaniraptorans which could be the basalmost family of birds (in the general sense) in the clade Avialae. Anchiornithids have been classified at varying positions in the maniraptoran tree, with some scientists classifying them as a distinct family, a basal subfamily of Troodontidae, members of Archaeopterygidae, or an assemblage of dinosaurs that are an evolutionary grade within Avialae or Paraves.

Dinosaur size

Size has been one of the most interesting aspects of dinosaur science to the general public and to scientists. Dinosaurs show some of the most extreme variations in size of any land animal group, ranging from the tiny hummingbirds, which can weigh as little as three grams, to the extinct titanosaurs, which could weigh as much as 90 tonnes (89 long tons; 99 short tons).Scientists will probably never be certain of the largest and smallest dinosaurs to have ever existed. This is because only a tiny fraction of animals ever fossilize, and most of these remain buried in the earth. Few of the specimens that are recovered are complete skeletons, and impressions of skin and other soft tissues are rare. Rebuilding a complete skeleton by comparing the size and morphology of bones to those of similar, better-known species is an inexact art, and reconstructing the muscles and other organs of the living animal is, at best, a process of educated guesswork. Weight estimates for dinosaurs are much more variable than length estimates, because estimating length for extinct animals is much more easily done from a skeleton than estimating weight. Estimating weight is most easily done with the laser scan skeleton technique that puts a "virtual" skin over it, but even this is only an estimate.Current evidence suggests that dinosaur average size varied through the Triassic, early Jurassic, late Jurassic and Cretaceous periods. Predatory theropod dinosaurs, which occupied most terrestrial carnivore niches during the Mesozoic, most often fall into the 100- to 1,000-kilogram (220 to 2,200 lb) category when sorted by estimated weight into categories based on order of magnitude, whereas recent predatory carnivoran mammals peak in the 10- to 100-kilogram (22 to 220 lb) category. The mode of Mesozoic dinosaur body masses is between one and ten metric tonnes. This contrasts sharply with the size of Cenozoic mammals, estimated by the National Museum of Natural History as about 2 to 5 kg (4.4 to 11.0 lb).

Feathered dinosaur

Since scientific research began on dinosaurs in the early 1800s, they were generally believed to be closely related to modern reptiles, such as lizards. The word "dinosaur" itself, coined in 1842 by paleontologist Richard Owen, comes from the Greek for "fearsome lizard". This view began to shift during the so-called dinosaur renaissance in scientific research in the late 1960s, and by the mid-1990s significant evidence had emerged that dinosaurs were much more closely related to birds, which descended directly from the theropod group of dinosaurs and are themselves a subgroup within the Dinosauria.

Understanding of the origin of feathers developed both as new fossils were discovered throughout the 2000s and 2010s and as technology has enabled scientists to study fossils more closely. Among non-avian dinosaurs, feathers or feather-like integument have been discovered in dozens of genera via direct and indirect fossil evidence. Although the vast majority of feather discoveries have been in coelurosaurian theropods, feather-like integument has also been discovered in at least three ornithischians, suggesting that feathers may have been present on the last common ancestor of the Ornithoscelida, a dinosaur group including both theropods and ornithischians. It is possible that feathers first developed in even earlier archosaurs, in light of the discovery of highly feather-like pycnofibers in pterosaurs. Crocodilians also possess beta keratin similar to those of birds, which suggests that they evolved from common ancestral genes.

Jeholornis

Jeholornis (meaning "Jehol bird") is a genus of avialans that lived between approximately 122 and 120 million years ago during the early Cretaceous Period in China. Fossil Jeholornis were first discovered in the Jiufotang Formation in Hebei Province, China (in what was previously Rehe Province, also known as Jehol—hence the name) and additional specimens have been found in the older Yixian Formation.Jeholornis had long tails and few small teeth, and were approximately the size of turkeys, making them among the largest avialans known until the Late Cretaceous. Their diet included seeds of cycads, Ginkgo or similar plants.

List of dinosaur genera

This list of dinosaurs is a comprehensive listing of all genera that have ever been included in the superorder Dinosauria, excluding class Aves (birds, both living and those known only from fossils) and purely vernacular terms.

The list includes all commonly accepted genera, but also genera that are now considered invalid, doubtful (nomen dubium), or were not formally published (nomen nudum), as well as junior synonyms of more established names, and genera that are no longer considered dinosaurs. Many listed names have been reclassified as everything from birds to crocodilians to petrified wood. The list contains 1559 names, of which approximately 1192 are considered either valid dinosaur genera or nomina dubia.

List of non-avian dinosaur species preserved with evidence of feathers

Several non-avian dinosaurs were feathered. Direct evidence of feathers exists for the following species, listed in the order currently accepted evidence was first published. In all examples, the evidence described consists of feather impressions, except those genera inferred to have had feathers based on skeletal or chemical evidence, such as the presence of quill knobs (the anchor points for wing feathers on the forelimb) or a pygostyle (the fused vertebrae at the tail tip which often supports large feathers).

Ostromia crassipes (1970)

Avimimus portentosus (inferred 1987: ulnar ridge)

Sinosauropteryx prima (1996)

Fulicopus lyellii, an ichnotaxon, possible squatting Dilophosaurus or similar. (1996)

Protarchaeopteryx robusta (1997)

GMV 2124 (1997)

Caudipteryx zoui (1998)

Rahonavis ostromi (inferred 1998: quill knobs; possibly avialan)

Shuvuuia deserti (1999)

Beipiaosaurus inexpectus (1999)

Sinornithosaurus millenii (1999)

Caudipteryx dongi (2000)

Caudipteryx sp. (2000)

Microraptor zhaoianus (2000)

Nomingia gobiensis (inferred 2000: pygostyle)

Psittacosaurus sp.? (2002)

Scansoriopteryx heilmanni (2002; possibly avialan)

Yixianosaurus longimanus (2003)

Dilong paradoxus (2004)

Pedopenna daohugouensis (2005; possibly avialan)

Jinfengopteryx elegans (2005)

Juravenator starki (2006)

Sinocalliopteryx gigas (2007)

Velociraptor mongoliensis (inferred 2007: quill knobs)

Epidexipteryx hui (2008; possibly avialan)

Similicaudipteryx yixianensis (inferred 2008: pygostyle; confirmed 2010)

Anchiornis huxleyi (2009; possibly avialan)

Tianyulong confuciusi? (2009)

Concavenator corcovatus? (inferred 2010: quill knobs?)

Xiaotingia zhengi (2011; possibly avialan)

Yutyrannus huali (2012)

Sciurumimus albersdoerferi (2012)

Ornithomimus edmontonicus (2012)

Ningyuansaurus wangi (2012)

Eosinopteryx brevipenna (2013; possibly avialan)

Jianchangosaurus yixianensis (2013)

Aurornis xui (2013; possibly avialan)

Changyuraptor yangi (2014)

Kulindadromeus zabaikalicus? (2014)

Citipati osmolskae (inferred 2014: pygostyle)

Conchoraptor gracilis (inferred 2014: pygostyle)

Deinocheirus mirificus? (inferred 2014: pygostyle)

Yi qi (2015)

Ornithomimus sp. (2015)

Zhenyuanlong suni (2015)

Dakotaraptor steini (inferred 2015: quill knobs)

Apatoraptor pennatus (inferred 2016: quill knobs)

Jianianhualong tengi (2017)

Serikornis sungei (2017)

Caihong juji (2018)

Xingtianosaurus ganqi (2019)

Ambopteryx longibrachium (2019)Note that the filamentous structures in some ornithischian dinosaurs (Psittacosaurus, Tianyulong and Kulindadromeus) and the pycnofibres found in some pterosaurs may or may not be homologous with the feathers of theropods.

Maniraptoriformes

Maniraptoriformes is a clade of dinosaurs with pennaceous feathers and wings that contains ornithomimosaurs and maniraptors. This group was named by Thomas Holtz, who defined it as "the most recent common ancestor of Ornithomimus and birds, and all descendants of that common ancestor."

Origin of birds

The scientific question of within which larger group of animals birds evolved, has traditionally been called the origin of birds. The present scientific consensus is that birds are a group of theropod dinosaurs that originated during the Mesozoic Era.

A close relationship between birds and dinosaurs was first proposed in the nineteenth century after the discovery of the primitive bird Archaeopteryx in Germany. Birds and extinct non-avian dinosaurs share many unique skeletal traits. Moreover, fossils of more than thirty species of non-avian dinosaur have been collected with preserved feathers. There are even very small dinosaurs, such as Microraptor and Anchiornis, which have long, vaned, arm and leg feathers forming wings. The Jurassic basal avialan Pedopenna also shows these long foot feathers. Paleontologist Lawrence Witmer concluded in 2009 that this evidence is sufficient to demonstrate that avian evolution went through a four-winged stage. Fossil evidence also demonstrates that birds and dinosaurs shared features such as hollow, pneumatized bones, gastroliths in the digestive system, nest-building and brooding behaviors.

Although the origin of birds has historically been a contentious topic within evolutionary biology, only a few scientists still debate the dinosaurian origin of birds, suggesting descent from other types of archosaurian reptiles. Within the consensus that supports dinosaurian ancestry, the exact sequence of evolutionary events that gave rise to the early birds within maniraptoran theropods is disputed. The origin of bird flight is a separate but related question for which there are also several proposed answers.

Paraves

Paraves are a widespread group of theropod dinosaurs that originated in the Late Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and scansoriopterygids, the group also contains the avialans, among which are the over ten thousand species of living birds. Primitive members of Paraves are well known for the possession of an enlarged claw on the second digit of the foot, which was held off the ground when walking in some species.

Planet Dinosaur

Planet Dinosaur, is a six-part documentary television series created by Nigel Paterson and Phil Dobree, produced by the BBC, and narrated by John Hurt. It first aired in the United Kingdom in 2011, with VFX studio Jellyfish Pictures as its producer. It is the first major dinosaur-related series for BBC One since Walking with Dinosaurs. There are more than 50 different prehistoric species featured, and they and their environments were created entirely as computer-generated images, for around a third of the production cost that was needed a decade earlier for Walking with Dinosaurs. Much of the series' plot is based on scientific discoveries made since Walking with Dinosaurs. The companion book to Planet Dinosaur was released on 8 September 2011, and the DVD and Blu-ray were released on 24 October 2011.

Saurornithoides

Saurornithoides ( saw-ROR-ni-THOY-deez) is a genus of troodontid maniraptoran dinosaur, which lived during the Late Cretaceous period. These creatures were predators, which could run fast on their hind legs and had excellent sight and hearing. The name is derived from the Greek stems saur~ (lizard), ornith~ (bird) and eides (form), referring to its bird-like skull.

Scansoriopterygidae

Scansoriopterygidae (meaning "climbing wings") is an extinct family of climbing and gliding maniraptoran dinosaurs. Scansoriopterygids are known from five well-preserved fossils, representing four species, unearthed in the Tiaojishan Formation fossil beds (dating to the mid-late Jurassic Period) of Liaoning and Hebei, China.

Scansoriopteryx heilmanni (and its likely synonym Epidendrosaurus ninchengensis) was the first non-avian dinosaur found that had clear adaptations to an arboreal or semi-arboreal lifestyle–it is likely that they spent much of their time in trees. Both specimens showed features indicating they were juveniles, which made it difficult to determine their exact relationship to other non-avian dinosaurs and birds. It was not until the description of Epidexipteryx hui in 2008 that an adult specimen was known. In 2015, the discovery of another, larger adult specimen belonging to the species Yi qi showed that scansoriopterygids were not only climbers but also had adaptations that could have been used for gliding flight. The recently discovered (in 2019) Ambopteryx also proves this.

Similicaudipteryx

Similicaudipteryx, meaning "similar to Caudipteryx" (which itself means "tail feather"), is a genus of theropod dinosaur of the family Caudipteridae.

Sinornithoides

Sinornithoides (meaning "Chinese bird form") is a genus of troodontid theropod dinosaurs containing the single species Sinornithoides youngi. S. youngi lived during the Early Cretaceous (Aptian/Albian stage, around 113 million years ago). It measured approximately one meter long (3.3 ft). It lived in Inner Mongolia, China, and probably ate invertebrates and other small prey. They live in what is now Mongolia, which was part of Laurasia.

Temporal paradox (paleontology)

The temporal paradox, or time problem is a controversial issue in the evolutionary relationships of birds. It was described by paleornithologist Alan Feduccia in 1994. It reflects a conflict between where chronologically fossils are expected to be found based on a proposed phylogeny, and where they are actually found. The temporal paradox can give rise to alternative phylogenies (i.e. the archosaurian hypothesis of bird evolution).

Statistical measures have been devised to assess the likelihood of proposed phylogenies based on the size and positioning of gaps in the fossil record.

Tiaojishan Formation

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 fossil plants, and 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 "Lanqui Formation."

Most researchers now agree that the Daohugou Bed, of formerly controversial dating, is a part of the Tiaojishan formation.

Yi (dinosaur)

Yi is a genus of scansoriopterygid dinosaurs from the Late Jurassic of China. Its only species, Yi qi (Mandarin pronunciation: [î tɕʰǐ]; from Chinese: 翼; pinyin: yì; literally: 'wing' and 奇; qí; 'strange'), is known from a single fossil specimen of an adult individual found in Middle or Late Jurassic Tiaojishan Formation of Hebei, China, approximately 160 million years ago. It was a small, possibly tree-dwelling (arboreal) animal. Like other scansoriopterygids, Yi possessed an unusual, elongated third finger, that appears to have helped to support a membranous gliding plane made of skin. The planes of Yi qi 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.

Zhongornis

Zhongornis (meaning "intermediate bird") is a genus of primitive avialan that lived during the Early Cretaceous. It was found in rocks of the Yixian Formation in Lingyuan City (China), and described by Gao et al. in 2008.Zhongornis has only one described species, Zhongornis haoae. The only specimen is a fossil slab and counterslab numbered D2455/6. It is in the collection of the Dalian Natural History Museum. It is a fairly complete skeleton about eight centimeters in length. Pores in the bones and unfused sutures in the skeleton indicate that the specimen was a juvenile, but the authors believe that it was developed enough to erect a new taxon on the basis of its unique morphological characters. There are feather impressions preserved on the right hand and also probable tail feathers preserved near the left foot. Zhongornis had a beaked mouth with no teeth. The tail is proportionately short, has thirteen vertebrae, and no pygostyle. The third finger has only two phalangeal bones, unlike non - avian dinosaurs and Confuciusornis, and more like Enantiornithes and more advanced avialans. These features and a cladistic analysis indicate that Zhongornis is the sister group to all pygostylia, meaning that it is intermediate between long - tailed Avialae like Archaeopteryx and more advanced taxa like Confuciusornis.

Coelurosauria

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