Epidexipteryx Temporal range: Callovian, | |
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Holotype fossil exhibited in Japan | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Family: | † Scansoriopterygidae |
Genus: | † Epidexipteryx Zhang et al., 2008 |
Species: | †E. hui |
Binomial name | |
†Epidexipteryx hui Zhang et al., 2008 | |
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, belonging to a subadult individual, is catalog number IVPP V 15471. [2] The specific name, Epidexipteryx hui ("Hu's display feather"), and its Chinese name Húshì Yàolóng ("Hu Yaoming's display dragon") were coined in memory of paleomammologist Hu Yaoming. [3]
Due to a pre-publication error, [4] 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]
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. [5]
In all, the skeleton of Epidexipteryx hui measures 25 cm (9.8 in) in length (44.5 cm (17.5 in) including the incomplete tail feathers), [6] and the authors estimated a weight of 164 grams, smaller than most other basal avialans. [2] Gregory S. Paul presented a length estimate of 30 cm (12 in) and body mass estimate of 220 g (0.49 lb). [7]
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]
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). [8] 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). [9] 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.
Cau (2024) cast doubt on the validity of Epidexipteryx, considering it a junior synonym of the contemporary Epidendrosaurus due to the diagnosis being based on incomplete tail remains within the holotype, the length of the presacral series, which is not fully known in Epidendrosaurus, and the co-ossification of the 10 distalmost cervical centra, which is interpreted as an ontologically variable trait. As such, the Epidexipteryx holotype would represent a more mature individual of Epidendrosaurus. [10]
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. [11] 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. [12] Agnolín and Novas (2013) recovered monophyletic Scansoriopterygidae as well, but found them to be non-paravian maniraptorans and the sister group to Oviraptorosauria. [13]
An abbreviated version of Zhang et al.'s 2008 cladogram is presented below.
Maniraptora |
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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] [14]
Epidexipteryx is known from the Middle Jurassic or Upper Jurassic age Daohugou Beds of Inner Mongolia, China (about 160 or 154 mya). [2]
Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discoveries of complete and articulated specimens, have helped to increase understanding about this group. Anatomical studies, particularly studies of the most primitive troodontids, like Sinovenator, demonstrate striking anatomical similarities with Archaeopteryx and primitive dromaeosaurids, and demonstrate that they are relatives comprising a clade called Paraves.
Deinonychosauria is a clade of paravian dinosaurs which lived from the Late Jurassic to the Late Cretaceous periods. Fossils have been found across the globe in North America, Europe, Africa, Asia, South America, and Antarctica, with fossilized teeth giving credence to the possibility that they inhabited Australia as well. This group of dinosaurs are known for their sickle-shaped toe claws and features in the shoulder bones.
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Dromaeosauridae, Troodontidae, Oviraptorosauria, and Therizinosauria. Ornitholestes and the Alvarezsauroidea are also often included. Together with the next closest sister group, the Ornithomimosauria, Maniraptora comprises the more inclusive clade Maniraptoriformes. Maniraptorans first appear in the fossil record during the Jurassic Period, and survive today as living birds.
Coelurosauria is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs.
Yixianosaurus is a maniraptoran theropod dinosaur genus from the Early Cretaceous of China.
Scansoriopteryx is a genus of maniraptoran dinosaur. Described from only a single juvenile fossil specimen found in Liaoning, China, Scansoriopteryx is a sparrow-sized animal that shows adaptations in the foot indicating an arboreal (tree-dwelling) lifestyle. It possessed an unusual, elongated third finger which may have supported a membranous wing, much like the related Yi qi. The type specimen of Scansoriopteryx also contains the fossilized impression of feathers.
Rahonavis is a genus of bird-like theropods from the Late Cretaceous of what is now northwestern Madagascar. It is known from a partial skeleton found by Catherine Forster and colleagues in Maevarano Formation rocks at a quarry near Berivotra, Mahajanga Province. Rahonavis was a small predator, at about 70 centimetres (2.3 ft) long and 0.45-2.27 kg, with the typical dromaesaurid-like raised sickle claw on the second toe. It was originally the first African coelurosaur until the discovery of Nqwebasaurus in 2000.
Scansoriopterygidae 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 of Liaoning and Hebei, China.
Velociraptorinae is a subfamily of the theropod group Dromaeosauridae. The earliest velociraptorines are probably Nuthetes from the United Kingdom, and possibly Deinonychus from North America. However, several indeterminate velociraptorines have also been discovered, dating to the Kimmeridgian stage, in the Late Jurassic Period. These fossils were discovered in the Langenberg quarry, Oker near Goslar, Germany.
Avialae is a clade containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally used.
Paraves are a widespread group of theropod dinosaurs that originated in the Middle Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and possibly the scansoriopterygids, the group also contains the avialans, which include diverse extinct taxa as well as the over 10,000 species of living birds. Basal 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. A number of differing scientific interpretations of the relationships between paravian taxa exist. New fossil discoveries and analyses make the classification of Paraves an active subject of research.
Xiaotingia is a genus of anchiornithid theropod dinosaur from Middle Jurassic or early Late Jurassic deposits of western Liaoning, China, containing a single species, Xiaotingia zhengi.
Eosinopteryx is an extinct genus of theropod dinosaurs known to the Late Jurassic epoch of China. It contains a single species, Eosinopteryx brevipenna.
Aurornis is an extinct genus of anchiornithid theropod dinosaurs from the Jurassic period of China. The genus Aurornis contains a single known species, Aurornis xui. Aurornis xui may be the most basal ("primitive") avialan dinosaur known to date, and it is one of the earliest avialans found to date. The fossil evidence for the animal pre-dates that of Archaeopteryx lithographica, often considered the earliest bird species, by about 10 million years.
Yi is a genus of scansoriopterygid dinosaurs from the Late Jurassic of China. Its only species, Yi qi, is known from a single fossil specimen of an adult individual found in Middle or Late Jurassic Tiaojishan Formation of Hebei, China, approximately 159 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.
Unenlagiidae is a proposed family of eumaniraptoran paravians that includes the subfamilies Unenlagiinae and possibly Halszkaraptorinae. Fossils of both subfamilies have been found in both Gondwanan and Laurasian deposits. The biology of the group suggests that some members were semiaquatic specialists.
Serikornis is a genus of small, feathered anchiornithid dinosaur from the Upper Jurassic Tiaojishan Formation of Liaoning, China. It is represented by the type species Serikornis sungei.
Anchiornithidae is a family of small paravian dinosaurs. Anchiornithids have been classified at varying positions in the paravian 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.
Overoraptor is an extinct genus of paravian theropod of uncertain affinities from the Late Cretaceous Huincul Formation of Argentinian Patagonia. The genus contains a single species, O. chimentoi, known from several bones of the hands, feet, and hips alongside some vertebrae.