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Epidexipteryx

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Epidexipteryx
Temporal range:
Callovian, 164 Ma
Holotype
Holotype fossil, Paleozoological Museum of China
Scientific classification Edit this 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 maniraptoran 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]

Discovery

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Replica of the holotype in Japan

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]

Description

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The scansoriopterygid genera Epidexipteryx (orange), Yi (green), and Scansoriopteryx (red) compared to a human in size

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]

Skeletal diagram showing known elements of the holotype

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]

Classification

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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]

Life restoration showing the animal without arm membranes
Reconstructions of the skull

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

Paleobiology

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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]

Paleoenvironment

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Epidexipteryx is known from the Middle Jurassic or Upper Jurassic age Daohugou Beds of Inner Mongolia, China (about 160 or 154 mya).[2]

References

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  1. ^ Morgan, James (2008-10-22). "New feathered dinosaur discovered". BBC. Retrieved 2009-07-02.
  2. ^ a b c d e f g h 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 (7216): 1105–1108. Bibcode:2008Natur.455.1105Z. doi:10.1038/nature07447. PMID 18948955. S2CID 4362560. 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. Archived from the original on February 2, 2009. Retrieved November 4, 2008.
  4. ^ Dr. Thomas Holtz, Jr. "The mistaken scansoripterygid". Message to the Dinosaur Mailing List <http://dml.cmnh.org/2008Oct/msg00008.html Archived 2016-03-03 at the Wayback Machine> (October 1, 2008)
  5. ^ "Epidexipteryx: Bizarre little strap-feathered maniraptoran : Tetrapod Zoology". Archived from the original on 2011-06-06. Retrieved 2010-12-10.
  6. ^ Zhang, F.; Zhou, Z.; Xu, X.; Wang, X.; Sullivan, C. (2008). ""A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers", Supplementary Information". Nature. 455 (7216): 1105–8. Bibcode:2008Natur.455.1105Z. doi:10.1038/nature07447. PMID 18948955. S2CID 4362560.
  7. ^ Paul, Gregory S. (2016). The Princeton Field Guide to Dinosaurs. Princeton University Press. p. 139. ISBN 978-1-78684-190-2. OCLC 985402380.
  8. ^ 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. Bibcode:2009Natur.461..640H. doi:10.1038/nature08322. PMID 19794491. S2CID 205218015.
  9. ^ 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. PMID 21437379.
  10. ^ Cau, Andrea (2024). "A Unified Framework for Predatory Dinosaur Macroevolution" (PDF). Bollettino della Società Paleontologica Italiana. 63 (1): 1-19. doi:10.4435/BSPI.2024.08.
  11. ^ 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. hdl:2246/6352. S2CID 83572446.
  12. ^ 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. Bibcode:2013NatCo...4.1394G. doi:10.1038/ncomms2389. PMID 23340434.
  13. ^ 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. ISBN 978-94-007-5636-6. S2CID 199493087.
  14. ^ Cau, A (2012), Il ritorno del paraviano pterosauro-mimo?, Theropoda, July 2012
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