Epidexipteryx
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''Epidexipteryx'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of small
paravian 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 aviala ...
dinosaurs, known from one fossil specimen in the collection of the
Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name suggest ...
in
Beijing } Beijing ( ; ; ), alternatively romanized as Peking ( ), is the capital of the People's Republic of China. It is the center of power and development of the country. Beijing is the world's most populous national capital city, with over 21 ...
. ''Epidexipteryx'' represents the earliest known example of ornamental feathers in the fossil record.


Discovery

The
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
is catalog number IVPP V 15471. 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. Due to a pre-publication error,Dr. Thomas Holtz, Jr. "The mistaken scansoripterygid". Message to the Dinosaur Mailing List (October 1, 2008) 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''.


Description

''E. hui'' is known from a well-preserved partial skeleton that includes four long feathers on the tail, composed of a central
rachis In biology, a rachis (from the grc, ῥάχις [], "backbone, spine") is a main axis or "shaft". In zoology and microbiology In vertebrates, ''rachis'' can refer to the series of articulated vertebrae, which encase the spinal cord. In this c ...
and vanes. However, unlike in modern-style
rectrices Flight feathers (''Pennae volatus'') are the long, stiff, asymmetrically shaped, but symmetrically paired pennaceous feathers on the wings or tail of a bird; those on the wings are called remiges (), singular remex (), while those on the tail ...
(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" at the base of each feather. It has been suggested that this may represent a stage in the evolution of the feather. In all, the skeleton of ''Epidexipteryx hui'' measures in length ( including the incomplete tail feathers), and the authors estimated a weight of 164 grams, smaller than most other basal avialans. Gregory S. Paul presented a length estimate of and body mass estimte of . The skull of ''Epidexipteryx'' is also unique in a number of features, and bears an overall similarity to the skull of ''
Sapeornis ''Sapeornis'' is a monotypic genus of avialan which lived during the early Cretaceous period ( late Barremian to early Aptian, roughly 125-120 mya). ''Sapeornis'' contains only one species, ''Sapeornis chaoyangensis''. Description ''Sapeornis' ...
'',
oviraptorosaur Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period (geology), Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like s ...
s and, to a lesser extent,
therizinosaur Therizinosaurs (once called segnosaurs) were large herbivorous theropod dinosaurs whose fossils have been found across the Early to Late Cretaceous deposits in Asia and North America. Various features of the forelimbs, skull and pelvis unite thes ...
oids. It had teeth only in the front of the jaws, with unusually long front teeth angled forward, a feature only seen in ''
Masiakasaurus ''Masiakasaurus'' is a genus of small predatory noasaurid theropod dinosaurs from the Late Cretaceous of Madagascar. In Malagasy, ''masiaka'' means "vicious"; thus, the genus name means "vicious lizard". The type species, ''Masiakasaurus knopfl ...
'' 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 The ischium () form ...
, 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 Pygostyle describes a skeletal condition in which the final few caudal vertebrae are fused into a single ossification, supporting the tail feathers and musculature. In modern birds, the rectrices attach to these. The pygostyle is the main compon ...
of modern birds and some oviraptorosaurs.


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 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 Formatio ...
and as a basal member of the clade
Avialae Avialae ("bird wings") 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 ...
; this was confirmed by the subsequent analysis conducted by Hu ''et al.'' (2009). 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 An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with the family
Alvarezsauridae Alvarezsauridae is a family of small, long-legged dinosaurs. Although originally thought to represent the earliest known flightless birds, they are now thought to be an early diverging branch of maniraptoran theropods. Alvarezsaurids were highly ...
and the clade Eumaniraptora (containing the clades Avialae and
Deinonychosauria 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,Case, J.A., Mar ...
). 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 Oviraptorosaurs ("egg thief lizards") are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot-like skulls, with or wit ...
required only one additional step. 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. 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. Agnolín and Novas (2013) recovered monophyletic Scansoriopterygidae as well, but found them to be non-paravian maniraptorans and the sister group to Oviraptorosauria. An abbreviated version of Zhang ''et al.s 2008
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
is presented below.


Paleobiology

''Epidexipteryx'' appears to have lacked
remiges Flight feathers (''Pennae volatus'') are the long, stiff, asymmetrically shaped, but symmetrically paired pennaceous feathers on the wings or tail of a bird; those on the wings are called remiges (), singular remex (), while those on the tail ...
(wing feathers), though based on the related '' Yi'', it may have possessed some sort of membrane wing to allow gliding.Cau, A (2012)
Il ritorno del paraviano pterosauro-mimo?
Theropoda, July 2012


Paleoenvironment

''Epidexipteryx'' is known from the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
or
Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
age
Daohugou Beds The Haifanggou Formation (), also known as the Jiulongshan Formation (), is a fossil-bearing rock deposit located near Daohugou () village of Ningcheng County, in Inner Mongolia, northeastern China. The formation consists of coarse conglomerates ...
of
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an autonomous region of the People's Republic of China. Its border includes most of the length of China's border with the country of Mongolia. Inner Mongolia also accounts for a ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
(about 160 or 154 mya).


References


External links


New feathered dinosaur discovered
''
BBC News BBC News is an operational business division of the British Broadcasting Corporation (BBC) responsible for the gathering and broadcasting of news and current affairs in the UK and around the world. The department is the world's largest broadca ...
'' {{Taxonbar, from=Q132600 Prehistoric paravians Jurassic dinosaurs of Asia Fossil taxa described in 2008 Feathered dinosaurs