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: ''For the Permian gliding reptile, see [[Coelurosauravus]].''
 
 
{{Automatic taxobox |
 
{{Automatic taxobox |
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| name = Coelurosaurians
|edit_link=
 
 
| fossil_range = <br>[[Late Jurassic]]&ndash;[[Holocene|Present]], {{Fossil range|165|0|earliest=196}}<small>Possible [[Early Jurassic]] record</small>
|color= {{Taxobox_colour|[[Animalia]]}}
 
 
| image = Zuni Coelurosaur.jpg
| name = Coelurosaurs
 
| fossil_range = [[Middle Jurassic]]&ndash;Present, {{Fossil range|165|0|earliest=196}}<small>Possible [[Early Jurassic]] record</small>
 
| image = Coelurus BW.jpg
 
 
| image_width = 250px
 
| image_width = 250px
| image_caption = Life restoration of ''[[Coelurus fragilis]]''
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| image_caption = Reconstructed coelurosaur skeleton, [[Wyoming Dinosaur Center]]
 
| authority = [[Friedrich von Huene|von Huene]], 1914
 
| authority = [[Friedrich von Huene|von Huene]], 1914
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| subdivision_ranks = Subgroups<ref name="Holtz2008">Holtz, Thomas R. Jr. (2012) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'' [http://www.geol.umd.edu/~tholtz/dinoappendix/HoltzappendixWinter2011.pdf Winter 2011 Appendix.]</ref>
| subdivision_ranks = Subgroups
 
| display_children = 1
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| subdivision =
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*{{extinct}}''[[Aniksosaurus]]''<ref name="Holtz2008"/>
| synonyms =
 
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*{{extinct}}''[[Bagaraatan]]''<ref name="Holtz2008"/>
* '''Tyrannoraptora'''? <small>[[Paul Sereno|Sereno]], 1999</small>
 
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*{{extinct}}''[[Bicentenaria]]''
* [[Avifilopluma]]? <small>[[Jacques Gauthier|Gauthier]] & [[Kevin de Queiroz|de Queiroz]], 2001</small>
 
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*{{extinct}}''[[Coelurus]]''?
* [[Avipluma]]? <small>Clarke ''et al.'', 2004</small>
 
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*{{extinct}}''[[Lourinhanosaurus]]''?<ref name=carranno2012>Carrano, M.T., Benson, R.B.J., and Sampson, S.D. 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). ''Journal of Systematic Palaeontology'', '''10'''(2): 211-300.</ref>
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*{{extinct}}''[[Nedcolbertia]]''<ref name="Holtz2008"/>
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*{{extinct}}''[[Phaedrolosaurus]]''<ref name="Holtz2008"/>
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*{{extinct}}''[[Richardoestesia]]''<ref name="Holtz2008"/>
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*{{extinct}}''[[Sciurumimus]]''?<ref name=AurornisNature>{{Cite journal | last1 = Godefroit | first1 = Pascal | last2 = Cau| first2 = Andrea | last3 = Hu | first3 = Dong-Yu| last4 = Escuillié | first4 = François| last5 = Wu | first5 = Wenhao| last6 = Dyke | first6 = Gareth| doi = 10.1038/nature12168 | title = A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds | journal = Nature | volume = 498| issue = 7454| pages =359–362| year = 2013 | pmid = 23719374| pmc = | bibcode = 2013Natur.498..359G }}</ref>
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*{{extinct}}''[[Tanycolagreus]]''?
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*{{extinct}}''[[Teinurosaurus]]''<ref name="Holtz2008">Holtz, Thomas R. Jr. (2012) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'' [http://www.geol.umd.edu/~tholtz/dinoappendix/HoltzappendixWinter2011.pdf Winter 2011 Appendix.]</ref>
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*{{extinct}}''[[Tugulusaurus]]''
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*{{extinct}}''[[Xinjiangovenator]]''<ref name="Holtz2008"/>
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*{{extinct}}''[[Zuolong]]''
 
*{{anchor|Tyrannoraptora}}'''Tyrannoraptora''' <small>[[Paul Sereno|Sereno]], 1999</small>
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**{{extinct}}''[[Aorun]]''
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**{{extinct}}''[[Ornitholestes]]''
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**{{extinct}}''[[Scipionyx]]''
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**{{extinct}}[[Compsognathidae]]
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**{{extinct}}[[Tyrannosauroidea]]
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**[[Maniraptoriformes]]
 
}}
 
}}
'''Coelurosauria''' {{IPAc-en|s|ɨ|ˌ|lj|ʊər|ə|ˈ|s|ɔr|i|ə}} (from [[Greek language|Greek]], meaning "hollow tailed lizards") is the [[clade]] containing all [[theropod]] [[dinosaur]]s more closely related to [[bird]]s than to [[carnosaurs]]. In the past, it was used to refer to all small theropods, although this classification has been abolished. Nonetheless, it is still a diverse group that includes [[Compsognathidae|compsognathids]], [[Tyrannosauroidea|tyrannosaurs]], [[Ornithomimosauria|ornithomimosaurs]], and [[Maniraptora|maniraptorans]]; Maniraptora includes [[bird]]s, the only dinosaurs alive today. Most [[feathered dinosaurs]] discovered so far have been coelurosaurs; [[Philip J. Currie]] considers it probable that all coelurosaurs were feathered.<ref name="currie-2005" />
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'''Coelurosauria''' ({{IPAc-en|s|ɪ|ˌ|l|jʊər|ə|ˈ|s|ɔːr|i|ə}};{{refn|{{cite web |url=https://www.oxforddictionaries.com/definition/english/coelurosaur |title=coelurosaur - definition of coelurosaur in English from the Oxford dictionary |publisher=[[OxfordDictionaries.com]] |access-date=2016-01-20 }}}}{{refn|{{Dictionary.com|coelurosaur}}}} from [[Greek language|Greek]], meaning "hollow tailed lizards") is the [[clade]] containing all [[theropod]] [[dinosaur]]s more closely related to [[bird]]s than to [[carnosaurs]].
  +
  +
Coelurosauria is a subgroup of theropod dinosaurs that includes [[Compsognathidae|compsognathids]], [[Tyrannosauroidea|tyrannosaurs]], [[Ornithomimosauria|ornithomimosaurs]], and [[maniraptora]]ns; Maniraptora includes [[bird]]s, the only dinosaur group alive today.<ref>Turner, A.H., Makovicky, P.J., and Norell, M.A. 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History 371: 1–206.</ref>
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  +
Most [[feathered dinosaurs]] discovered so far have been coelurosaurs. [[Philip J. Currie]] considers it probable that all coelurosaurs were feathered.<ref name="currie-2005" /> In the past, ''Coelurosauria'' was used to refer to all small theropods, although this classification has been abolished.
   
 
==Anatomy==
 
==Anatomy==
  +
 
===Bodyplan===
 
===Bodyplan===
All coelurosaurs were bipedal, and most were carnivores, though many groups exhibited a more varied diet including insectivory ([[alvarezsauridae]]), omnivory ([[oviraptoridae]] and [[troodontidae]]), and herbivory ([[therizinosauridae]]).<ref name="zannoetal2009" /> The group includes some of the largest (''[[Tyrannosaurus]]'') and smallest (''[[Microraptor]]'', ''[[Parvicursor]]'') carnivorous dinosaurs ever discovered. Characteristics that distinguish coelurosaurs include:
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The studying of anatomical traits in coelurosaurs indicates that the last common ancestor had evolved the ability to eat and digest plant matter, adapting to an omnivorous diet, an ability that could be a major contributor to the clade's success. Later groups would hold on to the omnivory, while others specialized in various directions, becoming insectivorous ([[Alvarezsauridae]]), herbivorous ([[Therizinosauridae]]) and carnivorous ([[Tyrannosauroidea]] and [[Dromaeosauridae]]).<ref>{{cite web|url=http://www.sciencedaily.com/releases/2010/12/101220163052.htm|title=Meat-eating dinosaurs not so carnivorous after all|work=ScienceDaily}}</ref> The group includes some of the largest (''[[Tyrannosaurus]]'') and smallest (''[[Microraptor]]'', ''[[Parvicursor]]'') carnivorous dinosaurs ever discovered. Characteristics that distinguish coelurosaurs include:
 
* a [[sacrum]] (series of vertebrae that attach to the hips) longer than in other dinosaurs
 
* a [[sacrum]] (series of vertebrae that attach to the hips) longer than in other dinosaurs
 
* a tail stiffened towards the tip
 
* a tail stiffened towards the tip
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===Integument===
 
===Integument===
 
{{Main|Feathered dinosaurs}}
 
{{Main|Feathered dinosaurs}}
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Fossil evidence shows that the skin of even the most primitive coelurosaurs was covered primarily in [[feather]]s. Fossil traces of feathers, though rare, have been found in members of most major coelurosaurian lineages. Most coelurosaurs also retained scales and scutes on some portion of their bodies, particularly the feet, though some primitive coelurosaurian species are known to have had scales on the upper legs and portions of the tail as well. These include [[tyrannosauroid]]s, ''[[Juravenator]]'', and ''[[Scansoriopteryx]]''. Fossils of at least some of these animals (''Scansoriopteryx'' and possibly ''Juravenator'') also preserve feathers elsewhere on the body.
   
  +
Though once thought to be a feature exclusive to coelurosaurs, feathers or feather-like structures are also known in some [[ornithischia]]n dinosaurs (like ''[[Tianyulong]]'' ''and [[Kulindadromeus]]''), and in [[pterosaur]]s. Though it is unknown whether these are related to true feathers, recent analysis has suggested that the feather like integument found in ornithischians may have evolved independently of coelurosaurs.<ref>{{Cite journal|url = http://rsbl.royalsocietypublishing.org/content/11/6/20150229|title = Evolution of Dinosaur Epidermal Structures|last = Barrett|first = Paul M.|date = 3 June 2015|journal = The Royal Society Biology Letters|doi = |pmid = |access-date = }}</ref>
To date, fossilized traces of feathers have been identified among most coelurosaurian lineages. Feathers of some type, or morphological features suggesting feathers, have been found in fossils of at least one species in all coelurosaur subgroups other than [[ornithomimosauria]]. Modern [[bird]]s are classified as coelurosaurs by nearly all palaeontologists.<ref name="mayeretal2005" /> The arrangements of feathers currently observed on Coelurosauria other than modern birds are without exception more primitive, and some coelurosaurian species are known to have had bare or scaly skin rather than feathers on at least some parts of their bodies. These include large tyrannosaurids, some compsognathids such as ''[[Juravenator]]'', and ''[[Scansoriopteryx]]''. All preserve the impression of scales from regions near the hind legs and tail, though ''Scansoriopteryx'' and possibly ''Juravenator'' also preserve feathers elsewhere on the body. Most coelurosaurs, including modern birds, retained scales on the feet, though in a few (such as ''[[Anchiornis]]'' and the modern [[Rock Ptarmigan]]) the feet and toes are also entirely covered in feathers.
 
 
Though once thought to be a feature exclusive to coelurosaurs, feathers or feather-like structures are also known in some [[ornithischia]]n dinosaurs (like ''[[Tianyulong]]''), though whether these are related to true feathers or novel structures that evolved independently is unknown.
 
   
 
===Nervous system and senses===
 
===Nervous system and senses===
Although rare, complete casts of theropod [[endocrania]] are known from fossils. Theropod endocrania can also be reconstructed from preserved braincases without damaging valuable specimens by using a computed tomography scan and 3D reconstruction software. These finds are of evolutionary significance because they help document the emergence of the neurology of modern birds from that of earlier reptiles. An increase in the proportion of the brain occupied by the cerebrum seems to have occurred with the advent of the Coelurosauria and "continued throughout the evolution of maniraptorans and early birds."<ref name="csaharicus-endo-abs-19" />
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Although rare, complete casts of theropod [[endocrania]] are known from fossils. Theropod endocrania can also be reconstructed from preserved braincases without damaging valuable specimens by using a [[X-ray computed tomography|computed tomography scan]] and 3D reconstruction software. These finds are of evolutionary significance because they help document the emergence of the neurology of modern birds from that of earlier reptiles. An increase in the proportion of the brain occupied by the cerebrum seems to have occurred with the advent of the Coelurosauria and "continued throughout the evolution of maniraptorans and early birds."<ref name="csaharicus-endo-abs-19" />
   
 
==Fossil evidence and age==
 
==Fossil evidence and age==
 
A few fossil traces tentatively associated with the Coelurosauria date back as far as the late [[Triassic]]. What has been found between then and the start of the late [[Jurassic]] is fragmentary. A typical example is ''[[Iliosuchus]]'', known only from two [[Ilium (bone)|ilia]] bones in the mid Jurassic. It was a 1.5&nbsp;m long carnivore from about 165&nbsp;Ma (million years ago) in Oxfordshire and is tentatively assigned to the [[Tyrannosauroidea]].
   
 
Many nearly complete fossil coelurosaurians are known from the late Jurassic. ''[[Archaeopteryx]]'' (incl. ''Wellnhoferia'') is known from Bavaria at 155-150&nbsp;Ma. ''[[Ornitholestes]]'', the troodontid [[WDC DML 001]], ''[[Coelurus|Coelurus fragilis]]'' and ''[[Tanycolagreus|Tanycolagreus topwilsoni]]'' are all known from the [[Morrison Formation]] in Wyoming at about 150 Ma. ''[[Epidendrosaurus]]'' and ''[[Pedopenna]]'' are known from the [[Daohugou Beds]] in China, whose age is still being debated, but may be about 160&nbsp;Ma or 145&nbsp;Ma.
A few fossil traces tentatively associated with the Coelurosauria date back as far as the late [[Triassic]].<ref name="Dinodata" /> What has been found between then and the start of the late [[Jurassic]] is fragmentary. A typical example is ''[[Iliosuchus]]'', known only from two [[ilia]] bones in the mid Jurassic. It was a 1.5&nbsp;m long carnivore from about 165&nbsp;Ma (million years ago) in Oxfordshire and is tentatively assigned to the [[Tyrannosauroidea]].
 
 
Many nearly complete fossil coelurosaurians are known from the late Jurassic. ''[[Archaeopteryx]]'' (incl. ''Wellnhoferia'') is known from Bavaria at 155-150&nbsp;Ma. ''[[Ornitholestes]]'', the troodontid WDC DML 110, ''[[Coelurus|Coelurus fragilis]]'' and ''[[Tanycolagreus|Tanycolagreus topwilsoni]]'' are all known from the [[Morrison Formation]] in Wyoming at about 150 Ma. ''[[Epidendrosaurus]]'' and ''[[Pedopenna]]'' are known from the [[Daohugou Beds]] in China, whose age is still being debated, but may be about 160&nbsp;Ma or 145&nbsp;Ma.
 
   
 
The wide range of fossils in the late Jurassic and morphological evidence suggests that coelurosaurian differentiation was virtually complete before the end of the Jurassic.
 
The wide range of fossils in the late Jurassic and morphological evidence suggests that coelurosaurian differentiation was virtually complete before the end of the Jurassic.
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There is consensus among paleontologists that birds are the descendants of coelurosaurs. Under modern [[cladistics|cladistic]] definitions, birds are considered the only living lineage of coelurosaurs. Birds are classified by most paleontologists as belonging to the subgroup [[Maniraptora]].<ref name=KP04 />
 
There is consensus among paleontologists that birds are the descendants of coelurosaurs. Under modern [[cladistics|cladistic]] definitions, birds are considered the only living lineage of coelurosaurs. Birds are classified by most paleontologists as belonging to the subgroup [[Maniraptora]].<ref name=KP04 />
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  +
A portion of a tail belonging to a juvenile coelurosaur was found in 2015, inside of a piece of amber.
   
 
==Classification==
 
==Classification==
The [[phylogeny]] and [[taxonomy]] of Coelurosauria has been subject to intensive research and revision. For many years, Coelurosauria was a 'dumping ground' for all small theropods. In the 1960s several distinctive lineages of coelurosaurs were recognized, and a number of new infraorders were erected, including the [[Ornithomimosauria]], [[Deinonychosauria]], and [[Oviraptorosauria]]. During the 1980s and 1990s, paleontologists began to give Coelurosauria a formal definition, usually as all animals closer to birds than to ''[[Allosaurus]]'', or equivalent specifiers. Under this modern definition, many small theropods are not classified as coelurosaurs at all and some large theropods, such as the [[Tyrannosauridae|tyrannosaurids]], were actually more advanced than allosaurs and therefore were reclassified as giant coelurosaurs. Even more drastically, the [[Therizinosaur|segnosaurs]], once not even regarded as theropods, have turned out to be non-carnivorous coelurosaurs related to ''[[Therizinosaurus]]''. Senter (2007) listed 59 different published phylogenies since 1984. Those since 2005 have followed almost the same pattern, and differ significantly from many older phylogenies.
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The [[phylogeny]] and [[Taxonomy (biology)|taxonomy]] of Coelurosauria has been subject to intensive research and revision. For many years, Coelurosauria was a 'dumping ground' for all small theropods. In the 1960s several distinctive lineages of coelurosaurs were recognized, and a number of new infraorders were erected, including the [[Ornithomimosauria]], [[Deinonychosauria]], and [[Oviraptorosauria]]. During the 1980s and 1990s, paleontologists began to give Coelurosauria a formal definition, usually as all animals closer to birds than to ''[[Allosaurus]]'', or equivalent specifiers. Under this modern definition, many small theropods are not classified as coelurosaurs at all and some large theropods, such as the [[Tyrannosauridae|tyrannosaurids]], were actually more advanced than allosaurs and therefore were reclassified as giant coelurosaurs. Even more drastically, the [[Therizinosaur|segnosaurs]], once not even regarded as theropods, have turned out to be non-carnivorous coelurosaurs related to ''[[Therizinosaurus]]''. Senter (2007) listed 59 different published phylogenies since 1984. Those since 2005 have followed almost the same pattern, and differ significantly from many older phylogenies.
   
  +
The following family tree illustrates a synthesis of the relationships of the major coelurosaurian groups based on various studies conducted in the 2010s.<ref name=theropodphylogeny2015>Hendrickx, C., Hartman, S.A., & Mateus, O. (2015). An Overview of Non- Avian Theropod Discoveries and Classification. ''PalArch’s Journal of Vertebrate Palaeontology'', '''12'''(1): 1-73.</ref>
The following phylogenetic results are taken from Senter (2007) "A new look at the Phylogeny of Coelurosauria (Dinosauria: Theropoda)."<ref name="senter2007" /> This cladogram does not represent time, but a crude estimate of the time can be inferred from morphological changes. The first coelurosaurs were similar to the [[Coeluridae|coelurids]] ''[[Tanycolagreus]]'' and ''[[Coelurus]]'', and differed only sightly from other early [[Tyrannosauroidea|tyrannosauroids]] ''[[Dilong (dinosaur)|Dilong]]'' and ''[[Eotyrannus]]''. The two most significant separations between subgroups are those between the [[Paraves]] and other coelurosaurs and between the paravian clades [[Avialae]] and [[Deinonychosauria]].
 
   
 
{{clade| style=font-size:100%;line-height:80%
{|width="80%" border="0" cellpadding="2" cellspacing="0"
 
 
|label1='''Coelurosauria'''
|-
 
 
|1={{clade
|
 
  +
|1=†''[[Bicentenaria]]''
{{Clade| style=font-size:80%;line-height:80%
 
  +
|2={{clade
|label1='''1'''
 
  +
|1=†''[[Zuolong]]''
|1={{Clade
 
|label1='''2''' |1={{Clade
+
|label2='''Tyrannoraptora'''
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|2={{clade
|label1='''3''' |1={{Clade |1=†''[[Tanycolagreus]]'' |2=†''[[Coelurus]]''}}
 
  +
|1=†[[Tyrannosauroidea]][[File:Rjpalmer tyrannosaurusrex (white background).jpg|130 px]]
|2={{Clade |1=†''[[Dilong (dinosaur)|Dilong]]'' |2={{Clade |1=†''[[Eotyrannus]]'' |2=†''[[Tyrannosaurus]]'' }} }} }} |2={{Clade
 
  +
|label2={{color|white|unnamed}}
|label1='''4''' |1={{Clade |1=†''[[Compsognathus]]'' |2=†''[[Sinosauropteryx]]''}}
 
|label2='''5''' |2={{Clade
+
|2={{clade
  +
|1=†''[[Aorun]]''
|label1='''6''' |1={{Clade |1=†''[[Harpymimus]]'' |2={{Clade |1=†''[[Pelecanimimus]]'' |2=†''[[Ornithomimus]]'' }} }}
 
  +
|2={{clade
|label2='''7''' |2={{Clade |1=†''[[Ornitholestes]]'' |2={{Clade
 
|label1='''8''' |1={{Clade |1=†''[[Falcarius]]'' |2={{Clade |1=†''[[Alxasaurus]]''
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|1=†''[[Scipionyx]]''
  +
|2={{clade
|label2='''9''' |2={{Clade |1=†''[[Segnosaurus]]'' |2=†''[[Therizinosaurus]]''
 
 
|1=†''[[Ornitholestes]]
}} }} }}
 
|2={{Clade
+
|2={{clade
  +
|1=†[[Compsognathidae]]
|label1='''10''' |1={{Clade |1=†''[[Alvarezsaurus]]'' |2=†''[[Shuvuuia]]'' }}
 
  +
|label2=[[Maniraptoriformes]]
|2='''11'''-'''14'''
 
  +
|2={{clade
}} }} }} }} }} }} }}
 
  +
|1=†[[Ornithomimosauria]][[File:Hypothetical Deinocheirus (flipped).jpg|120 px]]
|
 
  +
|2=[[Maniraptora]][[File:Deinonychus ewilloughby (flipped).png|80px]]
{{Clade| style=font-size:80%;line-height:80%
 
 
}} }} }} }} }} }} }} }} }}
|1={{Clade
 
|label1='''11''' |1={{Clade |1=†''[[Protarchaeopteryx]]'' |2={{Clade |1=†''[[Avimimus]]'' |2={{Clade |1=†''[[Caudipteryx]]''
 
|label2='''12''' |2={{Clade |1=†''[[Oviraptor]]'' |2={{Clade |1=†''[[Citipati]]''
 
|label2='''13''' |2={{Clade |1=†''[[Ingenia]]''}}
 
}} }} }} }} }}
 
|label2='''14''' |2={{Clade
 
|label1='''15''' |1={{Clade |1=†''[[Epidendrosaurus]]''
 
|label2='''16''' |2={{Clade |1=†''[[Archaeopteryx]]''
 
|label2='''17''' |2={{Clade |1=†''[[Jeholornis]]'' |2={{Clade |1=†''[[Sapeornis]]'' |2={{Clade |1=†''[[Confuciusornis]]''
 
|2={{Clade |1=†[[Enantiornithes]] |2={{Clade |1=†''[[Yanornis]]'' |2={{Clade |1=†''[[Hesperornis]]'' |2=[[Fossil birds|Modern Birds]]}} }} }}
 
}} }} }} }} }}
 
|label2='''18''' |2={{Clade
 
|label1='''19''' |1={{Clade |1=†''[[Sinovenator]]'' |2={{Clade |1=†''[[Mei (dinosaur)|Mei]]'' |2=†''[[Troodon]]'' }} }}
 
|label2='''20''' |2={{Clade
 
|label1='''21''' |1={{Clade |1=†''[[Unenlagia]]''}} |2={{Clade
 
|label1='''22''' |1={{Clade |1=†''[[Bambiraptor]]'' |2=†''[[Microraptor]]'' }} |2={{Clade
 
|1=†''[[Velociraptor]]'' |2={{Clade |1=†''[[Deinonychus]]''
 
|label2='''23''' |2={{Clade |1=†''[[Dromaeosaurus]]'' }}
 
}} }} }} }} }} }} }} }}
 
|}
 
<small>
 
'''1''' Coelurosauria, '''2''' †[[Tyrannosauroidea]], '''3''' †[[Coeluridae]], '''4''' †[[Compsognathidae]], '''5''' [[Maniraptoriformes]], '''6''' †[[Ornithomimosauria]], '''7''' [[Maniraptora]], '''8''' †[[Therizinosauroidea]], '''9''' †[[Therizinosauridae]], '''10''' †[[Alvarezsauridae]], '''11''' †[[Oviraptorosauria]], '''12''' †[[Oviraptoridae]], '''13''' †[[Ingeniinae]], '''14''' [[Paraves]], '''15''' [[Avialae]], '''16''' [[Aves]], '''17''' [[Ornithurae]], '''18''' †[[Deinonychosauria]], '''19''' †[[Troodontidae]], '''20''' †[[Dromaeosauridae]], '''21''' †[[Unenlagiinae]], '''22''' †[[Microraptoria]], '''23''' †[[Dromaeosaurinae]]</small>
 
   
 
==="Coelurosaurus"===
 
==="Coelurosaurus"===
"Coelurosaurus" is an [[nomen nudum|informal]] [[genus|generic]] name, attributed to [[Friedrich von Huene]], 1929, that is sometimes seen in lists of dinosaurs. The name is undescribed and has not been used seriously, although it has appeared in works of fiction.
+
"Coelurosaurus" is an [[nomen nudum|informal]] [[genus|generic]] name, attributed to [[Friedrich von Huene]], 1929, that is sometimes seen in lists of dinosaurs. It probably arose as a [[typo|typographical error]]; von Huene intended to assign indeterminate remains to Coelurosauria ''[[incertae sedis]]'', but at some point in the process of publication a revision to the text made it appear that he was creating a new generic name "Coelurosaurus" (as described by [[George Olshevsky]] in a 1999 post to the Dinosaur Mailing List). The name is undescribed and has not been used seriously, although it has appeared in works of fiction.
   
 
==See also==
 
==See also==
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*[[List of fossil birds]]
 
*[[List of fossil birds]]
   
==Footnotes==
+
==References==
{{Reflist|3|refs=
+
{{Reflist|2|refs=
<ref name="currie-2005">Currie (2005) Page 368.</ref>
+
<ref name="currie-2005">Currie (2005) p. 368.</ref>
 
<ref name="csaharicus-endo-abs-19">"Abstract," Larsson (2001). Page 19.</ref>
 
<ref name="csaharicus-endo-abs-19">"Abstract," Larsson (2001). Page 19.</ref>
 
<ref name=KP04>Padian (2004). Basal Avialae. Pages 210–231.</ref>}}
<ref name="Dinodata">[http://www.dinodata.org/index.php?option=com_content&task=view&id=1017&Itemid=104 '''Dinodata:''' Coelurosauria].</ref>
 
<ref name="mayeretal2005">Mayr, et al. (2005). Pages 1483-1486.</ref>
 
<ref name=KP04>Padian (2004). Basal Avialae. Pages 210–231.</ref>
 
<ref name="senter2007">Senter (2007).</ref>
 
<ref name="zannoetal2009">Zanno, et al. (2009).</ref>
 
}}
 
   
 
==References==
 
==References==
 
* {{cite book |last=Currie|first=Philip J.|authorlink=Philip J. Currie |year=2005 |title=Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed |publisher=[[Indiana University Press]]|isbn=0-253-34595-2 |page=368}}
 
* {{cite book |last=Currie|first=Philip J.|authorlink=Philip J. Currie |year=2005 |title=Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed |publisher=[[Indiana University Press]]|isbn=0-253-34595-2 |page=368}}
 
* Larsson, H.C.E. 2001. Endocranial anatomy of ''Carcharodontosaurus saharicus'' (Theropoda: Allosauroidea) and its implications for theropod brain evolution. pp.&nbsp;19–33. In: ''Mesozoic Vertebrate Life''. Ed.s Tanke, D. H., Carpenter, K., Skrepnick, M. W. Indiana University Press.
* [http://www.dinodata.org/index.php?option=com_content&task=view&id=1017&Itemid=104 '''Dinodata:''' Coelurosauria].
 
* Larsson, H.C.E. 2001. Endocranial anatomy of ''Carcharodontosaurus saharicus'' (Theropoda: Allosauroidea) and its implications for theropod brain evolution. pp.&nbsp;19–33. In: ''Mesozioc Vertebrate Life''. Ed.s Tanke, D. H., Carpenter, K., Skrepnick, M. W. Indiana University Press.
 
 
* Mayr, G., B. Pohl & D.S. Peters (2005). "A well-preserved ''Archaeopteryx'' specimen with theropod features". ''Science'', '''310'''(5753): 1483-1486.
 
* Mayr, G., B. Pohl & D.S. Peters (2005). "A well-preserved ''Archaeopteryx'' specimen with theropod features". ''Science'', '''310'''(5753): 1483-1486.
  +
* {{cite web | author = George Olshevsky | title = Re: What are these dinosaurs | url= http://dml.cmnh.org/1999Nov/msg00507.html | accessdate = 2007-01-29 }} (on "Coelurosaurus")
* Padian, K. (2004). Basal Avialae. In: Weishampel, D.B., Dodson, P., and Osmólska, H. (eds.). ''The Dinosauria'' (second edition). University of California Press, Berkeley, 210–231. ISBN 0-520-24209-2.
+
** Padian, K. (2004). Basal Avialae. In: Weishampel, D.B., Dodson, P., and Osmólska, H. (eds.). ''The Dinosauria'' (second edition). University of California Press, Berkeley, 210–231. ISBN 0-520-24209-2.
 
* Senter, P. (2007). "A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda)." ''Journal of Systematic Palaeontology'', ({{doi|10.1017/S1477201907002143}}).
 
* Senter, P. (2007). "A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda)." ''Journal of Systematic Palaeontology'', ({{doi|10.1017/S1477201907002143}}).
* Zanno, L.E., Gillette, D.D., Albright, L.B., and Titus, A.L. (2009). "A new North American therizinosaurid and the role of herbivory in 'predatory' dinosaur evolution." ''Proceedings of the Royal Society B'', Published online before print July 15, 2009, doi: 10.1098/rspb.2009.1029.
+
* Zanno, L.E., Gillette, D.D., Albright, L.B., and Titus, A.L. (2009). "A new North American therizinosaurid and the role of herbivory in 'predatory' dinosaur evolution." ''Proceedings of the Royal Society B'', Published online before print July 15, 2009, {{doi|10.1098/rspb.2009.1029}}.
  +
* http://news.nationalgeographic.com/2016/12/feathered-dinosaur-tail-amber-theropod-myanmar-burma-cretaceous/
   
 
==External links==
 
==External links==
  +
{{Wiktionary|Coelurosauria}}
 
* [http://paleodb.org/cgi-bin/bridge.pl?action=checkTaxonInfo&taxon_no=53940 '''Paleobiology Database:''' Coelurosauria]
 
* [http://paleodb.org/cgi-bin/bridge.pl?action=checkTaxonInfo&taxon_no=53940 '''Paleobiology Database:''' Coelurosauria]
 
* [http://www.ucmp.berkeley.edu/diapsids/saurischia/coelurosauria.html The Major Groups of Coelurosaurs]
 
* [http://www.ucmp.berkeley.edu/diapsids/saurischia/coelurosauria.html The Major Groups of Coelurosaurs]
* [http://www.thescelosaurus.com/coelurosauria.htm '''Thescelosaurus:''' Coelurosaurs]
+
* [http://www.palaeos.com/Vertebrates/Units/340Theropoda/340.500.html Palaeos: Coelurosauria]
* [http://www.palaeos.com/Vertebrates/Units/340Theropoda/340.500.html '''Palaeos:''' Coelurosauria]
+
* [http://tolweb.org/Coelurosauria/15769 Tree of Life Web: Coelurosauria]
* [http://tolweb.org/Coelurosauria/15769 '''Tree of Life Web:''' Coelurosauria]
 
* [http://www.dinodata.org/index.php?option=com_content&task=view&id=1017&Itemid=104 '''Dinodata:''' Coelurosauria]
 
   
 
{{Project Dinosaur Taxonomy}}
 
[[Category:Coelurosaurs| ]]
 
[[Category:Coelurosaurs| ]]
  +
[[Category:Late Jurassic dinosaurs]]
 
  +
[[Category:Cretaceous dinosaurs]]
{{Project Dinosaur Taxonomy}}
 
  +
[[Category:Cenozoic dinosaurs]]
  +
[[Category:Extant Late Jurassic first appearances]]

Latest revision as of 14:29, 27 April 2017

Coelurosaurians
Temporal range:
Late JurassicPresent, 165–0 Ma
PreЄ
Є
Є
O
O
S
S
D
D
C
C
P
P
T
T
J
J
Pg
Pg
N
N
Possible Early Jurassic record
File:Zuni Coelurosaur.jpg
Reconstructed coelurosaur skeleton, Wyoming Dinosaur Center
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauria
Clade: Avetheropoda
Clade: Coelurosauria
von Huene, 1914
Subgroups[1]

Coelurosauria ( /sɪˌljʊərəˈs[unsupported input]iə/;Template:RefnTemplate:Refn from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs.

Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyrannosaurs, ornithomimosaurs, and maniraptorans; Maniraptora includes birds, the only dinosaur group alive today.[4]

Most feathered dinosaurs discovered so far have been coelurosaurs. Philip J. Currie considers it probable that all coelurosaurs were feathered.[5] In the past, Coelurosauria was used to refer to all small theropods, although this classification has been abolished.

Anatomy

Bodyplan

The studying of anatomical traits in coelurosaurs indicates that the last common ancestor had evolved the ability to eat and digest plant matter, adapting to an omnivorous diet, an ability that could be a major contributor to the clade's success. Later groups would hold on to the omnivory, while others specialized in various directions, becoming insectivorous (Alvarezsauridae), herbivorous (Therizinosauridae) and carnivorous (Tyrannosauroidea and Dromaeosauridae).[6] The group includes some of the largest (Tyrannosaurus) and smallest (Microraptor, Parvicursor) carnivorous dinosaurs ever discovered. Characteristics that distinguish coelurosaurs include:

  • a sacrum (series of vertebrae that attach to the hips) longer than in other dinosaurs
  • a tail stiffened towards the tip
  • a bowed ulna (lower arm bone).
  • a tibia (lower leg bone) that is longer than the femur (upper leg bone)

Integument

Main article: Feathered dinosaurs

Fossil evidence shows that the skin of even the most primitive coelurosaurs was covered primarily in feathers. Fossil traces of feathers, though rare, have been found in members of most major coelurosaurian lineages. Most coelurosaurs also retained scales and scutes on some portion of their bodies, particularly the feet, though some primitive coelurosaurian species are known to have had scales on the upper legs and portions of the tail as well. These include tyrannosauroids, Juravenator, and Scansoriopteryx. Fossils of at least some of these animals (Scansoriopteryx and possibly Juravenator) also preserve feathers elsewhere on the body.

Though once thought to be a feature exclusive to coelurosaurs, feathers or feather-like structures are also known in some ornithischian dinosaurs (like Tianyulong and Kulindadromeus), and in pterosaurs. Though it is unknown whether these are related to true feathers, recent analysis has suggested that the feather like integument found in ornithischians may have evolved independently of coelurosaurs.[7]

Nervous system and senses

Although rare, complete casts of theropod endocrania are known from fossils. Theropod endocrania can also be reconstructed from preserved braincases without damaging valuable specimens by using a computed tomography scan and 3D reconstruction software. These finds are of evolutionary significance because they help document the emergence of the neurology of modern birds from that of earlier reptiles. An increase in the proportion of the brain occupied by the cerebrum seems to have occurred with the advent of the Coelurosauria and "continued throughout the evolution of maniraptorans and early birds."[8]

Fossil evidence and age

A few fossil traces tentatively associated with the Coelurosauria date back as far as the late Triassic. What has been found between then and the start of the late Jurassic is fragmentary. A typical example is Iliosuchus, known only from two ilia bones in the mid Jurassic. It was a 1.5 m long carnivore from about 165 Ma (million years ago) in Oxfordshire and is tentatively assigned to the Tyrannosauroidea.

Many nearly complete fossil coelurosaurians are known from the late Jurassic. Archaeopteryx (incl. Wellnhoferia) is known from Bavaria at 155-150 Ma. Ornitholestes, the troodontid WDC DML 001, Coelurus fragilis and Tanycolagreus topwilsoni are all known from the Morrison Formation in Wyoming at about 150 Ma. Epidendrosaurus and Pedopenna are known from the Daohugou Beds in China, whose age is still being debated, but may be about 160 Ma or 145 Ma.

The wide range of fossils in the late Jurassic and morphological evidence suggests that coelurosaurian differentiation was virtually complete before the end of the Jurassic.

In the early Cretaceous, a superb range of coelurosaurian fossils (including avians) are known from the Yixian Formation in Liaoning. All known theropod dinosaurs from the Yixian Formation are coelurosaurs. Many of the coelurosaurian lineages survived to the end of the Cretaceous period (about 65 Ma) and fossils of some lineages, such as the Tyrannosauroidea, are best known from the late Cretaceous. A majority of coelurosaur groups became extinct in the Cretaceous–Paleogene extinction event, including the Tyrannosauroidea, Ornithomimosauria, Oviraptorosauria, Deinonychosauria, Enantiornithes, and Hesperornithes. Only the Neornithes (modern birds) survived, and continued to diversify after the extinction of the other dinosaurs into the numerous forms found today.

There is consensus among paleontologists that birds are the descendants of coelurosaurs. Under modern cladistic definitions, birds are considered the only living lineage of coelurosaurs. Birds are classified by most paleontologists as belonging to the subgroup Maniraptora.[9]

A portion of a tail belonging to a juvenile coelurosaur was found in 2015, inside of a piece of amber.

Classification

The phylogeny and taxonomy of Coelurosauria has been subject to intensive research and revision. For many years, Coelurosauria was a 'dumping ground' for all small theropods. In the 1960s several distinctive lineages of coelurosaurs were recognized, and a number of new infraorders were erected, including the Ornithomimosauria, Deinonychosauria, and Oviraptorosauria. During the 1980s and 1990s, paleontologists began to give Coelurosauria a formal definition, usually as all animals closer to birds than to Allosaurus, or equivalent specifiers. Under this modern definition, many small theropods are not classified as coelurosaurs at all and some large theropods, such as the tyrannosaurids, were actually more advanced than allosaurs and therefore were reclassified as giant coelurosaurs. Even more drastically, the segnosaurs, once not even regarded as theropods, have turned out to be non-carnivorous coelurosaurs related to Therizinosaurus. Senter (2007) listed 59 different published phylogenies since 1984. Those since 2005 have followed almost the same pattern, and differ significantly from many older phylogenies.

The following family tree illustrates a synthesis of the relationships of the major coelurosaurian groups based on various studies conducted in the 2010s.[10]

Coelurosauria

Bicentenaria




Zuolong


Tyrannoraptora

TyrannosauroideaFile:Rjpalmer tyrannosaurusrex (white background).jpg


unnamed

Aorun




Scipionyx




Ornitholestes




Compsognathidae


Maniraptoriformes

OrnithomimosauriaFile:Hypothetical Deinocheirus (flipped).jpg



ManiraptoraFile:Deinonychus ewilloughby (flipped).png










"Coelurosaurus"

"Coelurosaurus" is an informal generic name, attributed to Friedrich von Huene, 1929, that is sometimes seen in lists of dinosaurs. It probably arose as a typographical error; von Huene intended to assign indeterminate remains to Coelurosauria incertae sedis, but at some point in the process of publication a revision to the text made it appear that he was creating a new generic name "Coelurosaurus" (as described by George Olshevsky in a 1999 post to the Dinosaur Mailing List). The name is undescribed and has not been used seriously, although it has appeared in works of fiction.

See also

Portal icon Dinosaurs portal

References

  1. ^ a b c d e f g h Holtz, Thomas R. Jr. (2012) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2011 Appendix.
  2. ^ Carrano, M.T., Benson, R.B.J., and Sampson, S.D. 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology, 10(2): 211-300.
  3. ^ Godefroit, Pascal; Cau, Andrea; Hu, Dong-Yu; Escuillié, François; Wu, Wenhao; Dyke, Gareth (2013). "A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds". Nature. 498 (7454): 359–362. Bibcode:2013Natur.498..359G. PMID 23719374. doi:10.1038/nature12168. 
  4. ^ Turner, A.H., Makovicky, P.J., and Norell, M.A. 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History 371: 1–206.
  5. ^ Currie (2005) p. 368.
  6. ^ "Meat-eating dinosaurs not so carnivorous after all". ScienceDaily. 
  7. ^ Barrett, Paul M. (3 June 2015). "Evolution of Dinosaur Epidermal Structures". The Royal Society Biology Letters. 
  8. ^ "Abstract," Larsson (2001). Page 19.
  9. ^ Padian (2004). Basal Avialae. Pages 210–231.
  10. ^ Hendrickx, C., Hartman, S.A., & Mateus, O. (2015). An Overview of Non- Avian Theropod Discoveries and Classification. PalArch’s Journal of Vertebrate Palaeontology, 12(1): 1-73.

References

  • Currie, Philip J. (2005). Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed. Indiana University Press. p. 368. ISBN 0-253-34595-2. 
  • Larsson, H.C.E. 2001. Endocranial anatomy of Carcharodontosaurus saharicus (Theropoda: Allosauroidea) and its implications for theropod brain evolution. pp. 19–33. In: Mesozoic Vertebrate Life. Ed.s Tanke, D. H., Carpenter, K., Skrepnick, M. W. Indiana University Press.
  • Mayr, G., B. Pohl & D.S. Peters (2005). "A well-preserved Archaeopteryx specimen with theropod features". Science, 310(5753): 1483-1486.
  • George Olshevsky. "Re: What are these dinosaurs". Retrieved 2007-01-29.  (on "Coelurosaurus")
    • Padian, K. (2004). Basal Avialae. In: Weishampel, D.B., Dodson, P., and Osmólska, H. (eds.). The Dinosauria (second edition). University of California Press, Berkeley, 210–231. ISBN 0-520-24209-2.
  • Senter, P. (2007). "A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda)." Journal of Systematic Palaeontology, (doi:10.1017/S1477201907002143).
  • Zanno, L.E., Gillette, D.D., Albright, L.B., and Titus, A.L. (2009). "A new North American therizinosaurid and the role of herbivory in 'predatory' dinosaur evolution." Proceedings of the Royal Society B, Published online before print July 15, 2009, doi:10.1098/rspb.2009.1029.
  • http://news.nationalgeographic.com/2016/12/feathered-dinosaur-tail-amber-theropod-myanmar-burma-cretaceous/

External links

Various dinosaurs This article is part of Project Dinosaur Taxonomy, a All Birds project that aims to write comprehensive articles on every order, family and other taxonomic rank related to dinosaurs.