Wikipedia - User contributions [en]

Oligoryzomys victus

2022-03-27T21:16:35Z

Wikkler:

{{Short description|Extinct species of rodent}}
{{speciesbox
| status = EX
| status_system = IUCN3.1
| extinct = c. 1892
| status_ref = <ref name="iucn status 14 November 2021">{{cite iucn |author=Turvey, S.T. |author2=Dávalos, L. |date=2019 |title=''Oligoryzomys victus'' |volume=2019 |page=e.T15255A22357957 |doi=10.2305/IUCN.UK.2019-1.RLTS.T15255A22357957.en |access-date=14 November 2021}}</ref>
| genus = Oligoryzomys
| species = victus
| authority = ([[Oldfield Thomas|Thomas]], 1898)
| synonyms =
*''Oryzomys victus'' Thomas, 1898
}}
'''''Oligoryzomys victus''''', also known as the '''St. Vincent colilargo'''<ref>Musser and Carleton, 2005</ref> or '''St. Vincent pygmy rice rat''',<ref name="iucn status 14 November 2021" /> is a species of [[rodent]] in the genus ''[[Oligoryzomys]]'' of the [[oryzomyini|oryzomyine]] tribe. Only one specimen is known, which was collected on [[Saint Vincent (island)|Saint Vincent]] in the [[Lesser Antilles]] in about 1892, and it is now presumed extinct.

==Taxonomy==
The only known specimen was collected by H. H. Smith at an unknown location on Saint Vincent and later presented to the [[British Museum of Natural History]], where it was registered as specimen BMNH 97.12.26.1.<ref>Ray, 1962, p. 48</ref> In 1898, [[Oldfield Thomas]] described the specimen as the [[holotype]] of a new species of ''[[Oryzomys]]'' which he named ''Oryzomys victus''. Although Thomas placed it close to species now placed in ''[[Oligoryzomys]]'', later compilators considered the affinities of ''O. victus'' as unknown; one study placed it in the ''Oryzomys tectus'' group (more or less = ''[[Oecomys]]''). In his 1962 study of Antillean oryzomyines, however, American paleontologist Clayton Ray reaffirmed its affinities with ''Oligoryzomys'', but he was unable to resolve its relation within the genus. On the one hand, he saw closer morphological similarities to small ''Oligoryzomys'' such as ''[[Oligoryzomys fulvescens|O. fulvescens]]'' and ''[[Oligoryzomys delicatus|O. delicatus]]'', but on the other hand larger species such as ''[[Oligoryzomys longicaudatus|O. longicaudatus]]'' are closer in size. Ray also considered the possibility that the St. Vincent population was in fact introduced from a still unknown mainland species, but considered this unlikely; no such species has since been found.<ref>Ray, 1962, pp. 57–61</ref>

==Description==
The holotype, a fluid-preserved adult female with the skull extracted, was described in Thomas' original description and in Ray's 1962 restudy. It is a relatively large ''Oligoryzomys'' and has the pelage dark reddish above and buffy white below. The short ears are brown in color. The tail nearly lacks hairs and is brown above and somewhat paler below. Like most oryzomyines, it has eight [[mammae]], including a pectoral pair.<ref>Thomas, 1898, p. 178, quoted in Ray, 1962, p. 51</ref> The head and body length is 96 mm, the tail length 121 mm, the hind foot length (without claws) 25 mm, the ear length 14 mm, the greatest length of the skull 27.2 mm, the length of the upper molars is 3.9 mm, and the length of the lower molars is 4.0 mm.<ref>Ray, 1962, table 6</ref>

==Behaviour==
Little is known about the habits or the ecology of ''O. victus''; in fact, the only direct information is a collector's note which calls it a "forest rat". Its morphology suggests that it was not [[arboreal]] or [[fossorial]].<ref>Ray, 1962, pp. 56–57</ref>

==Extinction==
The extinction of ''O. victus'' may be associated with the introduction of the [[small Asian mongoose]] to St. Vincent in the 1870s. Ray suggests that the rice rat was an easier prey for the mongoose than introduced ''[[Rattus]]'', which may have become partly arboreal, thus creating a competitive disadvantage for the rice rat. Any remaining populations may have been wiped out when the [[Soufrière (volcano)|Soufrière]] erupted in 1902, destroying the pristine vegetation on its slopes.<ref>Ray, 1962, pp. 48–50</ref>

==References==
{{Reflist}}

==Literature cited==
*{{MSW3 Muroidea | id = 13000783 | page = 1144}}
*Ray, C.E. 1962. The Oryzomyine Rodents of the Antillean Subregion. Doctor of Philosophy thesis, Harvard University, 211 pp.

{{Oryzomyini nav}}
{{Taxonbar|from=Q1761020}}

[[Category:Oligoryzomys]]
[[Category:Extinct rodents]]
[[Category:Endemic fauna of Saint Vincent and the Grenadines]]
[[Category:Fauna of Saint Vincent and the Grenadines]]
[[Category:Taxa named by Oldfield Thomas]]
[[Category:Mammal extinctions since 1500]]
[[Category:Taxonomy articles created by Polbot]]
[[Category:Mammals of the Caribbean]]

Dorygnathus purdoni

2018-08-09T01:37:16Z

Wikkler: ←Redirected page to Parapsicephalus

#REDIRECT [[Parapsicephalus]]

Megaconus

2018-02-20T23:09:13Z

Wikkler: non-generic names are not italicized

{{Automatic taxobox
| fossil_range = [[Middle Jurassic]], {{fossil range|165}}
| taxon = Megaconus
| authority = Zhou ''et al.'', 2013
| type_species = {{extinct}}'''''Megaconus mammaliaformis'''''
| type_species_authority = Zhou ''et al.'', 2013
}}
'''''Megaconus''''' is an extinct [[genus]] of [[mammaliaforms]] from the [[Middle Jurassic]] [[Tiaojishan Formation]] of [[Inner Mongolia]], China. The [[type species|type]] and only [[species]], '''''Megaconus mammaliaformis''''' (nicknamed '''Jurassic Squirrel'''),<ref>http://news.msn.com/science-technology/jurassic-squirrel-fossil-hair-came-before-mammals</ref> was first described in the journal ''[[Nature (journal)|Nature]]'' in 2013. ''Megaconus'' is thought to have been a [[herbivore]] that lived on the ground, having a similar posture to modern-day [[armadillo]]s and [[rock hyrax]]es. ''Megaconus'' is a member of a group called [[Haramiyida]]. A [[phylogenetic]] analysis published along with its first description showed that haramiyidans originated before the appearance of [[true mammals]], but an accompanying description of the haramiyidan ''[[Arboroharamiya]]'' in the same issue of ''Nature'' indicated that haramyidans were true mammals.<ref name=ZWML13>{{Cite journal | last1 = Zhou | first1 = C. F. | last2 = Wu | first2 = S. | last3 = Martin | first3 = T. | last4 = Luo | first4 = Z. X. | title = A Jurassic mammaliaform and the earliest mammalian evolutionary adaptations | doi = 10.1038/nature12429 | journal = [[Nature (journal)|Nature]] | volume = 500 | issue = 7461 | pages = 163 | year = 2013 | pmid = 23925238| pmc = }}</ref> If haramiyidans are not mammals, ''Megaconus'' would be one of the most [[basal (phylogenetics)|basal]] ("primitive") mammaliaforms to possess fur, and an indicator that fur evolved in the ancestors of mammals and not the mammals themselves.<ref>{{cite news | title = Jurassic squirrel’s secret is out | newspaper = The Hindu | date = August 9, 2013 | url = http://www.thehindu.com/opinion/op-ed/jurassic-squirrels-secret-is-out/article5004252.ece}}</ref>

==Description==
''Megaconus'' is one of the few early mammaliaforms known from a complete skeleton. The skeleton includes both the jaw bones and the teeth, which are the most informative features because they allow for comparisons with other mammaliaforms known only from dental features. ''Megaconus'' has a [[dentition]] similar to those of [[rodent]]s, with large [[incisor]]s at the front of the jaws and broad [[molar (tooth)|molar]]s in the back. One distinguishing feature of ''Megaconus'' is a pair of enlarged [[premolar]] teeth in the lower jaw. The teeth of ''Megaconus'' have many [[cusp (anatomy)|cusp]]s, allowing them to interlock tightly when the jaws are closed. If ''Megaconus'' is a non-mammalian mammaliaform, it is one of the most basal mammaliaforms to possess such complex teeth.<ref name=ZWML13/>

The [[middle ear]] of ''Megaconus'' is more primitive than that of modern mammals. The three bones that make up the middle ear in modern mammals — the malleus, incus, and stapes — originated from the lower jaw in the ancestors of mammals. In ''Megaconus'', these bones are still associated with the lower jaw, placed within a groove behind the [[dentary bone]] of the lower jaw.<ref name=ZWML13/>

''Megaconus'' is estimated to have weighed about {{convert|250|g|oz}}. It probably had an outwardly similar appearance to [[multituberculate]]s, a major group of [[Mesozoic]] mammals. However, its body is longer than those of multituberculates and most other Mesozoic mammaliaforms, having more back [[vertebrae]] (24) than other early mammals. The transition between the [[thoracic vertebrae|thoracic]] and [[lumbar vertebrae]] is more gradual in ''Megaconus'' than it is in multituberculates, and the boundary between the anterior and posterior [[Epaxial and hypaxial muscles|epaxial muscle]]s (the muscles that cover the front part and the back part of the back, respectively) is positioned farther back.<ref name=ZWML13/>

''Megaconus'' is inferred to have been [[ambulatory]], meaning that it walked on the ground. Its claws are short, meaning that they were not suitable for digging, and only slightly curved, meaning that they were not suitable for climbing. The lack of an elongated [[calcaneus]] or "heel" on the ankle means that it could not have run fast. However, ''Megaconus'' differs from most ambulatory mammals in having a fused [[tibia]] and [[fibula]] in its lower leg, a feature that is normally seen in jumping or [[bipedal]] mammals. Among the only living ambulatory mammals to have a fused tibia and fibula are the armadillo and the [[aardvark]].<ref name=ZWML13/>

Remains of fur are preserved on parts of the skeleton. The fur consists of dark [[guard hair]]s and a lighter layer of [[underfur]]. Fur seems to be absent on the underside of the [[abdomen]]. A [[keratinous]] spur on the rear leg may have been used to deliver poison, like the spur of the living [[platypus]].<ref name=ZWML13/>

==Discovery==
The [[holotype]] skeleton of ''Megaconus'' was discovered in the [[Daohugou Beds]] of the Tiaojishan Formation in Inner Mongolia, China. The layer in which ''Megaconus'' was found is about 165 million years old ([[Ma (unit)|Ma]]). In addition to ''Megaconus'', several other mammaliaforms are known from the Daohugou Beds: the non-mammalian mammaliaform ''[[Castorocauda]]'', the basal mammals ''[[Volaticotherium]]'' and ''[[Pseudotribos]]'', and the [[Eutheria|placental]] mammal ''[[Juramaia]]''.

The genus name ''Megaconus'' means "large cusp", coming from the [[Latin]] ''mega'' ("large") and the [[Greek language|Greek]] ''conus'' ("cusp"), in reference to the pair of large premolars in its lower jaw. The species name ''mammaliaformis'' references its ancestral mammalian features.<ref name=ZWML13/>

==Relationships==
''Megaconus'' is part of the [[clade]] Haramiyida, which is otherwise known almost exclusively from teeth. The relationships of haramiyidans to other early mammals is contested.

One idea is that haramiyidans are close relatives of Multituberculata, the most diverse group of Mesozoic mammals, and that both are part of the larger clade [[Allotheria]]. According to this evolutionary hypothesis ([[phylogeny]]), haramiyidans fall within the [[crown group]] Mammalia — the clade originating from the [[most recent common ancestor]] of living mammals. Within the crown group Mammalia, Haramiyida would be a basal offshoot of [[Boreosphenida]], the clade including [[marsupial]]s, [[eutherians]] (placental mammals), and all mammals more closely related to them than to [[monotreme]]s (which are part of the clade [[Australosphenida]]).

A second hypothesis holds that haramiyidans originated before the appearance of crown group Mammalia as more basal members of the larger clade Mammaliaformes and that multituberculates are deeply nested within the crown group, splitting up Allotheria. This phylogeny was supported by the phylogenetic analysis that included ''Megaconus''. Features of ''Megaconus'' that link it with basal mammaliaforms include a groove in the lower jaw holding the middle ear bones (in multituberculates and other true mammals, these bones are completely separated from the lower jaw) and a calcaneum or ankle bone that lacks an enlarged heel. Below is a [[cladogram]] modified from the analysis:<ref name=ZWML13/>

{{clade| style=font-size:85%;line-height:85%
|label1=[[Mammaliaformes]]
|1={{clade
|1=''[[Adelobasileus]]''
|2=''[[Sinoconodon]]''
|3={{clade
|1={{clade
|1=''[[Morganucodon]]''
|2=''[[Megazostrodon]]''}}
|2={{clade
|label1=[[Haramiyida]]
|1={{clade
|1={{clade
|1=''[[Thomasia]]''
|2=''[[Haramiyavia]]''}}
|label2=[[Eleutherodontidae]]
|2={{clade
|1='''''Megaconus'''''
|2={{clade
|1=''[[Eleutherodon]]''
|2=''[[Sineleutherus]]''}} }} }}
|2={{clade
|1=''[[Castorocauda]]''
|2=''[[Haldanodon]]''}}
|3=''[[Hadrocodium]]''
|label4=Crown group [[Mammalia]]
|4={{clade
|1=[[Australosphenida]] ([[monotreme]]s and extinct relatives)
|label2=[[Boreosphenida]]
|2={{clade
|1=[[Eutriconodonta]]
|2={{clade
|1=[[Multituberculata]]
|2={{clade
|1=[[Spalacotheroidea]]
|2=[[Cladotheria]] ([[marsupial]]s, [[placental mammal]]s, and extinct relatives)
}} }} }} }} }} }} }} }}

==References==
{{reflist}}

{{mammals}}

[[Category:Jurassic mammals]]
[[Category:Fossil taxa described in 2013]]
[[Category:Cynodont genera]]
[[Category:Fossils of China]]
[[Category:Paleontology in Inner Mongolia]]

Ankylosauria

2018-02-13T17:51:52Z

Wikkler: Undid revision 824642296 by 71.255.155.181 (talk)

{{Automatic taxobox
| name = Ankylosaurs
| fossil_range = [[Middle Jurassic]]—[[Late Cretaceous]], {{fossilrange|167|66}}
| image = Euoplocephalus-tutus-1.jpg
| image_caption = Mounted skeleton of ''[[Euoplocephalus tutus]]'', [[Senckenberg Museum]]
| authority = [[Henry Fairfield Osborn|Osborn]], 1923
| subdivision_ranks = Subgroups
| subdivision =
* {{extinct}}''[[Kunbarrasaurus]]''
* {{extinct}}''[[Sarcolestes]]''
* {{extinct}}[[Ankylosauridae]]
* {{extinct}}[[Nodosauridae]]
}}
'''Ankylosauria''' is a group of mainly [[herbivorous]] [[dinosaur]]s of the order [[Ornithischia]]. It includes the great majority of dinosaurs with [[Armour (zoology)|armor]] in the form of bony osteoderms. Ankylosaurs were bulky [[quadruped]]s, with short, powerful limbs. They are known to have first appeared in the early [[Jurassic Period]], and persisted until the end of the [[Cretaceous Period]]. They have been found on every continent except [[Africa]]. The first dinosaur ever discovered in [[Antarctica]] was the ankylosaurian ''[[Antarctopelta]]'', fossils of which were recovered from [[Ross Island]] in 1986.

Ankylosauria was first named by [[Henry Fairfield Osborn]] in 1923.<ref name = "osborn1923">Osborn, H. F. (1923). "Two Lower Cretaceous dinosaurs of Mongolia." ''American Museum Novitates'', '''95''': 1–10.[http://hdl.handle.net/2246/3267]</ref> In the [[Linnaean classification]] system, the group is usually considered either a suborder or an infraorder. It is contained within the group [[Thyreophora]], which also includes the [[stegosaur]]s, armored dinosaurs known for their combination of plates and spikes.

==Paleobiology==

They sported a very small brain size in proportion to their body, second only to the [[Saurischian]] [[sauropods]]. They were also rather slow moving, largely because of the shortness of the limbs combined with being incapable of running. Their top speed was likely less than 10 km/hour.

===Armor===
[[File:Edmontonia rugosidens armour AMNH 5381.jpg|thumb|right|Armour of ''Edmontonia'']]
All ankylosaurians had armor over much of their bodies, mostly scutes and nodules, with large spines in some cases. The scutes, or plates, are rectangular to oval objects organized in transverse (side to side) rows, often with keels on the upper surface. Smaller nodules and plates filled in the open spaces between large plates. In all three groups, the first two rows of plates tend to form a sort of half-ring around the neck; in nodosaurids, this comes from adjacent plates fusing with each other (and there is a third row as well), while ankylosaurids usually have the plates fused to the top of another band of bone. The skull has armor plastered on to it, including a distinctive piece on the outside-rear of the lower jaw.

===Diet and feeding===
Ankylosaurs were built low to the ground (typically one foot off the ground surface.) They had small, triangular teeth that were loosely packed, similar to [[stegosaurs]]. The large hyoid bones left in skeletons indicates that they had long, flexible tongues. They also had a large, side secondary palate. This means that they could breathe while chewing, unlike crocodiles. Their expanded gut region suggests the use of fermentation to digest their food, using symbiotic bacteria and gut flora. Their diet likely consisted of [[fern]]s, [[cycads]], and [[angiosperms]]. Mallon et al. (2013) examined herbivore coexistence on the island continent of [[Laramidia]] during the Late Cretaceous. It was concluded that ankylosaurs were generally restricted to feeding on vegetation at, or below, the height of 1 meter.<ref>{{cite journal|last=Mallon|first=Jordan C|author2=David C Evans |author3=Michael J Ryan |author4=Jason S Anderson |title=Feeding height stratification among the herbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta, Canada|journal=BMC Ecology|year=2013|volume=13|doi=10.1186/1472-6785-13-14|pages=14|pmid=23557203|pmc=3637170}}</ref>

===Reproduction===
Possible [[neonate]] sized ankylosaur fossils have been documented in the [[scientific literature]].<ref name="hadro-egg-intro-207">Tanke, D.H. and Brett-Surman, M.K. 2001. Evidence of Hatchling and Nestling-Size Hadrosaurs (Reptilia:Ornithischia) from Dinosaur Provincial Park (Dinosaur Park Formation: Campanian), Alberta, Canada. pp. 206-218. In: Mesozoic Vertebrate Life—New Research Inspired by the Paleontology of Philip J. Currie. Edited by D.H. Tanke and K. Carpenter. Indiana University Press: Bloomington. xviii + 577 pp.</ref>

==Classification==
Ankylosauria is usually split into two [[family (biology)|families]]: [[Nodosauridae]] (the nodosaurids) and [[Ankylosauridae]] (the ankylosaurids). A third family, the [[Polacanthidae]], is sometimes used,<ref name=armor2010>Hayashi, S., Carpenter, K., Scheyer, T.M., Watabe, M. and Suzuki. D. (2010). "Function and evolution of ankylosaur dermal armor." ''Acta Palaeontologica Polonica'', '''55'''(2): 213-228. {{doi|10.4202/app.2009.0103}}</ref> but is more often found to be a sub-group of one of the primary families.<ref name=phylogeny2012>Thompson, R.S., Parish, J.C., Maidment, S.C.R. and Barrett, P.M. (2012). "Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora)." ''Journal of Systematic Palaeontology'', '''10'''(2): 301-312. {{DOI|10.1080/14772019.2011.569091}}</ref>

The first formal definition of Ankylosauria as a [[clade]], a group containing all species of a certain evolutionary branch, was given in 1997 by [[Kenneth Carpenter|Carpenter]].<ref>Carpenter, K., 1997, "Ankylosauria" pp. 16-20 in: P.J. Currie and K. Padian (eds.), ''Encyclopedia of Dinosaurs'', Academic Press, San Diego</ref> He defined the group as all dinosaurs (more precisely: all thyreophoran Ornithischia) closer to ''Ankylosaurus'' than to ''Stegosaurus''. This definition is essentially followed by most paleontologists today. This "stem-based" definition means that the primitive armored dinosaur ''[[Scelidosaurus]]'', which is slightly closer to ankylosaurids than to stegosaurids, is technically a member of Ankylosauria. Upon the discovery of ''[[Bienosaurus]]'', [[Dong Zhiming]] (2001) erected the family [[Scelidosauridae]] for both of these primitive ankylosaurs.<ref>{{cite book|author=[[Dong Zhiming]]|year=2001|chapter=Primitive Armored Dinosaur from the Lufeng Basin, China|editor=Tanke, Darren H. |editor2=Carpenter, Kenneth |title=Mesozoic Vertebrate Life|pages=237–243|publisher= Indiana University Press|isbn=0-253-33907-3}}</ref> In 2001, Carpenter proposed a new group uniting ''Scelidosaurus'', [[Ankylosauridae]], [[Nodosauridae]], and [[Polacanthidae]], with ''[[Minmi (dinosaur)|Minmi]]'' (thought to be a primitive ankylosaurian), to the exclusion of ''[[Stegosaurus]]''.<ref name="carpenter2001">{{cite book|last1=Carpenter|first1=K.|editor1-last=Carpenter|editor1-first=Kenneth|title=The Armored Dinosaurs|date=2001|publisher=Indiana University Press|isbn=0-253-33964-2|page=455|url=https://books.google.ca/books?hl=en&lr=&id=04tQ5_qJN8MC|chapter=Phylogenetic Analysis of the Ankylosauria}}</ref> However, many taxonomists find that Ankylosauromorpha is an invalid group.<ref name="maidment06">{{cite journal|last1=Maidment|first1=S.C.R|last2=Wei|first2=G.|last3=Norman|first3=D.B.|title=Re-description of the postcranial skeleton of the middle Jurassic stegosaur ''Huayangosaurus taibaii''|journal=Journal of Vertebrate Paleontology|date=2006|volume=26|issue=4|pages=944–956|doi=10.1671/0272-4634(2006)26[944:ROTPSO]2.0.CO;2}}</ref>

===Nodosauridae===
This group traditionally includes ''[[Nodosaurus]]'', ''[[Edmontonia]]'', and ''[[Sauropelta]]''. The nodosauridae had longer snouts than their ankylosaurid cousins. They did not sport the archetypal 'clubs' at the ends of their tails, but rather, their most pronounced physical features were their spikes. Nodosaurids had very muscular shoulders, and a specialized knob of bone on each shoulder blade called the acromial process. It served as an attachment site for the muscles that held up their large parascapular spines. These spines would be used for self-defense against predators. They had wide, flaring hips and thick limbs. Most nodosaurid finds are from North America. They had smaller, narrow beaks than the ankylosaurids, which likely allowed them to be very selective over what plant matter they grazed on.

===Ankylosauridae===
Major differences distinguishing the ankylosaurids from the nodosaurids is that the ankylosaurids had bony [[Club (zoology)|clubs]] at the end of their tails, domed snouts in front of the eyes, and large squamosal plates projecting from the top and bottom of each side of the skull, all of which nodosaurids lacked. The traditional ankylosaurids are from later in the [[Cretaceous]]. They had much wider bodies and have even been discovered with bony eyelids. The large clubs at the end of their tails may have been used in self-defense (swung at predators) or in sexual selection. This family included ''[[Ankylosaurus]]'', ''[[Euoplocephalus]]'', and ''[[Pinacosaurus]]''. The clubs were made of several plates of bone that were permeated by soft tissue, allowing them to absorb thousands of pounds of force. Their beaks were larger and broader than the nodosaurids, indicating that these ankylosaurs were generalists in their diet.

===Polacanthidae===
The family [[Polacanthidae]] was named by [[George Reber Wieland]] in 1911 to refer to a group of ankylosaurs that seemed to him to be intermediate between the ankylosaurids and nodosaurids.<ref>G.R. Wieland, 1911, "Notes on the armored Dinosauria", ''The American Journal of Science, series 4'' '''31''': 112-124</ref> This grouping was ignored by most researchers until the late 1990s, when it was used as a subfamily (Polacanthinae) by Kirkland for a natural group recovered by his 1998 analysis suggesting that ''[[Polacanthus]]'', ''[[Gastonia (dinosaur)|Gastonia]]'', and ''[[Mymoorapelta]]'' were closely related within the family Ankylosauridae. Kenneth Carpenter resurrected the name Polacanthidae for a similar group that he also found to be closer to ankylosaurids than to nodosaurids. Carpenter became the first to define Polacanthidae as all dinosaurs closer to ''Gastonia'' than to either ''[[Edmontonia]]'' or ''[[Euoplocephalus]]''.<ref name="carpenter-2001">{{cite book|title=The Armored Dinosaurs|year=2001|chapter=Phylogenetic analysis of the Ankylosauria|editor=Carpenter, Kenneth |author=Carpenter K|pages=455–484|publisher=Indiana University Press|isbn=0-253-33964-2}}</ref> Most subsequent researchers placed polacanthines as primitive ankylosaurids, though mostly without any rigorous study to demonstrate this idea. The first comprehensive study of 'polacanthid' relationships, published in 2012, found that they are either an unnatural grouping of primitive nodosaurids, or a valid subfamily at the base of Nodosauridae.<ref name="phylogeny2012" />

==Taxonomy==
While ranked taxonomy has largely fallen out of favor among dinosaur paleontologists, a few 21st century publications have retained the use of ranks, though sources have differed on what its rank should be. Most have listed Thyreophora as an unranked taxon containing the traditional suborders Stegosauria and Ankylosauria, though Thyreophora is also sometimes classified as a suborder, with Ankylosauria and Stegosauria as infraorders. A simplified version of one possible classification follows:

* Order [[Ornithischia]] (the "bird hipped" dinosaurs)
** Clade [[Thyreophora]] (armored dinosaurs)
*** Clade [[Eurypoda]] (advanced armored dinosaurs)
**** Clade '''Ankylosauria'''
***** Family [[Ankylosauridae]]
****** Subfamily [[Ankylosaurinae]]
***** Family [[Nodosauridae]]
****** Subfamily [[Polacanthinae]]?

==See also==
{{Portal|Dinosaurs}}
* [[Timeline of ankylosaur research]]

==References==
{{reflist}}

{{Thyreophora|A.}}

[[Category:Ankylosaurs| ]]
[[Category:Bathonian first appearances]]
[[Category:Maastrichtian extinctions]]
[[Category:Taxa named by Henry Fairfield Osborn]]
[[Category:Fossil taxa described in 1923]]

Avaceratops

2018-02-11T16:34:32Z

Wikkler: /* Discovery and naming */ Greg Paul did NOT lump Avaceratops into Centrosaurus in Princeton Field Guide. I actually have the First Edition of Princeton Field Guide right here and can confirm that.

{{speciesbox
| fossil_range = [[Late Cretaceous]], {{fossilrange|77}}
| image = Academy of Natural Sciences, Philadelphia - IMG 7431.JPG
| image_caption = Mounted skeleton in the ANSP
| genus = Avaceratops
| parent_authority = [[Peter Dodson|Dodson]], 1986
| species = lammersi
| authority = Dodson, 1986
}}

'''''Avaceratops''''' is a [[genus]] of small herbivorous [[ceratopsia]]n [[dinosaur]]s which lived during the late [[Campanian]] during the [[Late Cretaceous]] [[Period (geology)|Period]] in what are now the Northwest [[United States]]. Most fossils come from the [[Judith River Formation]].

==Description==
[[File:Avaceratops BW.jpg|thumb|left|Restoration]]
''Avaceratops'' was originally by Dodson considered to have been an exceptionally small species. He estimated the holotype length at {{convert|2.3|m|ft}} and assumed it had almost attained adult size.<ref name="Dodson1986"/> However, the second skull (which does not necessarily belong to ''Avaceratops''),<ref name="ryan2016"/> MOR 692, indicates a body length of {{convert|4.2|m|ft}}.<ref name="Penkalski1999"/> Paul in 2010 estimated the weight of a four-metre-long animal at one tonne.<ref name="Paul2010"/> Another change in the image of ''Avaceratops'' brought about by MOR 692 concerned the brow horns. These were first thought to be rather short, although this was purely speculative, the holotype not having preserved them. The new skull showed postorbital horn cores with a length of {{convert|25|cm|in}}.<ref name="Penkalski1999"/>

''Avaceratops'' has a distinctive frill at the back of the skull. The squamosal, the element at the front of the frill side, is large with a continuously curving instead of a stepped edge. A raised area at the base of the squamosal divides it into two equal halves, whereas more derived species have an enlarged top part. The squamosal is separated from the [[parietal bone]] at the rear of the skull by a small indentation. The parietals however, show no indentation at the midline of the frill rear. Also the parietals are probably lacking ''[[Fenestra (anatomy)|fenestrae]]'' which are typical of many other genera except ''[[Triceratops]]'', resulting in a solid frill,<ref name="ageofdinosaursavaceratops"/> although damage to the holotype allows for a small opening to be present.<ref name="Dodson1986"/>

==Discovery and naming==
The first remains of ''Avaceratops'' were found by fossil dealer Eddie Cole in the [[Judith River Formation]] of [[Montana]], in 1981, on land of the Careless Creek Ranch, owned by rancher Arthur J. Lammers.<ref name="ageofdinosaursavaceratops">"Avaceratops." In: Dodson, Peter & Britt, Brooks & Carpenter, Kenneth & Forster, Catherine A. & Gillette, David D. & Norell, Mark A. & Olshevsky, George & Parrish, J. Michael & Weishampel, David B. ''The Age of Dinosaurs''. Publications International, LTD. p. 129. {{ISBN|0-7853-0443-6}}.</ref> They were preserved scattered throughout the remains of a prehistoric stream bed.<ref name="ageofdinosaursavaceratops"/> This ''Avaceratops'' specimen was likely buried in the sandbar after its body was swept downstream by the current.<ref name="ageofdinosaursavaceratops"/> The finds, displayed in Cole's fossil shop, were in October 1981 inspected by [[Peter Dodson]] who in July 1982 during a visit with Cole to the site discovered additional bones, which from 1984 were excavated by [[Anthony Fiorillo]].<ref name="Dodson1996">{{cite book|author=Dodson, P.|year=1996|title=The Horned Dinosaurs|publisher=Princeton University Press, Princeton, New Jersey|isbn=0-691-05900-4}}</ref>

The fossil was formally named and described by Dodson in 1986, as the [[type species]] ''Avaceratops lammersi''.<ref name="Dodson1986">{{cite journal|author=Dodson, P.|year=1986|title=Avaceratops lammersi: a new ceratopsid from the Judith River Formation of Montana.|journal=Proceedings of the Academy of Natural Sciences of Philadelphia|volume=138|issue=2|pages=305–317}}</ref> It was the first ceratopsid named since ''[[Pachyrhinosaurus]]'' in 1950.<ref name="Dodson1996"/> The genus was named after Ava Cole, Eddie's wife.<ref name="ageofdinosaursavaceratops"/> The specific epithet honors the Lammers family.<ref name="ageofdinosaursavaceratops"/> In 1990 George Olshevsky emended the name to ''A. lammersorum'', with the specific name in the genitive plural because it referred to several people.<ref>Olshevsky, G., 1991, ''A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia. Mesozoic Meanderings 2'' 196 pp</ref> However, Dodson objected against this change, arguing the genitive singular might also refer to a single family name.<ref name="Dodson1996"/> In 1990 [[Thomas Lehman]] renamed ''A. lammersi'' into ''[[Monoclonius]] lammersi'';<ref>Lehman, T.M., 1990, "The ceratopsian subfamily Chasmosaurinae: sexual dimorphism and systematics". In: K. Carpenter and P. J. Currie (eds.), ''Dinosaur Systematics: Perspectives and Approaches'', Cambridge University Press, Cambridge pp 211-229</ref> This alternative name has found no acceptance.

The [[holotype]], '''ANSP 15800''', consists of a partial skeleton containing the lower skull, a left lower jaw, vertebrae, a complete shoulder girdle and most elements of forelimbs and hindlimbs. The type specimen might represent a juvenile or a subadult individual, Dodson in 1986 being inclined to consider it almost fully grown.<ref name="Dodson1986"/> [[Kenneth Carpenter]] made a reconstruction of the skull, a cast of which was by Leroy Glenn combined with restored parts of the postcranial skeleton to create a mount that in 1986 was displayed in the [[Academy of Natural Sciences]] in [[Philadelphia]]; a copy of this skeletal mount was donated to the [[Upper Musselshell Valley Historical Museum]] in [[Harlowton]].<ref name="Dodson1996"/>

In 1993 [[Paul Penkalski]] referred two earlier found [[squamosal]]s to ''Avaceratops'', USNM 4802 and USNM 2415, of larger individuals.<ref>Penkalski, P.G., 1993, "The morphology of ''Avaceratops lammersi'', a primitive ceratopsid from the Campanian of Montana". ''Journal of Vertebrate Paleontology'' '''13'''(3, suppl.): 52A</ref> In 1999, Penkalski and Dodson described a second skull, MOR 692, again of a larger individual. This specimen includes the upper skull, with the nose and brow horns.<ref name="Penkalski1999">{{cite journal|doi=10.1080/02724634.1999.10011182|author1=Penkalski, P. |author2=Dodson, P. |lastauthoramp=yes |year=1999|title=The morphology and systematics of ''Avaceratops'', a primitive horned dinosaur from the Judith River Formation (Late Campanian) of Montana, with the description of a second skull.|journal=Journal of Vertebrate Paleontology|volume=19|issue=4|pages=692–711}}</ref> However, the assignment of the latter to ''Avaceratops'' has been questioned due to differences in relative chronological position, skull morphology, and lack of intermediate forms; thus, ''Avaceratops'' is only represented by the type specimen.<ref name="ryan2016">{{cite journal | last1 = Ryan | first1 = M.J. | last2 = Holmes | first2 = R. | last3 = Mallon | first3 = J. | last4 = Loewen | first4 = M. | last5 = Evans | first5 = D.C. | title = A basal ceratopsid (Centrosaurinae: Nasutoceratopsini) from the Oldman Formation (Campanian) of Alberta, Canada | doi = 10.1139/cjes-2016-0110 | journal = Canadian Journal of Earth Sciences | volume = 54 | date = 2017 | url = http://www.nrcresearchpress.com/doi/pdf/10.1139/cjes-2016-0110}}</ref>

== Classification ==
[[File:Avaceratops dinosaur.png|thumb|Restoration of two running specimens]]
''Avaceratops'' was by Dodson in 1986 assigned to the [[Ceratopsidae]] within the [[Ceratopsia]] (both names being derived from [[Ancient Greek]] for 'horned face'), a group of herbivorous dinosaurs with [[parrot]]-like beaks which thrived in what are now [[North America]] and [[Asia]], during the [[Cretaceous]] Period.

Apart from being a ceratopsian, little is certain about ''Avaceratops's'' taxonomic position. Because most of the skeleton is only known from a juvenile, and juveniles tend to express ancestral traits more strongly, a [[cladistic]] analysis would likely indicate a too basal, low, position in the evolutionary tree. For this reason, ''Avaceratops'' has often been excluded from such analyses. Penkalski and Dodson in 1999 concluded that ''Avaceratops'' most likely occupied a basal position within the [[Centrosaurinae]] but that it was also possible it was a basal member of the [[Ceratopsinae]] or just outside the [[clade]] formed by both groups. A phylogenetic analysis performed by Sampson et al. (2013) found that ''Avaceratops'' was the [[sister taxon]] of the new genus ''[[Nasutoceratops]]'', described and named in [[2013 in paleontology|2013]]. This clade containing both ''Avaceratops'' and ''Nasutoceratops'' was named the Nasutoceratopsini in 2016; a phylogenetic tree from the study in question is shown below.<ref name="ryan2016"/>

{{clade| style=font-size:85%; line-height:85%
|label1=[[Centrosaurinae]]
|1={{clade
|1=''[[Diabloceratops|Diabloceratops eatoni]]''
|2={{clade
|label2=Nasutoceratopsini
|1={{clade
|1={{clade
|1=''[[Albertaceratops|Albertaceratops nesmoi]]''
|2={{clade
|label1=Centrosaurini
|label2=Pachyrhinosaurini
|1={{clade
|1={{clade
|1=''[[Rubeosaurus|Rubeosaurus ovatus]]''
|2=''[[Styracosaurus|Styracosaurus albertensis]]''}}
|2={{clade
|1={{clade
|1=''[[Spinops|Spinops sternbergorum]]''
|2=''[[Centrosaurus|Centrosaurus apertus]]''}}
|2=''[[Coronosaurus|Coronosaurus brinkmani]]''}}}}
|2={{clade
|1=''[[Einiosaurus|Einiosaurus procurvicornis]]''
|2={{clade
|label2 = ''[[Pachyrhinosaurus]]''
|1=''[[Achelousaurus|Achelousaurus horneri]]''
|2={{clade
|1=''[[Pachyrhinosaurus|Pachyrhinosaurus canadensis]]''
|2={{clade
|1=''[[Pachyrhinosaurus|Pachyrhinosaurus lakustai]]''
|2=''[[Pachyrhinosaurus|Pachyrhinosaurus perotorum]]''}}}}}}}}}}
|3={{clade
|1=''[[Sinoceratops|Sinoceratops zhuchengensis]]''
|2=''[[Wendiceratops|Wendiceratops pinhornensis]]''}}}}
|2=''[[Xenoceratops|Xenoceratops foremostensis]]''}}
|2={{clade
|1='''''Avaceratops lammersi'' (ANSP 15800)'''
|2='''MOR 692'''
|3=CMN 8804
|4=''[[Nasutoceratops|Nasutoceratops titusi]]''
|5=Malta new taxon}}}}}}}}

== Paleobiology ==
''Avaceratops'', like all ceratopsians, was a [[herbivore]]. During the Cretaceous, flowering plants were "geographically limited on the landscape", so it is likely that this dinosaur fed on the predominant plants of the era: [[fern]]s, [[cycad]]s and [[conifer]]s. It would have used its sharp ceratopsian beak to bite off the leaves or needles. The [[habitat]] of ''Avaceratops'' was heavily forested and wet.<ref name="Paul2010"/>

==See also==
{{Portal|Dinosaurs}}
* [[Timeline of ceratopsian research]]

==References==

* https://web.archive.org/web/20050824122458/http://www.vertpaleo.org/jvp/19-692-711.html (online abstract of the preceding)
* https://web.archive.org/web/20090531083004/http://www.dinosaurvalley.com/Visiting_Drumheller/Kids_Zone/Groups_of_Dinosaurs/index.php
* ''[[The Humongous Book of Dinosaurs]]'', published by Stewart Tabori & Chang

==Notes==
{{Reflist|2}}

{{Marginocephalia|T.}}

[[Category:Centrosaurines]]
[[Category:Late Cretaceous dinosaurs of North America]]
[[Category:Fossil taxa described in 1986]]
[[Category:Taxa named by Peter Dodson]]
[[Category:Paleontology in Montana]]
[[Category:Campanian genus first appearances]]
[[Category:Campanian genus extinctions]]

Neochoristodera

2018-01-14T04:33:10Z

Wikkler: typo correction

{{Automatic taxobox
| name = Neochoristodera
| fossil_range = [[Early Cretaceous]] - [[Eocene]], {{fossilrange|130|40}}
| image = Champsosaurus_natator.jpg
| image_caption = Skeletal mount of ''[[Champsosaurus]]''
| taxon = Neochoristodera
| authority = [[Edward Drinker Cope|Cope]], 1884
| subdivision_ranks = Families
| subdivision =
*{{extinct}}[[Champsosauridae]]
*{{extinct}}[[Simoedosauridae]]
}}

'''Neochoristodera''' is a lineage of specialised [[crocodile]]-like fully aquatic [[choristodere]] [[reptile]]s. Noted for their long jaws and large size, these animals were predominant across the northern hemisphere (and possibly [[Australasia]]), occurring in freshwater and coastal environments across the [[Cretaceous]] and early [[Cenozoic]].

==Systematics==
Long-jawed choristoderes form a monophyletic group. They are generally divided into two main families: [[Champsosauridae]], which includes ''[[Champsosaurus]]'' and ''[[Eotomistoma]]'', and [[Simoedosauridae]] which includes ''[[Simoedosaurus]]'', ''[[Liaoxisaurus]]'', ''[[Ikechosaurus]]'' and ''[[Tchoiria]]''. Various taxa of uncertain affinities within this group are known, including a specimen from the [[Cedar Mountain Formation]] and from the [[Kuwajima Formation]].<ref>The first record of a long-snouted choristodere (Reptilia, Diapsida) from the Early Cretaceous of Ishikawa Prefecture, Japan
Article in Historical Biology 27(5):1-12 · December 2014
DOI: 10.1080/08912963.2014.898296</ref>

==Evolution==
[[File:Tchoiria_skeleton.PNG|thumb|left|Skeleton of ''[[Tchoiria]]'']]
Neochoristoderes first appear in the [[Early Cretaceous]] of [[Asia]], where they co-exist with other choristodere groups like [[monjurosuchidae|monjurosuchids]] and [[hyphalosauridae|hyphalosaurids]]. Here, a regional absence of aquatic [[crocodilian]]s, possibly due to colder temperatures, seems to have opened the ecological niche for these choristoderes to occupy a similar ecological niche.<ref>The first record of a long-snouted choristodere (Reptilia, Diapsida) from the Early Cretaceous of Ishikawa Prefecture, Japan
Article in Historical Biology 27(5):1-12 · December 2014
DOI: 10.1080/08912963.2014.898296</ref><ref>R. Matsumoto and S. E. Evans. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36(2):253-274</ref>

Other than a possible specimen from the [[Cedar Mountain Formation]], a large gap occurs between these Early Cretaceous faunas and the Late Cretaceous ones. There are no fossils of neochoristoderes in the Asian Late Cretaceous, but the subsequent distribution of ''[[Simoedosaurus]]'' in the [[Paleocene]] and [[Eocene]] implies that there were Asian taxa around this time, seeing as Simoedosauridae is predominantly Asian and ''[[Simoedosaurus]]'' only propagated widely after the KT event.<ref>R. Matsumoto and S. E. Evans. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36(2):253-274</ref>

[[File:Simoedosaurus_lemoinei_Cernay.JPG|thumb|right|Fossils of ''[[Simoedosaurus]]'']]
Choristoderes reappear in the fossil record in the [[Campanian]] of [[North America]] and [[Europe]] (and possibly [[Timor]]<ref>. H. F. Umbgrove, Structural History Of The East Indies</ref>) under the genus ''[[Champsosaurus]]''. This genus survives the KT event unscathed and diversifies in the aftermath, being soon after joined by ''[[Simoedosaurus]]''. Both taxa remain at high latitudes in [[North America]] and [[Europe]] until the Eocene, when they mysteriously disappear.<ref>R. Matsumoto and S. E. Evans. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36(2):253-274</ref>

Competition from [[crocodilia]]ns has been at times implicated in the extinction of neochoristoderes. There appears to be a niche partitioning between neochoristoderes and long-snouted crocodilians such as [[gharials]], which are absent from freshwater sites in Laurasia until well after neochoristoderes disappear, but competition between both groups, if it even existed, is currently unaccounted for. Ultimately, both ''[[Champsosaurus]]'' and ''[[Simoedosaurus]]'' co-existed with a variety of broad-snouted species like [[alligatorids]] and ''[[Borealosuchus]]''.<ref>R. Matsumoto and S. E. Evans. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36(2):253-274</ref><ref>The first record of a long-snouted choristodere (Reptilia, Diapsida) from the Early Cretaceous of Ishikawa Prefecture, Japan
Article in Historical Biology 27(5):1-12 · December 2014
DOI: 10.1080/08912963.2014.898296</ref>

==Biology==
[[File:Large_williston_champsosaurus.jpg|thumb|right|Lateral and dorsal views of a ''[[Champsosaurus]]'' skeletal mount]]
Neochoristoderes are thought to be fully aquatic. They possess laterally flattened, muscular tails and paddle-like limbs, their torsos are dorsoventrally flattened with short but massive ribs and their [[gastralia]] are heavily ossified, facilitating sinking. While they have their nostrils at the end of the snout as in crocodiles, they are oriented towards the tip instead of dorsally; their eye sockets are similarly forward-facing, implying that these animals did not surface often and probably simply rose the snout in a [[Snorkel (anatomy)|snorkel]]-like fashion. As in most choristoderes the skin lacks [[scute]]s, instead having small, non-overlapping scales.<ref>R. Matsumoto and S. E. Evans. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36(2):253-274</ref> In ''[[Champsosaurus]]'' adult females have particularly ossified limbs and pelvises, implying that they could crawl unto land, while adult males and juveniles could not.<ref>Yoshihiro Katsura, Fusion of sacrals and anatomy in Champsosaurus (Diapsida, Choristodera), doi:10.1080/08912960701374659</ref> As most choristoderes are thought to have been [[ovoviviparous]] or [[viviparous]], it is likely that at least some neochoristoderes were too.<ref>R. Matsumoto and S. E. Evans. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36(2):253-274</ref>

[[File:Champsosaurus laramiensis.JPG|thumb|left|''[[Champsosaurus]]'' skull]]
Most neochoristoderes have exceptionally large temporal fenestrae, suggesting powerful bite forces; ''[[Champsosaurus]]'' is estimated to have a bite force around 1194 to 1910 N, as opposed to the modern [[gharial]]'s 310-497 N.<ref>omateus.googlepages.com/Jacobsetal2009CretaceousSkeletoncoas.pdf
</ref> In spite of this, most are thought to have had a diet similar to that of modern gharials due to the long and slender jaws, though ''[[Simoedosaurus]]'' has a more robust and shorter snout and could have fed on larger prey.<ref>R. Matsumoto and S. E. Evans. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36(2):253-274</ref>

Compared with other choristoderes, which have rather simple teeth, neochoristoderes have teeth completely enveloped in striated [[Tooth enamel|enamel]] with an enamel infolding at the base, labiolingually compressed and hooked, the exception being ''[[Ikechosaurus]]'' which has still rather simple teeth aside from the start of an enamel infolding. There is some tooth differentiation, with the anterior teeth being larger than the posterior ones. As with most choristoderes, neochoristoderes have palatal teeth, indicating food manipulation in the mouth.<ref>Morphology and function of the palatal dentition in Choristodera Article in Journal of Anatomy 228(3):n/a-n/a · November 2015 DOI: 10.1111/joa.12414</ref>

Other than the possible Timor fossils, nearly all neochoristodere remains occur at high latitudes, suggesting a preference for colder climates.<ref>The first record of a long-snouted choristodere (Reptilia, Diapsida) from the Early Cretaceous of Ishikawa Prefecture, Japan
Article in Historical Biology 27(5):1-12 · December 2014
DOI: 10.1080/08912963.2014.898296</ref> ''[[Champsosaurus]]'' fossils are known in the [[Canada|Canadian]] [[Arctic]] and [[Greenland]].<ref>R. Matsumoto and S. E. Evans. 2010. Choristoderes and the freshwater assemblages of Laurasia. Journal of Iberian Geology 36(2):253-274</ref>

==References==
{{reflist}}

{{Choristodera}}

[[Category:Choristodera]]

Archosaur

2018-01-10T02:13:28Z

Wikkler:

{{Automatic taxobox
| name = Archosaurs
| taxon = Archosauria
| fossil_range = [[Early Triassic]]–[[Holocene|Present]], {{Fossil range|250|0}}
| image = Yellow-billed stork kazinga.jpg
| image_width = 250px
| image_caption = Birds and crocodilians (in this case a [[yellow-billed stork]] and a [[Nile crocodile]]) are the only living archosaur groups.
| authority = [[Edward Drinker Cope|Cope]], 1869
| subdivision_ranks = Subgroups
| subdivision =
* [[Avemetatarsalia]] (Ornithodira) ([[bird]]s and their extinct relatives)
* [[Pseudosuchia]] ([[crocodilia]]ns and their extinct relatives)
* {{extinct}}''[[Albisaurus]]''
* {{extinct}}''[[Avipes]]''
* {{extinct}}''[[Palaeosaurus]]''
* {{extinct}}''[[Sikannisuchus]]''
* {{extinct}}''[[Smok (archosaur)|Smok]]''
| synonyms =
Arctopoda [[Ernst Haeckel|Haeckel]], 1895 
Avesuchia Benton, 1999
}}

'''Archosaurs''' are a group of [[diapsid]] [[amniote]]s whose living representatives consist of [[birds]] and [[crocodilian]]s. This group also includes all extinct [[dinosaur|non-avian dinosaur]]s, extinct crocodilian relatives, and [[pterosaur]]s. '''Archosauria''', the archosaur [[clade]], is a [[crown group]] that includes the [[most recent common ancestor]] of living birds and crocodilians. It includes two main clades: [[Pseudosuchia]], which includes crocodilians and their extinct relatives, and [[Avemetatarsalia]], which includes birds and their extinct relatives (such as non-avian dinosaurs and [[pterosaurs]]).

==Distinguishing characteristics==
Archosaurs can be distinguished from other tetrapods on the basis of several [[Synapomorphy|synapomorphies]], or shared characteristics, first found in a common ancestor. The simplest and most widely agreed synapomorphies of archosaurs include teeth set in sockets, [[antorbital fenestra|antorbital]] and [[:File:Massospondylus Skull Steveoc 86.png|mandibular fenestrae]] (openings in front of the eyes and in the jaw, respectively),<ref name="Dyk, Kai 2011">{{cite book|title=Living Dinosaurs: The Evolutionary History of Modern Birds|year=2011|publisher=John Wiley & Sons|isbn=0470656662|pages=10|editor1=Gareth Dyke |editor2=Gary Kaiser }}</ref> and a [[fourth trochanter]] (a prominent ridge on the [[femur]]).<ref name=khanna2004>{{cite book|last=Khanna|first=D.R.|title=Biology Of Reptiles|year=2004|publisher=Discovery Publishing House|isbn=8171419070|pages=78ff}}</ref> Being set in sockets, the teeth were less likely to be torn loose during feeding. This feature is responsible for the name "[[Thecodontia|thecodont]]" (meaning "socket teeth"),<ref name="WhiKazAOver"/> which paleontologists used to apply to many Triassic archosaurs.<ref name=khanna2004/> Some archosaurs, such as birds, are secondarily toothless. Antorbital fenestrae reduced the weight of the skull, which was relatively large in early archosaurs, rather like that of modern [[crocodilian]]s. Mandibular fenestrae may also have reduced the weight of the jaw in some forms. The fourth trochanter provides a large site for the attachment of muscles on the femur. Stronger muscles allowed for erect gaits in early archosaurs, and may also be connected with the ability of the archosaurs or their immediate ancestors to survive the catastrophic [[Permian-Triassic extinction event]].{{Citation needed|date=April 2017}}

==Origins==
There is some debate about when archosaurs first appeared. Those who classify the [[Permian]] reptiles ''[[Archosaurus]] rossicus'' and/or ''[[Protorosaurus]] speneri'' as true archosaurs maintain that archosaurs first appeared in the late Permian. Some [[taxonomist]]s classify both ''Archosaurus rossicus'' and ''Protorosaurus speneri'' as [[archosauriform]]s (not true archosaurs but very closely related); these taxonomists maintain that archosaurs first evolved from archosauriform ancestors during the [[Olenekian]] stage of the [[Early Triassic]]. The earliest archosaurs were [[rauisuchia]]ns, such as ''[[Scythosuchus]]'' and ''[[Tsylmosuchus]]'', both of which have been found from [[Russia]] and date back to the Olenekian.<ref name=GS03>{{cite book |last1=Gower |first1=D. J. |last2=Sennikov |first2=A. G. |year=2003 |chapter=Early archosaurs from Russia |title=The Age of Dinosaurs in Russia and Mongolia |editor=Benton, M.J. |editor2=Shishkin, M.A. |editor3=Unwin, D.M. |publisher=Cambridge University Press |location=Cambridge |pages=140–159}}</ref>

==Archosaur takeover in the Triassic==
[[Synapsid]]s (a group including [[mammals]] and their extinct relatives, which are often referred to as "mammal-like reptiles") were the dominant land vertebrates throughout the [[Permian]], but most perished in the [[Permian-Triassic extinction event]]. Very few large synapsids survived the event, although one form, ''[[Lystrosaurus]]'' (an [[herbivorous]] [[dicynodont]]), attained a widespread distribution soon after the extinction.{{Citation needed|date=December 2017}}

But archosaurs quickly became the dominant land vertebrates in the early [[Triassic]]. The two most commonly suggested explanations for this are:{{Citation needed|date=February 2007}}
*Archosaurs made more rapid progress towards erect limbs than synapsids, and this gave them greater stamina by avoiding [[Carrier's constraint]]. An objection to this explanation is that archosaurs became dominant while they still had sprawling or semi-erect limbs, similar to those of ''[[Lystrosaurus]]'' and other synapsids.{{Citation needed|date=December 2017}}
*Archosaurs have more efficient respiratory systems featuring unidirectional air flow. Dr. Peter Ward suggests this may have proven advantageous in a suspected drop in oxygen levels at the end of the Permian.{{Citation needed|date=December 2017}}
*The early Triassic was predominantly arid, because most of the [[earth]]'s land was concentrated in the [[supercontinent]] [[Pangaea]]. Archosaurs were probably better at conserving water than early synapsids because:
**Modern [[diapsid]]s (lizards, snakes, crocodilians, birds) excrete [[uric acid]], which can be excreted as a paste, resulting in low water loss as opposed to a more dilute urine. It is reasonable to suppose that archosaurs (the ancestors of crocodilians, dinosaurs and birds) also excreted uric acid, and therefore were good at conserving water. The aglandular (glandless) skins of diapsids would also have helped to conserve water.{{Citation needed|date=December 2017}}
**Modern mammals excrete [[urea]], which requires a relatively high urinary rate to keep it from leaving the urine by diffusion in the kidney tubules. Their skins also contain many glands, which also lose water. Assuming that early synapsids had similar features, e.g., as argued by the authors of ''[[Palaeos]]'', they were at a disadvantage in a mainly arid world. The same well-respected site points out that "for much of Australia's [[Plio-Pleistocene]] history, where conditions were probably similar, the largest terrestrial predators were not mammals but gigantic [[varanid]] lizards (''[[Megalania]]'') and land crocs."<ref name="WhiKazAOver">{{cite web|author1=White, T. |author2=Kazlev, M. A. |title=Archosauromorpha: Overview |url=http://palaeos.com/Vertebrates/Units/270Archosauromorpha/270.000.html |publisher=Palaeos.com |accessdate=6 September 2012 |deadurl=yes |archiveurl=https://web.archive.org/web/20101220204743/http://palaeos.com/Vertebrates/Units/270Archosauromorpha/270.000.html |archivedate=December 20, 2010 }}</ref>

However, this theory has been questioned, since it implies synapsids were necessarily less advantaged in water retention, that synapsid decline coincides with climate changes or archosaur diversity (neither of which tested) and the fact that desert dwelling mammals as well adapted in this department as archosaurs,<ref>[[Darren Naish]], [http://tetzoo.com/podcast/2015/1/22/episode-38-a-not-too-shabby-podcarts Episode 38: A Not Too Shabby Podcarts]</ref> and some cynodonts like ''[[Trucidocynodon]]'' were large sized predators.<ref>{{cite journal | last1 = Oliveira | first1 = T.V. | last2 = Soares | first2 = M.B. | last3 = Schultz | first3 = C.L. | year = 2010 | title = Trucidocynodon riograndensis gen. nov. et sp. nov. (Eucynodontia), a new cynodont from the Brazilian Upper Triassic (Santa Maria Formation) | url = | journal = Zootaxa | volume = 2382 | issue = | pages = 1–71 }}</ref>

==Main forms==
[[File:Pancrocodylia diversity.jpg|thumb|left|200px|Examples of pseudosuchians. Clockwise from top-left: ''[[Longosuchus|Longosuchus meadei]]'' (an [[aetosaur]]), ''[[Gavialis gangeticus]]'', (a [[crocodilian]]), ''[[Saurosuchus|Saurosuchus galilei]]'' (a [[rauisuchia]]n), ''[[Pedeticosaurus|Pedeticosaurus leviseuri]]'' (a [[sphenosuchia]]n), ''[[Chenanisuchus|Chenanisuchus lateroculi]]'' (a [[Dyrosauridae|dyrosaurid]]), and ''[[Dakosaurus|Dakosaurus maximus]]'' (a [[thalattosuchia]]n).]]
[[File:Panaves diversity.jpg|thumb|right|200px|Examples of avemetatarsalians. Clockwise from top-left: ''[[Tupuxuara|Tupuxuara leonardi]]'' (a [[pterosaur]]), ''[[Alamosaurus|Alamosaurus sanjuanensis]]'', (a [[Sauropoda|sauropod]]), ''[[Tsintaosaurus|Tsintaosaurus spinorhinus]]'' (an [[Ornithopoda|ornithopod]]), ''[[Daspletosaurus|Daspletosaurus torosus]]'' (a [[Tyrannosauridae|tyrannosaur]]), ''[[Pentaceratops|Pentaceratops sternbergii]]'' (a [[ceratopsia]]n), and ''[[Common crane|Grus grus]]'' (a [[neornithes|neornithian]]).]]

Since the 1970s, scientists have classified archosaurs mainly on the basis of their ankles.<ref>[http://www.palaeos.com/Vertebrates/Units/270Archosauromorpha/270.500.html Archosauromorpha: Archosauria - Palaeos] {{webarchive|url=https://web.archive.org/web/20050405074210/http://palaeos.com/Vertebrates/Units/270Archosauromorpha/270.500.html |date=2005-04-05 }}</ref> The earliest archosaurs had "primitive mesotarsal" ankles: the [[Talus bone|astragalus]] and [[Calcaneus|calcaneum]] were fixed to the [[tibia]] and [[fibula]] by [[Suture (anatomical)|suture]]s and the joint bent about the contact between these bones and the foot.

The [[Pseudosuchia]] appeared early in the [[Triassic]]. In their ankles, the astragalus was joined to the tibia by a [[suture (anatomical)|suture]] and the joint rotated round a peg on the astragalus which fitted into a socket in the calcaneum. Early "crurotarsans" still walked with sprawling limbs, but some later crurotarsans developed fully erect limbs (most notably the [[Rauisuchia]]). Modern crocodilians are crurotarsans that can walk with their limbs sprawling or erect depending on speed of locomotion.

''[[Euparkeria]]'' and the [[Ornithosuchidae]] had "reversed crurotarsal" ankles, with a peg on the calcaneum and socket on the astragalus.

The earliest fossils of [[Avemetatarsalia]] ("bird ankles") appear in the [[Carnian]] age of the late [[Triassic]], but it is hard to see how they could have evolved from crurotarsans — possibly they actually evolved much earlier, or perhaps they evolved from the last of the "primitive mesotarsal" archosaurs. Ornithodires' "advanced mesotarsal" ankle had a very large astragalus and very small calcaneum, and could only move in one plane, like a simple hinge. This arrangement, which was only suitable for animals with erect limbs, provided more stability when the animals were running. The ornithodires differed from other archosaurs in other ways: they were lightly built and usually small, their necks were long and had an S-shaped curve, their skulls were much more lightly built, and many ornithodires were completely [[bipedalism|bipedal]]. The archosaurian fourth trochanter on the femur may have made it easier for ornithodires to become bipeds, because it provided more leverage for the thigh muscles. In the late Triassic, the ornithodires diversified to produce [[dinosaur]]s and [[pterosaur]]s.

==Classification==

===Modern classification===
Archosauria is normally defined as a [[crown group]], which means that it only includes descendants of the last common ancestors of its living representatives. In the case of archosaurs, these are birds and crocodilians. Archosauria is within the larger clade [[Archosauriformes]], which includes some close relatives of archosaurs, such as [[Proterochampsidae|proterochampsids]] and [[Euparkeriidae|euparkeriids]]. These relatives are often referred to as archosaurs despite being placed outside of the crown group Archosauria in a more [[basal (phylogenetics)|basal]] position within Archosauriformes.<ref name=GW96>{{cite journal |last1=Gower |first1=D. J. |last2=Wilkinson |first2=M. |year=1996 |title=Is there any consensus on basal archosaur phylogeny? |journal=Proceedings of the Royal Society B |volume=263 |issue=1375 |pages=1399–1406 |url=http://www.bmnh.org/PDFs/DG_96_basal.pdf | doi = 10.1098/rspb.1996.0205 }}</ref> Historically, many archosauriforms were described as archosaurs, including [[proterosuchid]]s and [[erythrosuchid]]s, based on the presence of an antorbital fenestra. While many researchers prefer to treat Archosauria as an unranked [[clade]], some continue to assign it a traditional biological rank. Traditionally, Archosauria has been treated as a Superorder, though a few 21st century researchers have assigned it to different ranks including Division<ref>Benton, M.J. (2005). ''Vertebrate Paleontology'', 3rd ed. Blackwell Science Ltd</ref> and Class.<ref>{{cite journal | last1 = Göhlich | first1 = U.B. | last2 = Chiappe | first2 = L.M. | last3 = Clark | first3 = J.M. | last4 = Sues | first4 = H.-D. | year = 2005 | title = The systematic position of the Late Jurassic alleged dinosaur ''Macelognathus'' (Crocodylomorpha: Sphenosuchia) | url = | journal = Canadian Journal of Earth Sciences | volume = 42 | issue = 3| pages = 307–321 | doi=10.1139/e05-005}}</ref>

===History of classification===
Archosauria as a term was first coined by American paleontologist [[Edward Drinker Cope]] in 1869, and included a wide range of taxa including [[dinosaur]]s, [[crocodilia]]ns, [[Thecodontia|thecodont]]s, [[sauropterygia]]ns (which may be related to turtles), [[rhynchocephalia]]ns (a group that according to Cope included [[rhynchosaur]]s, which nowadays are considered to be more basal [[archosauromorph]]s, and [[tuatara]]s, which are [[Lepidosauria|lepidosaurs]]), and [[anomodont]]s, which are now considered synapsids.<ref>{{cite journal |last=Cope |first=Edward Drinker |year=1869 |title=Synopsis of the extinct Batrachia, Reptilia and Aves of North America |journal=Transactions of the America Philosophical Society |volume=14 |pages=1–252 |url=https://archive.org/details/synopsisofextin00cope}}</ref> It was not until 1986 that Archosauria was defined as a crown-clade, restricting its use to more [[Synapomorphy|derived]] taxa.<ref name=BMJ04>{{cite book |last=Benton |first=M.J. |editor=Weishampel, D.B. |editor2=Dodson, P.r |editor3=Osmólska, H. |title=The Dinosauria |edition=2nd |year=2004 |publisher=University of California Press |location=Berkeley |isbn=0-520-24209-2 |pages=7–19 |chapter=Origin and relationships of Dinosauria}}</ref>
{{multiple image|perrow=2|align=left
| header = Archosaur ankle types: Adapted with permission from [https://web.archive.org/web/20050405074210/http://palaeos.com/Vertebrates/Units/270Archosauromorpha/270.500.html Palaeos] {{legend2| green|[[Tibia]]}}{{nbsp|5}}{{legend2| yellow|[[Fibula]]}}{{nbsp|5}}{{legend2|#ff6666|[[Talus bone|Astragalus]]}}{{nbsp|5}}{{legend2|#6600ff|[[Calcaneum]]}}
| header_background = inherit; font-weight:normal;
| image1 = PrimitiveMesotarsal01.gif| width1 = 250|caption1=Primitive mesotarsal ankle
| image2 = CrocNormal01.png| width2 = 250|caption2=Crocodilian form of [[crurotarsal]] ankle
| image3 = CrocReversed01.png| width3 = 250|caption3=Reversed [[crurotarsal]] ankle
| image4 = AdvMesotarsal.png| width4 = 250|caption4="Advanced" mesotarsal ankle
}}
Cope's term was a Greek-Latin [[hybrid word|hybrid]] intended to refer to the cranial arches, but has later also been understood as "leading reptiles" or "ruling reptiles" by association with Greek [[:wikt:ἀρχός|ἀρχός]] "leader, ruler".<ref>''Pamphlets on Biology: Kofoid collection'', vol. 2900 (1878), [https://books.google.com/books?id=BTgXAQAAIAAJ&q=%22den+Terminus+Archosauria+gebrauchten%22&dq=%22den+Terminus+Archosauria+gebrauchten%22&hl=de&ei=yAU8TdPwPMzY4gb7tuTACg&sa=X&oi=book_result&ct=result&resnum=2&ved=0CDIQ6AEwAQ p. 731]</ref>

The term "thecodont", now considered an obsolete term, was first used by the English paleontologist [[Richard Owen]] in 1859 to describe Triassic archosaurs, and it became widely used in the 20th century. Thecodonts were considered the "basal stock" from which the more advanced archosaurs descended. They did not possess features seen in later avian and crocodilian lines, and therefore were considered more primitive and ancestral to the two groups. With the [[Cladistics#History of cladistics|cladistic revolution]] of the 1980s and 90s, in which [[cladistics]] became the most widely used method of classifying organisms, thecodonts were no longer considered a valid grouping. Because they are considered a "basal stock", thecodonts are [[paraphyletic]], meaning that they form a group that does not include all descendants of its last common ancestor: in this case, the more derived crocodilians and birds are excluded from "Thecodontia" as it was formerly understood. The description of the basal [[ornithodire]]s ''[[Lagerpeton]]'' and ''[[Lagosuchus]]'' in the 1970s provided evidence that linked thecodonts with dinosaurs, and contributed to the disuse of the term "Thecodontia", which many cladists consider an artificial grouping.<ref name=SPC91>{{cite journal |doi=10.2307/3889336 |last=Sereno |first=P.C. |year=1991 |title=Basal archosaurs: phylogenetic relationships and functional implications |jstor=3889336 |journal=Memoir (Society of Vertebrate Paleontology) |volume=2 |pages=1–53}}</ref>

With the identification of "crocodilian normal" and "crocodilian reversed" ankles by [[Sankar Chatterjee]] in 1978, a basal split in Archosauria was identified. Chatterjee considered these two groups to be Pseudosuchia with the "normal" ankle and Ornithosuchidae with the "reversed" ankle. Ornithosuchids were thought to be ancestral to dinosaurs at this time. In 1979, [[A.R.I. Cruickshank]] identified the basal split and thought that the crurotarsan ankle developed independently in these two groups, but in opposite ways. Cruickshank also thought that the development of these ankle types progressed in each group to allow advanced members to have semi-erect (in the case of crocodilians) or erect (in the case of dinosaurs) gaits.<ref name=SPC91/>

===Phylogeny===
In many [[phylogenetic]] analyses, archosaurs have been shown to be a [[monophyletic]] grouping, thus forming a true clade. One of the first studies of archosaur phylogeny was authored by French paleontologist [[Jacques Gauthier]] in 1986. Gauthier split Archosauria into [[Pseudosuchia]], the crocodilian line, and [[Ornithosuchia]], the dinosaur and pterosaur line. Pseudosuchia was defined as all archosaurs more closely related to crocodiles, while Ornithosuchia was defined as all archosaurs more closely related to birds. Proterochampsids, erythrosuchids, and proterosuchids fell successively outside Archosauria in the resulting tree. Below is the [[cladogram]] from Gauthier (1986):<ref name=GJA86>{{cite book |last=Gauthier |first=J.A. |year=1986 |chapter=Saurischian monophyly and the origin of birds |title=The Origin of Birds and the Evolution of Flight. Memoirs of the California Academy of Sciences |editor=Padian, K. |volume=8 |publisher=California Academy of Sciences |location=San Francisco |pages=1–55}}</ref>
{{clade| style=font-size:80%;line-height:85%
|1={{clade
|1={{extinct}}[[Proterosuchidae]][[File:ProterosuchusDB flipped.jpg|80 px]]
|2={{clade
|1={{extinct}}[[Erythrosuchidae]][[File:Erythrosuchus afr12DB.jpg|80 px]]
|2={{clade
|1={{extinct}}[[Proterochampsidae]][[File:Chanaresuchus.jpg|80 px]]
|label2= '''Archosauria''' 
|2={{clade
|label1= [[Pseudosuchia]] 
|1={{clade
|1={{extinct}}[[Parasuchia]][[File:Smilosuchus adamanensis flipped.jpg|80px]]
|2={{clade
|1={{extinct}}[[Aetosauria]][[File:Desmatosuchus spurensis flipped.jpg|80px]]
|2={{clade
|1={{extinct}}[[Rauisuchia]][[File:Postosuchus kirkpatricki flipped.jpg|80px]]
|2=[[Crocodylomorpha]][[File:Description des reptiles nouveaux, ou, Imparfaitement connus de la collection du Muséum d'histoire naturelle et remarques sur la classification et les caractères des reptiles (1852) (Crocodylus moreletii).jpg|80px]]}} }} }}
|label2= [[Ornithosuchia]] 
|2={{clade
|1={{extinct}}''[[Euparkeria]]''[[File:Euparkeria BW flipped.jpg|80 px]]
|2={{clade
|1={{extinct}}[[Ornithosuchidae]][[File:Ornithosuchus BW white background.jpg|80 px]]
|2=[[Ornithodira]][[File:Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg|40 px]]
}} }} }} }} }} }} }}

In 1988, paleontologists [[Michael Benton]] and J.M. Clark produced a new tree in a phylogenetic study of basal archosaurs. As in Gauthier's tree, Benton and Clark's revealed a basal split within Archosauria. They referred to the two groups as Crocodylotarsi and Ornithosuchia. Crocodylotarsi was defined as an [[apomorphy]]-based taxon based on the presence of a "crocodile-normal" ankle joint (considered to be the defining apomorphy of the clade). Gauthier's Pseudosuchia, by contrast, was a [[stem-based taxon]]. Unlike Gauthier's tree, Benton and Clark's places ''Euparkeria'' outside Ornithosuchia and outside the crown group Archosauria altogether.<ref name=BC88>{{cite book |last1=Benton |first1=M. J. |last2=Clark |first2=J. M. |year=1985 |chapter=Archosaur phylogeny and the relationships of the Crocodylia |title=The Phylogeny and Classification of the Tetrapods |volume=1 |editor=Benton, M.J. |publisher=Clarendon Press |location=Oxford |pages=295–338 |isbn=0-19-857712-5}}</ref>

The clades Crurotarsi and Ornithodira were first used together in 1990 by paleontologist [[Paul Sereno]] and A.B. Arcucci in their phylogenetic study of archosaurs. They were the first to erect the clade Crurotarsi, while Ornithodira was named by Gauthier in 1986. Crurotarsi and Ornithodira replaced Pseudosuchia and Ornithosuchia, respectively, as the monophyly of both of these clades were questioned.<ref name=SPC91/><ref name=SA90>{{cite journal |last1=Sereno |first1=P. C. |last2=Arcucci |first2=A. B. |year=1990 |title=The monophyly of crurotarsal archosaurs and the origin of bird and crocodile ankle joints |journal=Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen |volume=180 |pages=21–52}}</ref> Sereno and Arcucci incorporated archosaur features other than ankle types in their analyses, which resulted in a different tree than previous analyses. Below is a cladogram based on Sereno (1991), which is similar to the one produced by Sereno and Arcucci:<ref name=SPC91/>
{{clade| style=font-size:80%;line-height:85%
|label1=[[Archosauriformes]] 
|1={{clade
|1={{extinct}}[[Proterosuchidae]][[File:ProterosuchusDB flipped.jpg|80 px]]
|2={{clade
|1={{extinct}}[[Erythrosuchidae]][[File:Erythrosuchus afr12DB.jpg|80 px]]
|2={{clade
|1={{extinct}}''[[Euparkeria]]''[[File:Euparkeria BW flipped.jpg|80 px]]
|2={{clade
|1={{extinct}}[[Proterochampsidae]][[File:Chanaresuchus.jpg|80 px]]
|label2= '''Archosauria''' 
|2={{clade
|label1= [[Crurotarsi]] 
|1={{clade
|1={{extinct}}[[Parasuchia]][[File:Smilosuchus adamanensis flipped.jpg|80px]]
|2={{clade
|1={{extinct}}[[Ornithosuchidae]][[File:Ornithosuchus BW white background.jpg|80 px]]
|2=[[Suchia]][[File:Description des reptiles nouveaux, ou, Imparfaitement connus de la collection du Muséum d'histoire naturelle et remarques sur la classification et les caractères des reptiles (1852) (Crocodylus moreletii).jpg|80px]]}} }}
|label2= [[Ornithodira]] 
|2={{clade
|1={{clade
|1={{extinct}}?''[[Scleromochlus]]''
|2={{extinct}}[[Pterosauria]][[File:Aerodactylus MCZ 1505.png|80 px]]}}
|2=[[Dinosauromorpha]][[File:Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg|40 px]]
}} }} }} }} }} }} }}

Ornithodira and Crurotarsi are both [[node-based taxon|node-based]] clades, meaning that they are defined to include the [[last common ancestor]] of two or more taxa and all of its descendants. Ornithodira includes the last common ancestor of pterosaurs and dinosaurs (which include birds), while Crurotarsi includes the last common ancestor of living crocodilians and three groups of Triassic archosaurs: [[ornithosuchid]]s, [[aetosaur]]s, and [[phytosaur]]s. These clades are not equivalent to "bird-line" and "crocodile-line" archosaurs, which would be [[stem-based taxon|branch-based]] clades defined as all taxa more closely related to one living group (either birds or crocodiles) than the another.

Benton proposed the name Avemetatarsalia in 1999 to include all bird-line archosaurs (under his definition, all archosaurs more closely related to dinosaurs than to crocodilians). His analysis of the small Triassic archosaur ''[[Scleromochlus]]'' placed it within bird-line archosaurs but outside Ornithodira, meaning that Ornithodira was no longer equivalent to bird-line archosaurs. Below is a cladogram modified from Benton (2004) showing this phylogeny:<ref name=BMJ04/>
{{clade| style=font-size:80%;line-height:85%
|label1='''Archosauria''' 
|1={{clade
|1={{extinct}}''[[Hyperodapedon]]'' ([[Rhynchosauria]]) [[File:Hyperodapedon BW2 white background.jpg|80 px]]
|2={{clade
|1={{extinct}}''[[Prolacerta]]'' ([[Prolacertiformes]]) [[File:Prolacerta broomi.jpg|80 px]]
|2={{clade
|1={{extinct}}''[[Proterosuchus]]'' ([[Proterosuchidae]])[[File:ProterosuchusDB flipped.jpg|80 px]]
|2={{clade
|1={{extinct}}''[[Euparkeria]]'' ([[Euparkeriidae]])[[File:Euparkeria BW flipped.jpg|80 px]]
|2={{clade
|1={{extinct}}''[[Proterochampsidae]]''[[File:Chanaresuchus.jpg|80 px]]
|label2=''Avesuchia'' |sublabel2= (Crown group Archosauria) 
|2={{clade
|label1= [[Crurotarsi]] 
|1={{clade
|1={{clade
|1={{extinct}}[[Phytosauridae]][[File:Smilosuchus adamanensis flipped.jpg|80px]]
|2={{extinct}}''[[Gracilisuchus]]''[[File:Gracilisuchus BW white background.jpg|80 px]]}}
|2={{extinct}}[[Ornithosuchidae]][[File:Ornithosuchus BW white background.jpg|80 px]]
|label3= [[Suchia]] 
|3={{clade
|1={{extinct}}[[Stagonolepididae]][[File:Desmatosuchus spurensis flipped.jpg|80px]]
|2={{clade
|1={{extinct}}''[[Postosuchus]]''[[File:Postosuchus kirkpatricki flipped.jpg|80px]]
|2=[[Crocodylomorpha]][[File:Description des reptiles nouveaux, ou, Imparfaitement connus de la collection du Muséum d'histoire naturelle et remarques sur la classification et les caractères des reptiles (1852) (Crocodylus moreletii).jpg|80px]]}}
|3={{clade
|1={{extinct}}''[[Fasolasuchus]]''
|label2= [[Prestosuchidae]]
|2={{clade
|1={{extinct}}''[[Ticinosuchus]]''[[File:Ticinosuchus BW white background.jpg|80px]]
|2={{extinct}}''[[Prestosuchus]]''[[File:Prestosuchus11DB.jpg|80px]]
|3={{extinct}}''[[Saurosuchus]]''[[File:Saurosuchus BW white background.jpg|80px]]}} }} }} }}
|label2= [[Avemetatarsalia]] 
|2={{clade
|1={{extinct}}''[[Scleromochlus]]''
|label2= [[Ornithodira]] 
|2={{clade
|1={{extinct}}[[Pterosauria]][[File:Aerodactylus MCZ 1505.png|80 px]]
|label2= [[Dinosauromorpha]]
|2={{clade
|1={{extinct}}''[[Lagerpeton]]''
|label2= [[Dinosauriformes]] 
|2={{clade
|1={{extinct}}''[[Marasuchus]]''[[File:Marasuchus flipped.jpg|80 px]]
|label2=[[Dinosauria]]
|2={{clade
|1={{extinct}}[[Ornithischia]][[File:Triceratops liveDB.jpg|80px]]
|label2= [[Saurischia]] 
|2={{clade
|1={{extinct}}[[Sauropodomorpha]][[File:Barapasaurus DB.jpg|80 px]]
|label2= [[Theropoda]] 
|2={{clade
|1={{extinct}}''[[Herrerasaurus]]''[[File:Herrerasaurus BW flipped.jpg|80 px]]
|2=[[Neotheropoda]][[File:Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg|40 px]]
}} }} }} }} }} }} }} }} }} }} }} }} }} }}

In [[Sterling Nesbitt]]'s 2011 monograph on early archosaurs, a phylogenetic analysis found strong support for phytosaurs falling outside Archosauria. Many subsequent studies supported this phylogeny. Because Crurotarsi is defined by the inclusion of phytosaurs, the placement of phytosaurs outside Archosauria means that Crurotarsi must include all of Archosauria. Nesbitt reinstated Pseudosuchia as a clade name for crocodile-line archosaurs, using it as a stem-based taxon. Below is a cladogram modified from Nesbitt (2011):<ref name=NSJ11>{{cite journal |last=Nesbitt |first=S.J. |year=2011 |title=The early evolution of archosaurs: relationships and the origin of major clades |journal=Bulletin of the American Museum of Natural History |volume=352 |pages=1–292 |url=http://digitallibrary.amnh.org/dspace/bitstream/2246/6112/1/B352.pdf |doi=10.1206/352.1}}</ref>

{{clade| style=font-size:80%;line-height:85%
|1={{clade
|1={{extinct}}[[Phytosauria]][[File:Smilosuchus adamanensis flipped.jpg|80px]]
|label2='''Archosauria'''
|2={{clade
|label1='''[[Pseudosuchia]]'''
|1={{clade
|1={{extinct}}[[Ornithosuchidae]][[File:Ornithosuchus BW white background.jpg|80 px]]
|label2='''[[Suchia]]'''
|2={{clade
|1={{extinct}}''[[Gracilisuchus]]''[[File:Gracilisuchus BW white background.jpg|80 px]]
|2={{extinct}}''[[Turfanosuchus]]''
|3={{clade
|1={{extinct}}''[[Revueltosaurus]]''
|2={{extinct}}[[Aetosauria]][[File:Desmatosuchus spurensis flipped.jpg|80px]]}}
|4={{clade
|1={{extinct}}''[[Ticinosuchus]]''[[File:Ticinosuchus BW white background.jpg|80px]]
|label2='''[[Paracrocodylomorpha]]'''
|2={{clade
|1={{extinct}}[[Poposauroidea]][[File:Poposaurus gracilis (1) flipped.jpg|80px]]
|label2='''[[Loricata]]'''
|2={{clade
|1={{extinct}}''[[Prestosuchus]]''[[File:Prestosuchus11DB.jpg|80px]]
|2={{clade
|1={{extinct}}''[[Saurosuchus]]''[[File:Saurosuchus BW white background.jpg|80px]]
|2={{clade
|1={{extinct}}''[[Batrachotomus]]''
|2={{clade
|1={{extinct}}''[[Fasolasuchus]]''
|2={{clade
|1={{extinct}}[[Rauisuchidae]][[File:Postosuchus kirkpatricki flipped.jpg|80px]]
|2='''[[Crocodylomorpha]]'''[[File:Description des reptiles nouveaux, ou, Imparfaitement connus de la collection du Muséum d'histoire naturelle et remarques sur la classification et les caractères des reptiles (1852) (Crocodylus moreletii).jpg|80px]]}} }} }} }} }} }} }} }} }}
|label2='''[[Avemetatarsalia]]'''/
|sublabel2= '''[[Ornithodira]]'''*
|2={{clade
|1={{extinct}}[[Pterosauromorpha]][[File:Aerodactylus MCZ 1505.png|80 px]]
|label2='''[[Dinosauromorpha]]'''
|2={{clade
|1={{extinct}}[[Lagerpetidae]]
|label2='''[[Dinosauriformes]]'''
|2={{clade
|1={{extinct}}''[[Marasuchus]]''[[File:Marasuchus flipped.jpg|80 px]]
|2={{clade
|1={{extinct}}[[Silesauridae]][[File:Silesaurus opolensis flipped.jpg|80 px]]
|label2='''[[Dinosauria]]'''
|2={{clade
|1={{extinct}}[[Ornithischia]][[File:Triceratops liveDB.jpg|80px]]
|label2='''[[Saurischia]]'''
|2={{clade
|1={{extinct}}[[Sauropodomorpha]][[File:Barapasaurus DB.jpg|80 px]]
|2='''[[Theropoda]]'''[[File:Meyers grosses Konversations-Lexikon - ein Nachschlagewerk des allgemeinen Wissens (1908) (Antwerpener Breiftaube).jpg|40 px]]
}} }} }} }} }} }} }} }} * Nesbitt did not include ''Scleromochlus'' in the analysis, meaning that Avemetatarsalia and Ornithodira occupy the same place in this cladogram }}

==Extinction and survival==
Crocodilians, pterosaurs and dinosaurs survived the [[Triassic–Jurassic extinction event]] about 200 million years ago, but other archosaurs became extinct.

Non-avian dinosaurs and [[pterosaur]]s perished in the [[Cretaceous–Paleogene extinction event]], which occurred approximately {{period start|Paleogene}} million years ago, but birds (the only remaining dinosaur group) and crocodilians survived. Both are descendants of archosaurs, and are therefore archosaurs themselves under [[phylogenetic taxonomy]].

Crocodilians (which include all modern [[crocodile]]s, [[alligator]]s, and [[gharial]]s) and birds flourish today. It is generally agreed that birds have the most species of all terrestrial vertebrates.{{Citation needed|date=January 2014}}

==Archosaur lifestyle==

===Hip joints and locomotion===
[[File:Sprawling and erect hip joints - horizontal.svg|thumb|right|250px|Hip joints and hindlimb postures.]]
Like the early [[tetrapod]]s, early archosaurs had a sprawling gait because their hip sockets faced sideways, and the knobs at the tops of their [[femur]]s were in line with the femur.
In the early to middle [[Triassic]], some archosaur groups developed hip joints that allowed (or required) a more erect gait. This gave them greater stamina, because it avoided [[Carrier's constraint]], i.e. they could run and breathe easily at the same time. There were two main types of joint which allowed erect legs:
*The hip sockets faced sideways, but the knobs on the femurs were at right angles to the rest of the femur, which therefore pointed downwards. Dinosaurs evolved from archosaurs with this hip arrangement.
*The hip sockets faced downwards and the knobs on the femurs were in line with the femur. This "pillar-erect" arrangement appears to have evolved independently in various archosaur lineages, for example it was common in [[Rauisuchia]] and also appeared in some [[aetosaur]]s.
It has been pointed out{{by whom|date=January 2016}} that an upright stance requires more energy, so it may indicate a higher metabolism and a higher body temperature.<ref>Desmond, Adrián J., The hot-blooded dinosaurs: a revolution in palaeontology. 1976, Dial Press, Page 87.</ref>

===Diet===
Most were large predators, but members of various lines diversified into other niches. [[Aetosaur]]s were herbivores and some developed extensive armor. A few crocodilians were herbivores, e.g., ''[[Simosuchus]]'', ''[[Phyllodontosuchus]]''. The large crocodilian ''[[Stomatosuchus]]'' may have been a [[filter feeder]]. [[Sauropodomorpha|Sauropodomorphs]] and [[ornithischia]]n dinosaurs were herbivores with diverse adaptations for feeding [[biomechanics]].

===Land, water and air===
Archosaurs are mainly portrayed as [[Terrestrial animal|land]] animals, but:
*The phytosaurs and crocodilians dominated the rivers and swamps and even invaded the seas (e.g., the [[Teleosauridae|teleosaurs]], [[Metriorhynchidae]] and [[Dyrosauridae]]). The Metriorhynchidae were rather dolphin-like, with paddle-like forelimbs, a tail fluke and smooth, unarmoured skins.
*Two clades of ornithodirans, the [[pterosaurs]] and the birds, dominated the air after becoming adapted to a volant lifestyle.

===Metabolism===
The metabolism of archosaurs is still a controversial topic. They certainly evolved from cold-blooded ancestors, and the surviving non-dinosaurian archosaurs, crocodilians, are cold-blooded. But crocodilians have some features which are normally associated with a warm-blooded metabolism because they improve the animal's oxygen supply:
*4-chambered hearts. Both birds and mammals have 4-chambered hearts, which completely separate the flows of oxygenated and de-oxygenated [[blood]]. Non-crocodilian reptiles have 3-chambered [[heart]]s, which are less efficient because they let the two flows mix and thus send some de-oxygenated blood out to the body instead of to the lungs. Modern crocodilians' hearts are 4-chambered, but are smaller relative to body size and run at lower pressure than those of modern birds and [[mammal]]s. They also have a [[pulmonary bypass]], which makes them functionally 3-chambered when under water, conserving [[oxygen]].
*a [[secondary palate]], which allows the animal to eat and breathe at the same time.
*a [[Reptile#Respiratory system|hepatic piston]] mechanism for pumping the [[lungs]]. This is different from the lung-pumping mechanisms of mammals and birds, but similar to what some researchers claim to have found in some dinosaurs.<ref>{{cite journal|author=Ruben, J. | title=The metabolic status of some Late Cretaceous dinosaurs | journal=Science | issue=5279 | pages=120–147 | year=1996 |volume=273|doi=10.1126/science.273.5279.1204|display-authors=etal}}</ref><ref>{{cite journal|author=Ruben, J. | title=Lung structure and ventilation in theropod dinosaurs and early birds | journal=Science | issue=5341 | pages= 1267–1270| year=1997 |volume=278|doi=10.1126/science.278.5341.1267|display-authors=etal}}</ref>

So, why did [[natural selection]] favour the development of these features, which are very important for active warm-blooded creatures, but of little apparent use to cold-blooded aquatic ambush [[predator]]s that spend the vast majority of their time floating in water or lying on river banks?
[[File:Terrestrisuchus BW.jpg|thumb|right|150px|''Terrestrisuchus'']]
[[File:Isochirotherium Footprint - geograph.org.uk - 2359261.jpg|thumb|right|150px|''[[Chirotherium]]'' footprint in Triassic sediments]]
Paleontological evidence shows that the ancestors of living crocodilians were active and endothermic (warm-blooded). Some experts believe that their archosaur ancestors were warm-blooded as well. Physiological, anatomical, and developmental features of the crocodilian heart support the paleontological evidence and show that the lineage reverted to ectothermy when it invaded the aquatic, ambush predator niche. Crocodilian embryos develop fully 4-chambered hearts at an early stage. Modifications to the growing heart form a pulmonary bypass shunt that includes the left [[aorta|aortic arch]], which originates from the right [[Ventricle (heart)|ventricle]], the [[foramen of Panizza]] between the left and right aortic arches, and the cog‐tooth valve at the base of the [[pulmonary artery]]. The shunt is used during diving to make the heart function as 3-chambered heart, providing the crocodilian with the neurally controlled shunting used by ectotherms. The researchers concluded that the ancestors of living crocodilians had fully 4-chambered hearts, and were therefore warm-blooded, before they reverted to a cold-blooded or ectothermic metabolism. The authors also provide other evidence for endothermy in stem archosaurs.<ref>{{cite journal|author=Seymour, R. S.|author2=Bennett-Stamper, C. L.|author3=Johnston, S. D.|author4=Carrier, D. R.|author5=Grigg, G. C.|last-author-amp=yes | title=Evidence for endothermic ancestors of crocodiles at the stem of archosaur evolution |journal=Physiol. Biochem. Zool. | volume=77 | pages=1051–1067 | year=2004 |doi=10.1086/422766|pmid=15674775|issue=6}}</ref><ref>{{cite journal|author=Summers, A.P. | title=Evolution: Warm-hearted crocs | journal=Nature | volume=434 | pages=833–834 | year=2005 |doi=10.1038/434833a|pmid=15829945|issue=7035}}</ref> It is reasonable to suggest that later crocodilians developed the pulmonary bypass shunt as they became cold-blooded, aquatic, and less active.

If the original crocodilians and other [[Triassic]] archosaurs were warm-blooded, this would help to resolve some evolutionary puzzles:
*The earliest crocodilians, e.g., ''[[Terrestrisuchus]]'', were slim, leggy terrestrial predators whose build suggests a fairly active lifestyle, which requires a fairly fast metabolism. And some other crurotarsan archosaurs appear to have had erect limbs, while those of [[rauisuchia]]ns are very poorly adapted for any other posture. Erect limbs are advantageous for active animals because they avoid [[Carrier's constraint]], but disadvantageous for more sluggish animals because they increase the energy costs of standing up and lying down.
*If early archosaurs were completely cold-blooded and (as seems most likely) [[Physiology of dinosaurs|dinosaurs were at least fairly warm-blooded]], dinosaurs would have had to evolve warm-blooded metabolisms in less than half the time it took for [[synapsida|synapsids]] to do the same.

===Respiratory system===
A recent study of the lungs of the [[American alligator]] has shown that the airflow through them is unidirectional, moving in the same direction during inhalation and exhalation.<ref name=FK10>{{Cite journal |last1=Farmer |first1=C. G. |last2=Sanders |first2=K. |year=2010 |title=Unidirectional airflow in the lungs of alligators |journal=Science |volume=327 |issue=5963 |pages=338–340 |doi=10.1126/science.1180219 |pmid=20075253}}</ref> This is also seen in birds and many non-avian dinosaurs, which have [[Bird anatomy#Respiratory system|air sacs]] to further aid in respiration. Both birds and alligators achieve unidirectional air flow through the presence of [[Bird anatomy#Respiratory system|parabronchi]], which are responsible for gas exchange. The study has found that in alligators, air enters through the second [[Bronchus|bronchial branch]], moves through the parabronchi, and exits through the first bronchial branch. Unidirectional airflow in both birds and alligators suggests that this type of respiration was present in basal Triassic archosaurs and their non-dinosaurian descendants, including phytosaurs, aetosaurs, rauisuchians, crocodylomorphs, and pterosaurs.<ref name=FK10/> The use of unidirectional airflow in the lungs of archosaurs may have given the group an advantage over synapsids, which had lungs where air moved tidally in and out through a network of bronchi that terminated in [[Pulmonary alveolus|alveoli]], which were cul-de-sacs. The better efficiency in gas transfer seen in archosaur lungs may have been advantageous during the times of low atmospheric oxygen which are thought to have existed during the Mesozoic.<ref name=SN>{{cite news |title=Alligators breathe like birds |author=Lisa Grossman |newspaper=Science News |date=January 14, 2010 |url=http://www.sciencenews.org/view/generic/id/54464/title/Alligators_breathe_like_birds |accessdate=January 14, 2010}}</ref>

==Reproduction==
Most archosaurs are [[oviparous]]. Birds and crocodilians lay hard-shelled eggs, as did extinct dinosaurs, and crocodylomorphs. Hard-shelled eggs are present in both dinosaurs and crocodilians, which has been used as an explanation for the absence of [[vivipary]] or [[ovovivipary]] in archosaurs.<ref>{{Cite journal|title=Natural History of Reptilian Development: Constraints on the Evolution of Viviparity |author1=Robin M. Andrews |author2=Tom Mathies |journal=BioScience |year=2000 |volume=50 |issue=3 |pages=227–238 |doi=10.1641/0006-3568(2000)050[0227:NHORDC]2.3.CO;2 }}</ref> However, both pterosaurs<ref>{{cite journal | last1 = Ji | first1 = Q | last2 = Ji | first2 = SA | last3 = Cheng | first3 = YN | display-authors = 3 | last4 = et al | year = | title = "(December 2004). "Palaeontology: pterosaur egg with a leathery shell | url = | journal = Nature | volume = 432 | issue = 7017| page = 572 | doi = 10.1038/432572a | pmid = 15577900 | date=December 2004}}</ref> and [[baurusuchids]]<ref>{{cite journal | last1 = Oliveira | first1 = C.E.M. | last2 = Santucci | first2 = R.M. | last3 = Andrade | first3 = M.B. | last4 = Fulfaro | first4 = V.J. | last5 = Basílo | first5 = J.A.F. | last6 = Benton | first6 = M.J. | year = 2011 | title = Crocodylomorph eggs and eggshells from the Adamantina Formation (Bauru Group), Upper Cretaceous of Brazil | url = | journal = Palaeontology | volume = 54 | issue = 2| pages = 309–321 | doi = 10.1111/j.1475-4983.2010.01028.x }}</ref> have soft-shelled eggs, implying that hard shells are not a plesiomorphic condition. The pelvic anatomy of ''[[Cricosaurus]]'' and other [[metriorhynchidae|metriorhynchids]]<ref>{{Cite journal|last=Herrera|first=Yanina|last2=Fernández|first2=Marta S.|last3=Lamas|first3=Susana G.|last4=Campos|first4=Lisandro|last5=Talevi|first5=Marianella|last6=Gasparini|first6=Zulma|date=2017-02-01|title=Morphology of the sacral region and reproductive strategies of Metriorhynchidae: a counter-inductive approach|url=https://www.cambridge.org/core/journals/earth-and-environmental-science-transactions-of-royal-society-of-edinburgh/article/div-classtitlemorphology-of-the-sacral-region-and-reproductive-strategies-of-metriorhynchidae-a-counter-inductive-approachdiv/AE3A4183C5925CFE4F6F3108B3B2C147|journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh|volume=106|issue=4|pages=247–255|doi=10.1017/S1755691016000165|issn=1755-6910}}</ref> and fossilized embryos belonging to the non-archosaur archosauromorph ''[[Dinocephalosaurus]],''<ref>{{Cite journal|last=Liu|first=Jun|last2=Organ|first2=Chris L.|last3=Benton|first3=Michael J.|last4=Brandley|first4=Matthew C.|last5=Aitchison|first5=Jonathan C.|date=2017-02-14|title=Live birth in an archosauromorph reptile|url=http://www.nature.com/articles/ncomms14445|journal=Nature Communications|language=en|volume=8|doi=10.1038/ncomms14445|issn=2041-1723|page=14445}}</ref> together suggest that the lack of viviparity among archosaurs may be a consequence of lineage-specific restrictions.

Archosaurs are ancestrally [[superprecocial]] as evidenced in various dinosaurs, pterosaurs, and crocodylomorphs.<ref>Mark P. Witton (2013), Pterosaurs: Natural History, Evolution, Anatomy, Princeton University Press, {{ISBN|978-0-691-15061-1}}</ref> However, parental care did evolve independently multiple times in crocodilians, dinosaurs, and [[aetosaur]]s.<ref>Avanzini, M.; Dalla vecchia, F.M.; Mietto, P; Piubelli, D; Preto, N; Rigo, M; Roghi, G (2007). "A vertebrate nesting site in northeastern Italy reveals unexpectedly complex behavior for late Carnian reptiles". PALAIOS. 22 (5): 465–475. {{doi|10.2110/palo.2005.p05-137r}}.</ref> In most such species the animals bury their eggs and rely on [[temperature-dependent sex determination]]. The notable exception are [[Neornithes]] which incubate their eggs and rely on genetic sex determination – a trait that might have given them a survival advantage over other dinosaurs.<ref>Eggshell Porosity Provides Insight on Evolution of Nesting in Dinosaurs, Published: November 25, 2015 https://dx.doi.org/10.1371/journal.pone.0142829
</ref>

==References==
{{reflist|colwidth=30em}}

==Sources==
* {{cite book|authorlink=Michael J. Benton|last=Benton|first=M. J.|year=2004|title=Vertebrate Paleontology|edition=3rd |publisher=Blackwell Science}}
* {{cite book|authorlink=Robert L. Carroll|last=Carroll|first=R. L.|year=1988|title=Vertebrate Paleontology and Evolution'|publisher=W. H. Freeman |location=New York}}

==External links==
* [http://www.ucmp.berkeley.edu/diapsids/archosauria.html UCMP]
* [https://web.archive.org/web/20050405074210/http://palaeos.com/Vertebrates/Units/270Archosauromorpha/270.500.html Paleos] reviews the messy history of archosaur phylogeny (family tree) and has an excellent image of the various archosaur ankle types.
* [https://web.archive.org/web/20050819095217/http://www.fmnh.helsinki.fi/users/haaramo/metazoa/Deuterostoma/Chordata/Archosauria/Archosauria.htm Mikko's Phylogeny Archive] Archosauria

{{Chordata}}
{{Archosauromorpha|state=collapsed}}
{{Archosauriformes|B.}}
{{taxonbar}}
{{portal bar |Dinosaurs|Evolutionary biology|Paleontology}}

[[Category:Archosaurs| ]]
[[Category:Extant Early Triassic first appearances]]
[[Category:Taxa named by Edward Drinker Cope]]

Expedition (book)

2018-01-05T01:28:55Z

Wikkler: /* See also */

{{refimprove|article|date=August 2014}}

{{Infobox book
| name = Expedition
| title_orig =
| translator =
| image = Expedition cover.jpg
| image_size = 250px
| caption = ''Expedition'' book cover
| author = [[Wayne Barlowe|Wayne Douglas Barlowe]]
| illustrator = [[Wayne Barlowe|Wayne Douglas Barlowe]]
| cover_artist = [[Wayne Barlowe|Wayne Douglas Barlowe]]
| country =
| language = English
| series =
| subject = [[Extraterrestrial life]]
| genre = Science fiction, [[speculative evolution]]
| published = 1990 (Workman Publishing Company)
| media_type = Print
| pages =
| isbn = 978-0894809828
| oclc =
| preceded_by =
| followed_by =
}}


'''''Expedition''''' is a [[science fiction]] and [[speculative fiction]] book by artist-author [[Wayne Douglas Barlowe]]. Subtitled "Being an Account in Words and Artwork of the 2358 A.D. Voyage to Darwin IV", it is written as though published in the year 2366, five years after Barlowe's participation in a voyage to an alien planet, dubbed [[Darwin IV]] in honor of [[Charles Darwin]].

In the [[24th century]] the exploitation of the Earth's ecosystem has created an environment so toxic that mass extinctions have wiped out nearly half of its animal population. Most of the remaining fauna, save humans, have suffered horrible mutation. Aided by the benevolent and technologically superior alien race, the Yma, humanity begins to repair their ravaged world while simultaneously learning more about the universe around them. When an unmanned Yma probe discovers evidence of alien life on another planet, the titular "expedition" is sent to investigate.

Barlowe writes as a sort of 24th century [[John James Audubon|Audubon]], presenting his findings in a collection of paintings, sketches, field notes, and diary entries from his explorations of Darwin IV. He details a bewildering variety of [[Extraterrestrial life|alien lifeforms]] such as Gyrosprinters, Arrowtongues, Grovebacks, Daggerwrists, Skewers, Emperor Sea Striders, and Eosapians. Unlike the aliens presented in much of popular science fiction, which often seem to be variations of terrestrial lifeforms, Barlowe's creatures are truly alien: none of them possess eyes or true jaws; their body structures are often unlike any found on Earth; and they have unique modes of locomotion, sensing, and eating. Very late in the expedition, the explorer encounters lifeforms which use tools (the Eosapiens), giving a very strong indication they are intelligent.

A conservationist theme is present throughout the book. The expedition is designed to have as minimal an impact as possible on Darwin IV's environment. When two of the expedition's members suffer a fatal accident, Yma technology is used to remove all traces of the accident from Darwin IV's environment. At the conclusion of the expedition, Darwin IV is left in the same pristine state it was in prior to the expedition, with the exception of a metal obelisk placed in a remote area by the expedition.

== Television ==
The [[Discovery Channel]] produced a television special adapted from Barlowe's ''Expedition'', entitled ''[[Alien Planet]]'',<ref>http://www.imdb.com/title/tt0453446/</ref> which first aired on May 14, 2005. This program was faithful to the book in its presentation of the lifeforms found on Darwin IV. However, instead of being presented as the artist's own experiences, the program is presented as the findings of two autonomous robotic probes.

== See also ==
*''[[Aurelia and Blue Moon|Alien Worlds ''or'' Extraterrestrial]]'', a similar program aired on Channel 4 and the National Geographic Channel

== References ==
{{Reflist}}

[[Category:1990 books]]
[[Category:24th century in fiction]]
[[Category:Books about extraterrestrial life]]
[[Category:Books by Wayne Barlowe]]
[[Category:Science fiction books]]
[[Category:Speculative evolution]]

{{1990s-sf-novel-stub}}

Caenagnathidae

2017-12-16T22:30:52Z

Wikkler: There's also a distinct possibility that Nomingia is a caenagnathid as well. It comes up as caenagnathid sometimes. Cladistics is messy.

{{automatic Taxobox
| name = Caenagnathids
| fossil_range = [[Late Cretaceous]], {{fossil range|91|65|}}
| image = Chirostenotes skull.jpg
| image_caption = Reconstructed skull of ''[[Anzu wyliei]]''
| authority = [[Charles Hazelius Sternberg|Sternberg]], 1940
| type_species = ''[[Caenagnathus collinsi]]''
| type_species_authority = Sternberg, 1940
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
*{{extinct}}''[[Beibeilong]]''
*{{extinct}}''[[Epichirostenotes]]''
*{{extinct}}''[[Gigantoraptor]]''?
*{{extinct}}''[[Hagryphus]]''
*{{extinct}}''[[Microvenator]]''?
*{{extinct}}''[[Nomingia]]''?
*{{extinct}}''[[Ojoraptorsaurus]]''
*{{extinct}}'''Caenagnathinae'''
**{{extinct}}''[[Anzu wyliei|Anzu]]''
**{{extinct}}''[[Caenagnathus]]''
*{{extinct}}'''Elmisaurinae'''
**{{extinct}}''[[Apatoraptor]]''
**{{extinct}}''[[Caenagnathasia]]''
**{{extinct}}''[[Chirostenotes]]''
**{{extinct}}''[[Elmisaurus]]''
**{{extinct}}''[[Leptorhynchos (dinosaur)|Leptorhynchos]]''
| synonyms =
* '''Elmisauridae''' [[Halszka Osmólska|Osmólska]], 1981
}}
'''Caenagnathidae''' is a family of bird-like [[maniraptora]]n [[theropod]] [[dinosaur]]s from the Late Cretaceous of North America and Asia. They are a member of the [[Oviraptorosauria]], and close relatives of the [[Oviraptoridae]].<ref>Osmólska, H., P. J. Currie, et al. (2004). Oviraptorosauria. The Dinosauria. D. B. Weishampel, P. Dodson and H. Osmolska. Berkeley, University of California Press: 165-183.</ref> Like other oviraptorosaurs, caenagnathids had specialized beaks,<ref name="Currie, P. J. 1993">{{cite journal | last1 = Currie | first1 = P. J. | last2 = Godfrey | first2 = S. J. | display-authors = 2 | last3 = et al | year = 1993 | title = New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia | url = | journal = Canadian Journal of Earth Sciences | volume = 30 | issue = 10-11| pages = 2255–2272 | doi=10.1139/e93-196}}</ref> long necks,<ref name="Sues, H. D. 1997">{{cite journal | last1 = Sues | first1 = H. D. | year = 1997 | title = On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from western North America | url = | journal = Journal of Vertebrate Paleontology | volume = 17 | issue = 4| pages = 698–716 | doi=10.1080/02724634.1997.10011018}}</ref> and short tails,<ref name = "barsboldetal2000">{{cite journal | last1 = Barsbold | first1 = R. | last2 = Osmolska | first2 = H. | last3 = Watabe | first3 = M. | last4 = Currie | first4 = P. J. | last5 = Tsogtbaatar | first5 = K. | year = 2000 | title = New oviraptorosaur (Dinosauria, Theropoda) from Mongolia: The first dinosaur with a pygostyle | url = http://www.app.pan.pl/archive/published/app45/app45-097.pdf | format=PDF| journal = Acta Palaeontologica Polonica | volume = 45 | issue = 2| pages = 97–106 }}</ref> and would have been covered in feathers. The relationships of caenagnathids were long a puzzle. The family was originally named by [[Charles Hazelius Sternberg]] in 1940 <ref name = "sternberg1940">{{cite journal | last1 = Sternberg | first1 = R.M. | year = 1940 | title = A toothless bird from the Cretaceous of Alberta | url = | journal = Journal of Paleontology | volume = 14 | issue = 1| pages = 81–85 }}</ref> as a family of flightless birds. The discovery of skeletons of the related oviraptorids revealed that they were in fact non-avian theropods,<ref>{{cite journal | last1 = Osmólska | first1 = H | year = 1976 | title = New light on the skull anatomy and systematic position of Oviraptor | url = | journal = Nature | volume = 262 | issue = | pages = 683–684 | doi=10.1038/262683a0}}</ref> and the discovery of more complete caenagnathid remains <ref name="Sues, H. D. 1997"/><ref name = "currierussell1988">{{cite journal | last1 = Currie | first1 = P.J. | last2 = Russell | first2 = D.A. | year = 1988 | title = Osteology and relationships of ''Chirostenotes pergracilis'' (Saurischia, Theropoda) from the Judith River Oldman Formation of Alberta | doi = 10.1139/e88-097 | journal = Canadian Journal of Earth Sciences | volume = 25 | issue = 3| pages = 972–986 }}</ref> revealed that ''Chirostenotes pergracilis'', originally named on the basis of a pair of hands, and ''"Ornithomimus" elegans'', named from a foot, were caenagnathids as well.

==Anatomy==
[[File:Oviraptorid Clean.png|thumb|left|''Anzu wyliei'' skeleton cast in the [[Rocky Mountain Dinosaur Resource Center]] in Woodland Park, Colorado, USA.]]
Overall, the anatomy of the caenagnathids is similar to that of the closely related Oviraptoridae, but there are a number of differences. In particular, caenagnathid jaws exhibited a distinct suite of specializations not seen in other oviraptorosaurs. Compared to the oviraptorids, the jaws tended to be relatively long and shallow, suggesting that the bite was not as powerful. The inside of the lower jaws also bore a complex series of ridges and toothlike processes, as well as a pair of horizontal, shelf-like structures. Furthermore, the jaws were unusual in being hollow and air filled, apparently being connected to the air sac system.<ref name="Currie, P. J. 1993"/> Caenagnathids also tended to be more lightly built than the oviraptorids. They had slender arms and long, gracile legs,<ref name="currierussell1988"/> although they lacked the extreme cursorial specializations seen in avimimids and ''Caudipteryx''.

==Etymology==
The name ''Caenagnathus'' (and hence Caenagnathidae) means "recent jaws"—when first discovered, it was thought that caenagnathids were close relatives of [[paleognathae|paleognath]] birds (such as the [[ostrich]]) based on features of the lower jaw. Since it would be unusual to find a recent group of birds in the Cretaceous, the name "recent jaws" was applied. Most paleontologists, however, now think that the birdlike features of the jaw were acquired [[convergent evolution|convergently]] with modern birds.<ref name = "cracraft1971">{{cite journal | last1 = Cracraft | first1 = J. | year = 1971 | title = Caenagnathiformes: Cretaceous birds convergent in jaw mechanism to dicynodont reptiles | url = | journal = Journal of Paleontology | volume = 45 | issue = | pages = 805–809 }}</ref><ref name = "barsboldetal1990">[[Rinchen Barsbold|Barsbold, R.]], [[Teresa Maryańska|Maryańska, T.]], and Osmólska, H. (1990). "Oviraptorosauria." pg. 249-258 ''in'' Weishampel, Dodson, and Osmolska (eds.) ''The Dinosauria'', University of California Press (Berkeley).</ref>

==Evolution==
The earliest known caenagnathid is ''Caenagnathasia martinsoni'', from the Turonian Bissekty Formation of Uzbekistan.<ref name="Currie, P. J. 1993"/> The jaw of ''Caenagnathasia'' already has the specialized ridges and crushing surfaces seen in later forms. This suggests that caenagnathids originated well before the Turonian, but currently, there are no Early Cretaceous caenagnathid fossils. Caenagnathids later appeared in western North America, during the Campanian, suggesting that they may have originated in Asia, then migrated into western North America. Caenagnathids showed considerable variation in form. The tiny jaws of ''Caenagnathasia'' suggest a small animal, perhaps the size of a turkey. An unnamed species from the Hell Creek Formation <ref name="Currie, P. J. 1993"/> indicates a much larger animal, considerably larger than a human. If ''Gigantoraptor erlianensis'' is a caenagnathid, then it would represent far and away the largest member of the group, measuring up to 8 meters in length and weighing up to 1.4 tons. Their beaks also show considerable variation; that of ''Caenagnathasia'' is relatively short and deep, while that of ''Caenagnathus'' is long and shovel-shaped. This variation in size and beak shape suggests that caenagnathids evolved to exploit a range of ecological niches. Caenagnathids persisted up until the end of the Cretaceous period, as shown by the presence of at least two species in the late Maastrichtian Hell Creek Formation, before vanishing at the end of the Cretaceous along with all other non-avian dinosaurs.

==Classification==
The family Caenagnathidae, together with its sister group the [[Oviraptoridae]], comprises the superfamily [[Caenagnathoidea]]. In [[phylogenetic taxonomy]], the [[clade]] Caenagnathidae is defined as the most inclusive group containing ''[[Chirostenotes|Chirostenotes pergracilis]]'' but not ''[[Oviraptor|Oviraptor philoceratops]]''. While before 2010s only about two to six species were commonly recognized as belonging to the Caenagnathidae, currently that number may be much greater, with new discoveries and theories about older species that may inflate this number to up to ten. Much of this historical difference centers on the first caenagnathid to be described, ''Chirostenotes pergracilis''. Due to the poor preservation of most caenagnathid remains and resulting misidentifications, different bones and different specimens of ''Chirostenotes'' have historically been assigned to a number of different species. For example, the feet of one species, named ''Macrophalangia canadensis'',<ref name = "sternberg1932">{{cite journal | last1 = Sternberg | first1 = C. H. | year = 1932 | title = Two new theropod dinosaurs from the Belly River Formation of Alberta | url = | journal = The Canadian Field-Naturalist | volume = 46 | issue = | pages = 99–105 }}</ref> were known from the same region from which ''Chirostenotes pergracilis'' was recovered, but the discovery of a new specimen with both hands and feet preserved<ref name="currierussell1988"/> provided the support to combine them, while the later discovery of a partial skull with hands and feet <ref name="Sues, H. D. 1997"/> suggested that ''Chirostenotes'' and ''Caenagnathus'' were the same animal, and current studies of caenagnathid relationships continue to find them as closely related genera.<ref name=leptorhynchos>{{Cite journal | last1 = Longrich | first1 = N. R. | last2 = Barnes | first2 = K. | last3 = Clark | first3 = S. | last4 = Millar | first4 = L. | title = Caenagnathidae from the Upper Campanian Aguja Formation of West Texas, and a Revision of the Caenagnathinae | doi = 10.3374/014.054.0102 | journal = Bulletin of the Peabody Museum of Natural History | volume = 54 | pages = 23–49 | year = 2013 | pmid = | pmc = }}</ref>
[[File:Caenagnathidae.jpg|thumb|Caenagnathid skeletons to scale]]
Longrich and colleagues (2013) defined a subgroup of Caenagnathidae, the '''Caenagnathinae''', as all caenagnathids more closely related to ''Caenagnathus collinsi'' than to ''Caenagnathasia martinsoni'' or ''Elmisaurus rarus''.<ref name=leptorhynchos/> In 2015, the group '''Elmisaurinae''' was defined, including all species more closely related to ''Elmisaurus rarus'' than to ''Caenagnathus collinsi''.<ref>Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.</ref><ref name="currie15">{{cite journal|last=Currie|first=P.J.|authorlink=Philip J. Currie|last2=Funston|first2=G.F.|last3=Osmólska|first3=H.†|authorlink3=Halszka Osmólska|year=2015|title=New specimens of the crested theropod dinosaur ''Elmisaurus rarus'' from Mongolia|url=http://app.pan.pl/archive/published/app60/app001302014_acc.pdf|journal=Acta Palaeontologica Polonica|volume=XX|issue=XX|pages=XXX–XXX|doi=10.4202/app.00130.2014}}</ref>

The [[cladogram]] below follows an analysis by Funston & Currie in 2016.<ref name=apatoraptor1>{{cite journal |author1=Gregory F. Funston |author2=Philip J. Currie |year=2016 |title=A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae |journal=Journal of Vertebrate Paleontology |volume=Online edition |issue= |pages=e1160910 |doi=10.1080/02724634.2016.1160910 }}</ref>

{{clade| style=font-size:100%;line-height:100%
|label1=Caenagnathidae
|1={{clade
|1=''[[Microvenator celer]]''
|label2=unnamed
|2={{clade
|1=''[[Gigantoraptor erlianensis]]''
|label2=unnamed
|2={{clade
|1=''[[Hagryphus giganteus]]''
|2={{clade
|1={{clade
|1=''[[Epichirostenotes curriei]]''
|2=''[[Anzu wyliei]]'' }}
|2={{clade
|1=''[[Caenagnathus collinsi]]''
|label2='''Elmisaurinae'''
|2={{clade
|1=''[[Caenagnathasia martinsoni]]''
|2={{clade
|1=''[[Chirostenotes pergracilis]]''
|2={{clade
|1=''[[Leptorhynchos elegans]]''
|2={{clade
|1=''[[Apatoraptor pennatus]]''
|2=''[[Elmisaurus rarus]]''
}} }} }} }} }} }} }} }} }} }}

===Species===
Roughly a dozen caenagnathid species have been named, but it remains unclear how many are valid. Many species are known from fragmentary remains, such as jaws, hands, or feet, making comparisons between them difficult. ''Caenagnathus sternbergi'', for example, was described on the basis of a jaw bone. It has been interpreted as either the jaws of ''Chirostenotes pergracilis'' (described on the basis of a pair of hands) or ''Chirostenotes elegans''<ref name="Sues, H. D. 1997"/> (described on the basis of a foot), but because no complete skeleton is known, it is difficult to be certain which animal it belongs to. The relationships of other species remain in doubt. ''Gigantoraptor'' was originally interpreted as an oviraptorid, but may in fact represent a primitive caenagnathid.<ref name="Longrichetal10">{{cite journal |year=2010 |title=A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia |journal=Palaeontology |volume=53 |issue=5 |pages=945–960 |doi=10.1111/j.1475-4983.2010.00968.x |url=http://onlinelibrary.wiley.com/doi/10.1111/j.1475-4983.2010.00968.x/abstract|author1=Nicholas R. Longrich |author2=Philip J. Currie |author3=Dong Zhi-Ming }}</ref>

* ''[[Anzu wyliei]]'' - ([[Hell Creek Formation]], Montana, United States)<ref name="anzu">{{Cite journal | doi = 10.1371/journal.pone.0092022| title = A New Large-Bodied Oviraptorosaurian Theropod Dinosaur from the Latest Cretaceous of Western North America| journal = PLoS ONE| volume = 9| issue = 3| pages = e92022| year = 2014| last1 = Lamanna | first1 = M. C. | last2 = Sues | first2 = H. D. | last3 = Schachner | first3 = E. R. | last4 = Lyson | first4 = T. R. | pmid=24647078 | pmc=3960162}}</ref><ref name=triebold>Varricchio, D. J. (2001). Late Cretaceous Oviraptorosaur (Theropoda) dinosaurs from Montana. Mesozoic Vertebrate Life. D. H. Tanke and K. Carpenter. Bloomington, Indiana University Press: 42-57.</ref>
* [[Apatoraptor|''Apatoraptor pennatus'']] - ([[Horseshoe Canyon Formation]], Alberta)
* ''[[Caenagnathasia martinsoni]]'' - ([[Bissekty Formation]], Mongolia)
* ''[[Chirostenotes pergracilis]]'' - ([[Dinosaur Park Formation]], Alberta, Canada)
* ''[[Caenagnathus collinsi]]'' - ([[Dinosaur Park Formation]], Alberta, Canada)
* ''[[Elmisaurus rarus]]'' - ([[Nemegt Formation]], Mongolia)
* ''[[Epichirostenotes curriei]]'' - ([[Horseshoe Canyon Formation]], Alberta, Canada)
* ''[[Gigantoraptor erlianensis]]'' - ([[Iren Dabasu Formation]], Inner Mongolia, China)
* ''[[Hagryphus giganteus]]'' - ([[Kaiparowits Formation]], Utah, United States)
* ''[[Leptorhynchos elegans]]'' - ([[Dinosaur Park Formation]], Alberta, Canada)
* ''[[Leptorhynchos gaddisi]]'' - ([[Aguja Formation]], Texas, United States)<ref name=leptorhynchos/>
* ''[[Ojoraptorsaurus boerei]]'' - ([[Ojo Alamo Formation]], New Mexico, United States)

Caenagnathids are only known from the Late Cretaceous of North America and Asia. The earliest and most primitive known caenagnathid is ''[[Caenagnathasia|Caenagnathasia martinsoni]]'', from the Bissekty Formation of Uzbekistan.<ref name = "CurrieGodfreyNesov1994">{{cite journal | last1 = Currie | first1 = P.J. | last2 = Godfrey | first2 = S.J. | last3 = Nesov | first3 = L.A. | year = 1994 | title = New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia | url = | journal = Canadian Journal of Earth Sciences | volume = 30 | issue = | pages = 2255–2272 | doi=10.1139/e93-196}}</ref> The largest is the enormous ''Gigantoraptor erlianensis''.<ref name="Longrichetal10"/>

==See also==
{{Portal|Dinosaurs}}
* [[Timeline of oviraptorosaur research]]

==References==
{{Reflist}}

==External links==
* [http://qilong.8m.com/Caenagnathidae.html Overview of Caenagnathidae by Jaime Headden.]
* [https://web.archive.org/web/20060827031308/http://www.carnegiemnh.org/ditw/ovi/res_mount.htm Photo of the Triebold caenagnathid, on display at the Carnegie Museum].
* [https://web.archive.org/web/20070312081952/http://www.skeletaldrawing.com/psgallery/pages/chirostenotes.html Skeletal reconstruction of the Triebold specimens.]

{{Oviraptorosauria}}

[[Category:Oviraptorosaurs]]
[[Category:Caenagnathids]]

Caenagnathidae

2017-12-15T21:05:18Z

Wikkler: True, but it's enough. I'm not sure why when I'm adding uncertainty because of many studies it gets reverted yet when Chilesaurus gets recovered in ONE paper as an Ornithischian it goes unquestioned.

{{automatic Taxobox
| name = Caenagnathids
| fossil_range = [[Late Cretaceous]], {{fossil range|91|65|}}
| image = Chirostenotes skull.jpg
| image_caption = Reconstructed skull of ''[[Anzu wyliei]]''
| authority = [[Charles Hazelius Sternberg|Sternberg]], 1940
| type_species = ''[[Caenagnathus collinsi]]''
| type_species_authority = Sternberg, 1940
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
*{{extinct}}''[[Beibeilong]]''
*{{extinct}}''[[Epichirostenotes]]''
*{{extinct}}''[[Gigantoraptor]]''?
*{{extinct}}''[[Hagryphus]]''
*{{extinct}}''[[Microvenator]]''?
*{{extinct}}''[[Ojoraptorsaurus]]''
*{{extinct}}'''Caenagnathinae'''
**{{extinct}}''[[Anzu wyliei|Anzu]]''
**{{extinct}}''[[Caenagnathus]]''
*{{extinct}}'''Elmisaurinae'''
**{{extinct}}''[[Apatoraptor]]''
**{{extinct}}''[[Caenagnathasia]]''
**{{extinct}}''[[Chirostenotes]]''
**{{extinct}}''[[Elmisaurus]]''
**{{extinct}}''[[Leptorhynchos (dinosaur)|Leptorhynchos]]''
| synonyms =
* '''Elmisauridae''' [[Halszka Osmólska|Osmólska]], 1981
}}
'''Caenagnathidae''' is a family of bird-like [[maniraptora]]n [[theropod]] [[dinosaur]]s from the Late Cretaceous of North America and Asia. They are a member of the [[Oviraptorosauria]], and close relatives of the [[Oviraptoridae]].<ref>Osmólska, H., P. J. Currie, et al. (2004). Oviraptorosauria. The Dinosauria. D. B. Weishampel, P. Dodson and H. Osmolska. Berkeley, University of California Press: 165-183.</ref> Like other oviraptorosaurs, caenagnathids had specialized beaks,<ref name="Currie, P. J. 1993">{{cite journal | last1 = Currie | first1 = P. J. | last2 = Godfrey | first2 = S. J. | display-authors = 2 | last3 = et al | year = 1993 | title = New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia | url = | journal = Canadian Journal of Earth Sciences | volume = 30 | issue = 10-11| pages = 2255–2272 | doi=10.1139/e93-196}}</ref> long necks,<ref name="Sues, H. D. 1997">{{cite journal | last1 = Sues | first1 = H. D. | year = 1997 | title = On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from western North America | url = | journal = Journal of Vertebrate Paleontology | volume = 17 | issue = 4| pages = 698–716 | doi=10.1080/02724634.1997.10011018}}</ref> and short tails,<ref name = "barsboldetal2000">{{cite journal | last1 = Barsbold | first1 = R. | last2 = Osmolska | first2 = H. | last3 = Watabe | first3 = M. | last4 = Currie | first4 = P. J. | last5 = Tsogtbaatar | first5 = K. | year = 2000 | title = New oviraptorosaur (Dinosauria, Theropoda) from Mongolia: The first dinosaur with a pygostyle | url = http://www.app.pan.pl/archive/published/app45/app45-097.pdf | format=PDF| journal = Acta Palaeontologica Polonica | volume = 45 | issue = 2| pages = 97–106 }}</ref> and would have been covered in feathers. The relationships of caenagnathids were long a puzzle. The family was originally named by [[Charles Hazelius Sternberg]] in 1940 <ref name = "sternberg1940">{{cite journal | last1 = Sternberg | first1 = R.M. | year = 1940 | title = A toothless bird from the Cretaceous of Alberta | url = | journal = Journal of Paleontology | volume = 14 | issue = 1| pages = 81–85 }}</ref> as a family of flightless birds. The discovery of skeletons of the related oviraptorids revealed that they were in fact non-avian theropods,<ref>{{cite journal | last1 = Osmólska | first1 = H | year = 1976 | title = New light on the skull anatomy and systematic position of Oviraptor | url = | journal = Nature | volume = 262 | issue = | pages = 683–684 | doi=10.1038/262683a0}}</ref> and the discovery of more complete caenagnathid remains <ref name="Sues, H. D. 1997"/><ref name = "currierussell1988">{{cite journal | last1 = Currie | first1 = P.J. | last2 = Russell | first2 = D.A. | year = 1988 | title = Osteology and relationships of ''Chirostenotes pergracilis'' (Saurischia, Theropoda) from the Judith River Oldman Formation of Alberta | doi = 10.1139/e88-097 | journal = Canadian Journal of Earth Sciences | volume = 25 | issue = 3| pages = 972–986 }}</ref> revealed that ''Chirostenotes pergracilis'', originally named on the basis of a pair of hands, and ''"Ornithomimus" elegans'', named from a foot, were caenagnathids as well.

==Anatomy==
[[File:Oviraptorid Clean.png|thumb|left|''Anzu wyliei'' skeleton cast in the [[Rocky Mountain Dinosaur Resource Center]] in Woodland Park, Colorado, USA.]]
Overall, the anatomy of the caenagnathids is similar to that of the closely related Oviraptoridae, but there are a number of differences. In particular, caenagnathid jaws exhibited a distinct suite of specializations not seen in other oviraptorosaurs. Compared to the oviraptorids, the jaws tended to be relatively long and shallow, suggesting that the bite was not as powerful. The inside of the lower jaws also bore a complex series of ridges and toothlike processes, as well as a pair of horizontal, shelf-like structures. Furthermore, the jaws were unusual in being hollow and air filled, apparently being connected to the air sac system.<ref name="Currie, P. J. 1993"/> Caenagnathids also tended to be more lightly built than the oviraptorids. They had slender arms and long, gracile legs,<ref name="currierussell1988"/> although they lacked the extreme cursorial specializations seen in avimimids and ''Caudipteryx''.

==Etymology==
The name ''Caenagnathus'' (and hence Caenagnathidae) means "recent jaws"—when first discovered, it was thought that caenagnathids were close relatives of [[paleognathae|paleognath]] birds (such as the [[ostrich]]) based on features of the lower jaw. Since it would be unusual to find a recent group of birds in the Cretaceous, the name "recent jaws" was applied. Most paleontologists, however, now think that the birdlike features of the jaw were acquired [[convergent evolution|convergently]] with modern birds.<ref name = "cracraft1971">{{cite journal | last1 = Cracraft | first1 = J. | year = 1971 | title = Caenagnathiformes: Cretaceous birds convergent in jaw mechanism to dicynodont reptiles | url = | journal = Journal of Paleontology | volume = 45 | issue = | pages = 805–809 }}</ref><ref name = "barsboldetal1990">[[Rinchen Barsbold|Barsbold, R.]], [[Teresa Maryańska|Maryańska, T.]], and Osmólska, H. (1990). "Oviraptorosauria." pg. 249-258 ''in'' Weishampel, Dodson, and Osmolska (eds.) ''The Dinosauria'', University of California Press (Berkeley).</ref>

==Evolution==
The earliest known caenagnathid is ''Caenagnathasia martinsoni'', from the Turonian Bissekty Formation of Uzbekistan.<ref name="Currie, P. J. 1993"/> The jaw of ''Caenagnathasia'' already has the specialized ridges and crushing surfaces seen in later forms. This suggests that caenagnathids originated well before the Turonian, but currently, there are no Early Cretaceous caenagnathid fossils. Caenagnathids later appeared in western North America, during the Campanian, suggesting that they may have originated in Asia, then migrated into western North America. Caenagnathids showed considerable variation in form. The tiny jaws of ''Caenagnathasia'' suggest a small animal, perhaps the size of a turkey. An unnamed species from the Hell Creek Formation <ref name="Currie, P. J. 1993"/> indicates a much larger animal, considerably larger than a human. If ''Gigantoraptor erlianensis'' is a caenagnathid, then it would represent far and away the largest member of the group, measuring up to 8 meters in length and weighing up to 1.4 tons. Their beaks also show considerable variation; that of ''Caenagnathasia'' is relatively short and deep, while that of ''Caenagnathus'' is long and shovel-shaped. This variation in size and beak shape suggests that caenagnathids evolved to exploit a range of ecological niches. Caenagnathids persisted up until the end of the Cretaceous period, as shown by the presence of at least two species in the late Maastrichtian Hell Creek Formation, before vanishing at the end of the Cretaceous along with all other non-avian dinosaurs.

==Classification==
The family Caenagnathidae, together with its sister group the [[Oviraptoridae]], comprises the superfamily [[Caenagnathoidea]]. In [[phylogenetic taxonomy]], the [[clade]] Caenagnathidae is defined as the most inclusive group containing ''[[Chirostenotes|Chirostenotes pergracilis]]'' but not ''[[Oviraptor|Oviraptor philoceratops]]''. While before 2010s only about two to six species were commonly recognized as belonging to the Caenagnathidae, currently that number may be much greater, with new discoveries and theories about older species that may inflate this number to up to ten. Much of this historical difference centers on the first caenagnathid to be described, ''Chirostenotes pergracilis''. Due to the poor preservation of most caenagnathid remains and resulting misidentifications, different bones and different specimens of ''Chirostenotes'' have historically been assigned to a number of different species. For example, the feet of one species, named ''Macrophalangia canadensis'',<ref name = "sternberg1932">{{cite journal | last1 = Sternberg | first1 = C. H. | year = 1932 | title = Two new theropod dinosaurs from the Belly River Formation of Alberta | url = | journal = The Canadian Field-Naturalist | volume = 46 | issue = | pages = 99–105 }}</ref> were known from the same region from which ''Chirostenotes pergracilis'' was recovered, but the discovery of a new specimen with both hands and feet preserved<ref name="currierussell1988"/> provided the support to combine them, while the later discovery of a partial skull with hands and feet <ref name="Sues, H. D. 1997"/> suggested that ''Chirostenotes'' and ''Caenagnathus'' were the same animal, and current studies of caenagnathid relationships continue to find them as closely related genera.<ref name=leptorhynchos>{{Cite journal | last1 = Longrich | first1 = N. R. | last2 = Barnes | first2 = K. | last3 = Clark | first3 = S. | last4 = Millar | first4 = L. | title = Caenagnathidae from the Upper Campanian Aguja Formation of West Texas, and a Revision of the Caenagnathinae | doi = 10.3374/014.054.0102 | journal = Bulletin of the Peabody Museum of Natural History | volume = 54 | pages = 23–49 | year = 2013 | pmid = | pmc = }}</ref>
[[File:Caenagnathidae.jpg|thumb|Caenagnathid skeletons to scale]]
Longrich and colleagues (2013) defined a subgroup of Caenagnathidae, the '''Caenagnathinae''', as all caenagnathids more closely related to ''Caenagnathus collinsi'' than to ''Caenagnathasia martinsoni'' or ''Elmisaurus rarus''.<ref name=leptorhynchos/> In 2015, the group '''Elmisaurinae''' was defined, including all species more closely related to ''Elmisaurus rarus'' than to ''Caenagnathus collinsi''.<ref>Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.</ref><ref name="currie15">{{cite journal|last=Currie|first=P.J.|authorlink=Philip J. Currie|last2=Funston|first2=G.F.|last3=Osmólska|first3=H.†|authorlink3=Halszka Osmólska|year=2015|title=New specimens of the crested theropod dinosaur ''Elmisaurus rarus'' from Mongolia|url=http://app.pan.pl/archive/published/app60/app001302014_acc.pdf|journal=Acta Palaeontologica Polonica|volume=XX|issue=XX|pages=XXX–XXX|doi=10.4202/app.00130.2014}}</ref>

The [[cladogram]] below follows an analysis by Funston & Currie in 2016.<ref name=apatoraptor1>{{cite journal |author1=Gregory F. Funston |author2=Philip J. Currie |year=2016 |title=A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae |journal=Journal of Vertebrate Paleontology |volume=Online edition |issue= |pages=e1160910 |doi=10.1080/02724634.2016.1160910 }}</ref>

{{clade| style=font-size:100%;line-height:100%
|label1=Caenagnathidae
|1={{clade
|1=''[[Microvenator celer]]''
|label2=unnamed
|2={{clade
|1=''[[Gigantoraptor erlianensis]]''
|label2=unnamed
|2={{clade
|1=''[[Hagryphus giganteus]]''
|2={{clade
|1={{clade
|1=''[[Epichirostenotes curriei]]''
|2=''[[Anzu wyliei]]'' }}
|2={{clade
|1=''[[Caenagnathus collinsi]]''
|label2='''Elmisaurinae'''
|2={{clade
|1=''[[Caenagnathasia martinsoni]]''
|2={{clade
|1=''[[Chirostenotes pergracilis]]''
|2={{clade
|1=''[[Leptorhynchos elegans]]''
|2={{clade
|1=''[[Apatoraptor pennatus]]''
|2=''[[Elmisaurus rarus]]''
}} }} }} }} }} }} }} }} }} }}

===Species===
Roughly a dozen caenagnathid species have been named, but it remains unclear how many are valid. Many species are known from fragmentary remains, such as jaws, hands, or feet, making comparisons between them difficult. ''Caenagnathus sternbergi'', for example, was described on the basis of a jaw bone. It has been interpreted as either the jaws of ''Chirostenotes pergracilis'' (described on the basis of a pair of hands) or ''Chirostenotes elegans''<ref name="Sues, H. D. 1997"/> (described on the basis of a foot), but because no complete skeleton is known, it is difficult to be certain which animal it belongs to. The relationships of other species remain in doubt. ''Gigantoraptor'' was originally interpreted as an oviraptorid, but may in fact represent a primitive caenagnathid.<ref name="Longrichetal10">{{cite journal |year=2010 |title=A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia |journal=Palaeontology |volume=53 |issue=5 |pages=945–960 |doi=10.1111/j.1475-4983.2010.00968.x |url=http://onlinelibrary.wiley.com/doi/10.1111/j.1475-4983.2010.00968.x/abstract|author1=Nicholas R. Longrich |author2=Philip J. Currie |author3=Dong Zhi-Ming }}</ref>

* ''[[Anzu wyliei]]'' - ([[Hell Creek Formation]], Montana, United States)<ref name="anzu">{{Cite journal | doi = 10.1371/journal.pone.0092022| title = A New Large-Bodied Oviraptorosaurian Theropod Dinosaur from the Latest Cretaceous of Western North America| journal = PLoS ONE| volume = 9| issue = 3| pages = e92022| year = 2014| last1 = Lamanna | first1 = M. C. | last2 = Sues | first2 = H. D. | last3 = Schachner | first3 = E. R. | last4 = Lyson | first4 = T. R. | pmid=24647078 | pmc=3960162}}</ref><ref name=triebold>Varricchio, D. J. (2001). Late Cretaceous Oviraptorosaur (Theropoda) dinosaurs from Montana. Mesozoic Vertebrate Life. D. H. Tanke and K. Carpenter. Bloomington, Indiana University Press: 42-57.</ref>
* [[Apatoraptor|''Apatoraptor pennatus'']] - ([[Horseshoe Canyon Formation]], Alberta)
* ''[[Caenagnathasia martinsoni]]'' - ([[Bissekty Formation]], Mongolia)
* ''[[Chirostenotes pergracilis]]'' - ([[Dinosaur Park Formation]], Alberta, Canada)
* ''[[Caenagnathus collinsi]]'' - ([[Dinosaur Park Formation]], Alberta, Canada)
* ''[[Elmisaurus rarus]]'' - ([[Nemegt Formation]], Mongolia)
* ''[[Epichirostenotes curriei]]'' - ([[Horseshoe Canyon Formation]], Alberta, Canada)
* ''[[Gigantoraptor erlianensis]]'' - ([[Iren Dabasu Formation]], Inner Mongolia, China)
* ''[[Hagryphus giganteus]]'' - ([[Kaiparowits Formation]], Utah, United States)
* ''[[Leptorhynchos elegans]]'' - ([[Dinosaur Park Formation]], Alberta, Canada)
* ''[[Leptorhynchos gaddisi]]'' - ([[Aguja Formation]], Texas, United States)<ref name=leptorhynchos/>
* ''[[Ojoraptorsaurus boerei]]'' - ([[Ojo Alamo Formation]], New Mexico, United States)

Caenagnathids are only known from the Late Cretaceous of North America and Asia. The earliest and most primitive known caenagnathid is ''[[Caenagnathasia|Caenagnathasia martinsoni]]'', from the Bissekty Formation of Uzbekistan.<ref name = "CurrieGodfreyNesov1994">{{cite journal | last1 = Currie | first1 = P.J. | last2 = Godfrey | first2 = S.J. | last3 = Nesov | first3 = L.A. | year = 1994 | title = New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia | url = | journal = Canadian Journal of Earth Sciences | volume = 30 | issue = | pages = 2255–2272 | doi=10.1139/e93-196}}</ref> The largest is the enormous ''Gigantoraptor erlianensis''.<ref name="Longrichetal10"/>

==See also==
{{Portal|Dinosaurs}}
* [[Timeline of oviraptorosaur research]]

==References==
{{Reflist}}

==External links==
* [http://qilong.8m.com/Caenagnathidae.html Overview of Caenagnathidae by Jaime Headden.]
* [https://web.archive.org/web/20060827031308/http://www.carnegiemnh.org/ditw/ovi/res_mount.htm Photo of the Triebold caenagnathid, on display at the Carnegie Museum].
* [https://web.archive.org/web/20070312081952/http://www.skeletaldrawing.com/psgallery/pages/chirostenotes.html Skeletal reconstruction of the Triebold specimens.]

{{Oviraptorosauria}}

[[Category:Oviraptorosaurs]]
[[Category:Caenagnathids]]

Purple finch

2017-12-15T20:59:52Z

Wikkler:

{{Taxobox
| name = Purple finch
| status = LC
| status_system = IUCN3.1
| status_ref = <ref>{{IUCN|id=22720553 |title=''Carpodacus purpureus'' |assessor=BirdLife International |assessor-link=BirdLife International |version=2013.2 |year=2012 |accessdate=9 April 2014}}</ref>
| image = Carpodacus purpureus CT3.jpg
| image_caption = Male
| image2 = Carpodacus purpureus CT4.jpg
| image2_caption = Female
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[bird|Aves]]
| ordo = [[passerine|Passeriformes]]
| familia = [[Finch|Fringillidae]]
| genus = ''[[Haemorhous]]''
| species = '''''H. purpureus'''''
| binomial = ''Haemorhous purpureus''
| binomial_authority = ([[Johann Friedrich Gmelin|Gmelin]], 1789)
| synonyms = ''Burrica purpurea'' 
''Carpodacus purpureus''
| range_map = Carpodacus purpureus map.svg
| range_map_caption = Range of ''C. purpureus'' {{leftlegend|#FFD0D0|Breeding range|outline=gray}}{{leftlegend|#3CE67B|Year-round range|outline=gray}}{{leftlegend|#F8A20C|Wintering range|outline=gray}}
}}

The '''purple finch''' (''Haemorhous purpureus'') is a bird in the finch [[family (biology)|family]], [[Fringillidae]].

==Taxonomy==
This species and the other "American rosefinches" were formerly included with the rosefinches of Eurasia in the genus ''[[Carpodacus]]''; however, the three North American species are not closely related to the rosefinches of the Old World, and have thus been moved to the genus ''Haemorhous'' by most taxonomic authorities.

It is included in the finch [[family (biology)|family]], [[Fringillidae]], which is made up of [[passerine]] [[bird]]s found in the northern hemisphere, Africa, and South America. The purple finch was originally described by [[Johann Friedrich Gmelin]] in 1789.<ref name="ITIS">{{ITIS|taxon=''Carpodacus purpureus''|id=179186| accessdate = 2008-07-18}}</ref>

There are two subspecies of the purple finch, ''H. p. purpureus'' and ''H. p. californicus''. ''H. p. californicus'' was identified by [[Spencer F. Baird]] in 1858.<ref name="ITIS" /> It differs from the nominate subspecies in that it has a longer tail and shorter wings. The [[plumage]] of both males and females is darker, and the coloration of the females is more greenish.<ref>{{cite book
| last = Bailey
| first = Florence Merriam
|author2=Fuertes, Louis Agassiz
| title = Handbook of Birds of the Western United States
| publisher = Houghton Mifflin
| year = 1921
| pages = 310
| url = https://books.google.com/books?id=nQ0LAAAAIAAJ&pg=PA310&dq=Carpodacus+purpureus}}</ref> The bill of ''C. p. californicus'' is also longer than that of the nominate subspecies.<ref>{{cite book
| last = Kaufman
| first = Kenneth
| title = A Field Guide to Advanced Birding
| publisher = HMCo Field Guides
| year = 1999
| pages = 267–268
| url = https://books.google.com/books?id=Jlb7Gyhi3IIC&pg=PA267&dq=Carpodacus+purpureus+%2B+red#PPA268,M1
| isbn = 0-395-97500-X}}</ref>

==Description==
Adults have a short forked brown tail and brown wings and are about {{convert|15|cm|in|abbr=on}} in length and weigh {{convert|34|g|oz|abbr=on}}.<ref>{{cite book
| last = Maehr
| first = David S.
| last2 = Kale, II
| first2 = Herbert W.
| title = Florida's Birds: A Field Guide and Reference
| publisher =Pineapple Press
| year = 2005
| location =
| pages = 211
| url = https://books.google.com/books?id=gIThorxtoU4C&printsec=frontcover&dq=Florida%27s+Birds:+A+Field+Guide+and+Reference#PPA211,M1
| isbn =1-56164-335-1}}</ref> Adult males are [[Raspberry (color)|raspberry]] red on the head, breast, back and rump; their back is streaked. Adult females have light brown upperparts and white underparts with dark brown streaks throughout; they have a white line on the face above the eye.

==Habitat and distribution==
Their breeding habitat is [[conifer]]ous and mixed forest in Canada and the northeastern United States, as well as various wooded areas along the U.S. Pacific coast. They nest on a horizontal branch or in a fork of a tree.{{Citation needed|date=January 2010}}

Birds from northern Canada [[bird migration|migrate]] to the southern United States; other birds are permanent residents.{{Citation needed|date=January 2010}}

The purple finch population has declined sharply in the East due to the [[house finch]]. Most of the time, when these two species collide, the house finch outcompetes the purple finch. This bird has been also displaced from some habitat by the introduced [[house sparrow]].<ref>{{cite journal|last=Wootton|first= J. T.| year = 1987|title=Interspecific Competition between Introduced House Finch Populations and Two Associated Passerine Species|journal=Oecologia |volume=71 |issue=3|pages= 325–331| doi=10.1007/BF00378703|pmid= 28312977}}</ref>

==Behavior==

===Diet===
These birds forage in trees and bushes, sometimes in ground vegetation. They mainly eat seeds, berries, and insects. They are fond of sunflower seeds, millet, and thistle.

==Cultural depictions==
This is the state bird of [[New Hampshire]].
In 1763, [[Richard Brookes]] made the description of the female purple finch in [[Mexico]] with the name of "chiantototl" ([[Salvia hispanica|chia]] seed bird).<ref>Brookes, Richard (1763). ''The Natural History of Birds.'' Vol 2, p 205.</ref>

==References==

{{reflist|2}}

==External links==
{{Commons|Haemorhous purpureus}}
{{Wikispecies|Haemorhous purpureus}}
*[http://www.birdhouses101.com/purple-finch.asp Interesting Purple Finch Facts at BirdHouses101.com]
*{{InternetBirdCollection|purple-finch-carpodacus-purpureus|Purple Finch}}
*{{VIREO|Purple+Finch|Purple Finch}}
*[http://www.birds.cornell.edu/AllAboutBirds/BirdGuide/Purple_Finch.html Purple Finch Species Account] – Cornell Lab of Ornithology

{{taxonbar}}

[[Category:Haemorhous]]
[[Category:Birds described in 1789]]
[[Category:Endemic birds of North America]]

Troodon

2017-12-09T19:39:14Z

Wikkler: /* History of discovery */

{{Italic title}}{{automatic taxobox
| name = ''Trododon''
| fossil_range = [[Late Cretaceous]], {{fossilrange|77.5|76.5|earliest=77.5|latest=76.5}}
| image = Troodon formosus.jpg
| image_width = 250px
| image_caption = Illustration of the ''T. formosus'' holotype tooth
| parent_authority = [[Charles W. Gilmore|Gilmore]], 1924
| authority = [[Joseph Leidy|Leidy]], [[1856 in paleontology|1856]]
| type_species = {{extinct}}'''''Troodon formosus'''''
| type_species_authority = Leidy, 1856
}}
'''''Troodon''''' ({{IPAc-en|ˈ|t|r|oʊ|.|ə|d|ɒ|n}} {{respell|TROH|ə-don}}; '''''Troödon''''' in older sources) is a dubious [[genus]] of relatively small, [[bird]]-like [[dinosaur]]s known definitively from the [[Campanian]] age of the [[Cretaceous]] [[Period (geology)|period]] (about 77 [[mya (unit)|mya]]). It includes at least one species, '''''Troodon formosus''''', known from [[Montana]]. Discovered in October 1855, ''T. formosus'' was among the first dinosaurs found in [[North America]], although it was thought to be a [[lizard]] until 1877.

The genus name is [[Ancient Greek|Greek]] for "wounding tooth", referring to the teeth, which were different from those of most other [[theropod]]s known at the time of their discovery. The teeth bear prominent, apically oriented serrations. These "wounding" serrations, however, are [[morphometric]]ally more similar to those of herbivorous reptiles, and suggest a possibly omnivorous diet.<ref name="Holtzetal1998">Holtz, Thomas R., Brinkman, Daniel L., Chandler, Chistine L. (1998) Denticle Morphometrics and a Possibly Omnivorous Feeding Habit for the Theropod Dinosaur Troodon. Gaia number 15. December 1998. pp. 159-166.</ref>

== History of discovery ==
The name was originally spelled ''Troödon'' (with a [[Diaeresis (diacritic)|diaeresis]]) by [[Joseph Leidy]] in 1856, which was officially amended to its current status by Sauvage in 1876.<ref>H.-E. Sauvage, 1876, "Notes sur les reptiles fossiles", ''Bulletin de la Société Géologique de France, 3e série'' '''4''': 435-444</ref> The [[type (zoology)|type specimen]] of ''Troodon'' has caused problems with classification, as the entire genus is based only on a single tooth from the [[Judith River Formation]]. ''Troodon'' has historically been a highly unstable classification and has been the subject of numerous conflicting synonymies with similar theropod specimens.<ref name=Talos/>

The ''Troodon'' tooth was originally classified as a "lacertilian" ([[lizard]]) by Leidy, but reassigned as a [[megalosaurid]] dinosaur by Nopcsa in 1901 (Megalosauridae having historically been a [[wastebin taxon]] for most carnivorous dinosaurs). In 1924, Gilmore suggested that the tooth belonged to the herbivorous [[pachycephalosaur]] ''[[Stegoceras]]'', and that ''Stegoceras'' was in fact a junior synonym of ''Troodon'' (the similarity of troodontid teeth to those of herbivorous dinosaurs continues to lead many paleontologists to believe that these animals were omnivores). The classification of ''Troodon'' as a pachycephalosaur was followed for many years, during which the family [[Pachycephalosauridae]] was known as [[Troodontidae]]. In 1945, [[Charles Mortram Sternberg]] rejected the possibility that ''Troodon'' was a pachycephalosaur due to its stronger similarity to the teeth of other carnivorous dinosaurs. With ''Troodon'' now classified as a theropod, the family Troodontidae could no longer be used for the dome-headed dinosaurs, so Sternberg named a new family for them, Pachycephalosauridae.<ref name=sternberg1945>{{Cite journal | last1 = Sternberg | first1 = C. | year = 1945 | title = Pachycephalosauridae proposed for domeheaded dinosaurs, ''Stegoceras lambei'' n. sp., described | url = | journal = Journal of Paleontology | volume = 19 | issue = | pages = 534–538 }}</ref>

The first specimens assigned to ''Troodon'' that were not teeth were both found by Sternberg in the early 1930s, in the [[Dinosaur Park Formation]] of [[Alberta]]. The first was named ''[[Stenonychosaurus inequalis]]'' by Sternberg in 1932, based on a foot, fragments of a hand, and some tail vertebrae. A remarkable feature of these remains was the enlarged claw on the second toe, which is now recognized as characteristic of early [[Paraves|paravians]]. Sternberg initially classified ''Stenonychosaurus'' as a member of the family [[Coeluridae]]. The second, a partial lower jaw bone, was described by Gilmore (1932) as a new species of lizard which he named ''[[Polyodontosaurus|Polyodontosaurus grandis]]''. Later, in 1951, Sternberg recognized ''P. grandis'' as a possible synonym of ''Troodon'', and speculated that since ''Stenonychosaurus'' had a "very peculiar [[pes (zoology)|pes]]" and ''Troodon'' "equally unusual teeth", they may be closely related. Unfortunately, no comparable specimens were available at that time to test the idea. In a recent revision of the material by van der Reest & Currie, ''Polyodontosaurus'' was determined to be a nomen dubium, not fit fot synonymy with other taxa.<ref name=LS2017/>

A more complete skeleton of ''Stenonychosaurus'' was described by [[Dale Russell]] in 1969 from the Dinosaur Park Formation, which eventually formed the scientific foundation for a famous life-sized sculpture of ''Stenonychosaurus'' accompanied by its fictional, [[humanoid]] descendant, the "dinosauroid".<ref name=russell1982>{{Cite journal | last1 = Russell | first1 = D. A. | last2 = Séguin | first2 = R. | year = 1982 | title = Reconstruction of the small Cretaceous theropod ''Stenonychosaurus inequalis'' and a hypothetical dinosauroid | url = https://archive.org/details/syllogeus37nati | journal = Syllogeus | volume = 37 | issue = | pages = 1–43 }}</ref> ''Stenonychosaurus'' became a well-known theropod in the 1980s, when the feet and braincase were described in more detail. Along with ''[[Saurornithoides]]'', it formed the family [[Saurornithoididae]]. Based on differences in tooth structure, and the extremely fragmentary nature of the original ''Troodon formosus'' specimens, saurornithoidids were thought to be close relatives while ''Troodon'' was considered a dubious possible relative of the family. [[Phil Currie]], reviewing the pertinent specimens in 1987, showed that supposed differences in tooth and jaw structure among troodontids and saurornithoidids were based on age and position of the tooth in the jaw, rather than a difference in species. He reclassified ''Stenonychosaurus inequalis'' as well as ''Polyodontosaurus grandis'' and ''[[Pectinodon bakkeri]]'' as junior synonyms of ''Troodon formosus''. Currie also made Saurornithoididae a junior synonym of Troodontidae.<ref name=currie1987>{{Cite journal | last1 = Currie | first1 = P. | year = 1987 | title = Theropods of the Judith River Formation | url = | journal = Occasional Paper of the Tyrrell Museum of Palaeontology | volume = 3 | issue = | pages = 52–60 }}</ref> In 1988, [[Gregory S. Paul]] went farther and included ''Saurornithoides mongoliensis'' in the genus ''Troodon'' as ''T. mongoliensis'',<ref name=paul1988b>{{cite book|last=Paul |first=G.S. |year=1988 |title=Predatory Dinosaurs of the World |location=New York |publisher=Simon and Schuster |pages=398–399|isbn=0-671-61946-2 }}</ref> but this reclassification, along with many other unilateral synonymizations of well known genera, was not adopted by other researchers. Currie's classification of all North American troodontid material in the single species ''Troodon formosus'' became widely adopted by other [[paleontology|paleontologists]], and all of the specimens once called ''Stenonychosaurus'' were referred to as ''Troodon'' in the scientific literature through the early 21st century.
[[File:The Childrens Museum of Indianapolis - Troodon teeth.jpg|thumb|Teeth from South Dakota assigned to ''T. formosus'', with a [[US dime]] coin for scale, [[Children's Museum of Indianapolis]]]]
However, the concept that all Late Cretaceous North American troodontids belong to one species began to be questioned soon after Currie's 1987 paper was published, including by Currie himself. Currie and colleagues (1990) noted that, while they believed the Judith River troodontids were all ''T. formosus'', troodontid fossils from other formations, such as the [[Hell Creek Formation]] and [[Lance Formation]], might belong to different species. In 1991, George Olshevsky assigned the Lance formation fossils, which had first been named ''Pectinodon bakkeri'' but later synonymized with ''Troodon formosus'' to the species ''Troodon bakkeri'', and several other researchers (including Currie) have reverted to keeping the Dinosaur Park Formation fossils separate as ''Troodon inequalis'' (now ''Stenonychosaurus inequalis'').<ref name=currie2005>Currie, P. (2005). "Theropods, including birds." in Currie and Koppelhus (eds). ''Dinosaur Provincial Park, a spectacular ecosystem revealed, Part Two, Flora and Fauna from the park.'' Indiana University Press, Bloomington. Pp 367–397.</ref>

In 2011, Zanno and colleagues reviewed the convoluted history of troodontid classification in Late Cretaceous North America. They followed Longrich (2008) in treating ''Pectinodon bakkeri'' as a valid genus, and noted that it is likely the numerous Late Cretaceous specimens currently assigned to ''Troodon formosus'' almost certainly represent numerous new species, but that a more thorough review of the specimens is required. Because the holotype of ''T. formosus'' is a single tooth, this renders ''Troodon'' a [[nomen dubium]].<ref name=Talos/>

In 2017, Evans and colleagues further discussed the undiagnostic nature of the holotype of ''Troodon formosus'' and suggested that ''Stenonychosaurus'' be used for troodontid skeletal material from the Dinosaur Park Formation. <ref name=Alberavenator>{{cite journal|last1=Evans|first1=D. C.|last2=Cullen|first2=T.M.|last3=Larson|first3=D.W.|last4=Rego|first4=A.|title=A new species of troodontid theropod (Dinosauria: Maniraptora) from the Horseshoe Canyon Formation (Maastrichtian) of Alberta, Canada|journal=Canadian Journal of Earth Sciences|date=2017|pages=813–826|doi=10.1139/cjes-2017-0034|url=http://www.nrcresearchpress.com/doi/10.1139/cjes-2017-0034#.Wd5p19PytE4}}</ref> Later in the same year, Aaron J. van der Reest and Currie came to a similar conclusion as Evans and colleagues, and also split much of the material assigned to ''Stenonychosaurus'' into a new genus: ''[[Latenivenatrix]]''. <ref name=LS2017>{{cite journal|last1=van der Reest|first1=A. J.|last2=Currie|first2=P. J.|title=Troodontids (Theropoda) from the Dinosaur Park Formation, Alberta, with a description of a unique new taxon: implications for deinonychosaur diversity in North America|journal=Canadian Journal of Earth Sciences|date=2017|pages=919–935|doi=10.1139/cjes-2017-0031|url=http://www.nrcresearchpress.com/doi/abs/10.1139/cjes-2017-0031#.WYnhUlGGOUm}}</ref>

==Classification==
''Troodon'' is considered to be one of the most derived members of its family. Along with ''[[Zanabazar (dinosaur)|Zanabazar]]'', ''[[Saurornithoides]]'' and ''[[Talos (dinosaur)|Talos]]'' it forms a clade of specialized troodontids.<ref name=Talos/>

Below is a cladogram of Troodontidae by Zanno ''et al.'' in 2011.<ref name=Talos/>

{{clade| style=font-size:100%; line-height:100%
|label1=[[Troodontidae]]
|1={{clade
|1=''[[Sinovenator changii]]'' | state1=dashed
|2={{clade
|1={{clade
|1=''Sinovenator changii'' | state1=dashed
|2=''[[Mei long]]'' }}
|2={{clade
|1=IGM 100/44
|2={{clade
|1=''[[Sinornithoides youngi]]''
|2={{clade
|1=''[[Talos sampsoni]]'' | state1=dashed
|2={{clade
|1=''[[Byronosaurus jaffei]]''
|2={{clade
|1=''Talos sampsoni'' | state1=dashed
|2={{clade
|1={{clade
|1=''Talos sampsoni'' | state1=dashed
|2=''[[Saurornithoides mongoliensis]]'' }}
|2={{clade
|1=''[[Zanabazar junior]]''
|2='''''Troodon formosus''''' }} }} }} }} }} }} }} }} }} }}

==Paleobiology==
One study was based on multiple ''Troodon'' teeth that have been collected from Late Cretaceous deposits from northern Alaska. These teeth are much larger than those collected from more southern sites, providing evidence that northern Alaskan populations of ''Troodon'' grew to larger average body size. This study also provides an analysis of the proportions and wear patterns of a large sample of ''Troodon'' teeth. It proposes that the wear patterns of all ''Troodon'' teeth suggest a diet of soft foods - inconsistent with bone chewing, invertebrate exoskeletons, or tough plant items. This study hypothesizes a diet primarily consisting of meat.<ref name="Fiorillo2008">Fiorillo, Anthony R. (2008) "On the Occurrence of Exceptionally Large Teeth of ''Troodon'' (Dinosauria: Saurischia) from the Late Cretaceous of Northern Alaska" Palaios volume 23 pp.322-328</ref>

==Paleoecology==
[[File:Troodon Perot Museum.jpg|thumb|upright|Restored skeleton of an unnamed Alaskan species, [[Perot Museum]]]]
The type specimen of ''Troodon formosus'' was found in the [[Judith River Formation]] of [[Montana]]. The rocks of the Judith River Formation are equivalent in age with the [[Oldman Formation]] of [[Alberta]],<ref name=eberth1997>{{cite book |last=Eberth |first=David A. |year=1997 |chapter=Judith River Wedge |editor=[[Phil Currie|Currie, Philip J.]] |editor2=Padian, Kevin |title=Encyclopedia of Dinosaurs |publisher= Academic Press |location=San Diego |pages=199–204 |isbn=0-12-226810-5}}</ref> which has been dated to between 77.5 and 76.5 million years ago.<ref name=ABS09>{{cite journal |last=Arbour |first=V. M. |author2=Burns, M. E. |author3= Sissons, R. L. |year=2009 |title=A redescription of the ankylosaurid dinosaur ''Dyoplosaurus acutosquameus'' Parks, 1924 (Ornithischia: Ankylosauria) and a revision of the genus |journal=Journal of Vertebrate Paleontology |volume=29 |issue=4 |pages=1117–1135 |doi=10.1671/039.029.0405}}</ref>

In the past, remains have been attributed to the same genus as the Judith River ''Troodon'' from a wide variety of other geological formations. It is now recognized as unlikely that all of these fossils, which come from localities hundreds or thousands of miles apart, separated by millions of years of time, represent a single species or genus of troodontid dinosaurs. Further study and more fossils are needed to determine how many species of ''Troodon'' existed. It is questionable that, after further study, any additional species can be referred to ''Troodon'', in which case the genus would be considered a [[nomen dubium]].<ref name=Talos>{{Cite journal|author=Lindsay E. Zanno, David J. Varricchio, Patrick M. O'Connor, Alan L. Titus and Michael J. Knell |year=2011 |title=A new troodontid theropod, ''Talos sampsoni'' gen. et sp. nov., from the Upper Cretaceous Western Interior Basin of North America |url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0024487 |journal=PLoS ONE |volume=6 |issue=9 |pages=e24487 |doi=10.1371/journal.pone.0024487 |pmid=21949721 |pmc=3176273|editor1-last=Lalueza-Fox|editor1-first=Carles}}</ref>

Additional specimens currently referred to ''Troodon'' come from the upper [[Two Medicine Formation]] of [[Montana]] and the [[Prince Creek Formation]] of [[Alaska]]. There is some evidence that ''Troodon'' favored cooler climates, as its species seem to have been particularly abundant in northern and even [[arctic]] areas and during cooler intervals, such as the early [[Maastrichtian]].<ref name=alaska>{{Cite journal | doi = 10.1671/0272-4634(2000)020[0675:TTFTPC]2.0.CO;2 | last1 = Fiorillo | first1 = Anthony R.| last2 = Gangloff | first2 = Roland A.| year = 2000 | title = Theropod teeth from the Prince Creek Formation (Cretaceous) of Northern Alaska, with speculations on Arctic dinosaur paleoecology | url = | journal = Journal of Vertebrate Paleontology | volume = 20 | issue = 4| pages = 675–682 }}</ref> ''Troodon''-like teeth have been found in the lower [[Javelina Formation]] of [[Texas]] and the Naashoibito Member of the [[Ojo Alamo Formation]] in [[New Mexico]].<ref name=texas>Langston, Standhardt and Stevens, (1989). "Fossil vertebrate collecting in the Big Bend - History and retrospective." in ''Vertebrate Paleontology, Biostratigraphy and Depositional Environments, Latest Cretaceous and Tertiary, Big Bend Area, Texas''. Guidebook Field Trip Numbers 1 a, B, and 49th Annual Meeting of the Society of Vertebrate Paleontology, Austin, Texas, 29 October - 1 November 1989. 11-21.</ref><ref name=weil&williamson2000>Weil and Williamson, (2000). "Diverse Maastrichtian terrestrial vertebrate fauna of the Naashoibito Member, Kirtland Formation (San Juan Basin, New Mexico) confirms "Lancian" faunal heterogeneity in western North America." ''Geological Society of America Abstracts with Programs'', '''32''': A-498.</ref>

==See also==
{{Portal|Dinosaurs}}
* [[Timeline of troodontid research]]

==References==
{{Reflist|2}}
* Russell, D. A. (1987). "Models and paintings of North American dinosaurs." In: Czerkas, S. J. & Olson, E. C. (eds) ''Dinosaurs Past and Present, Volume I.'' Natural History Museum of Los Angeles County/University of Washington Press (Seattle and Washington), pp. 114–131.

== External links ==
{{commons category|Troodon}}
* [http://www.daviddarling.info/encyclopedia/D/dinosaurintell.html Dinosauroid]

{{Troodontidae}}

[[Category:Late Cretaceous dinosaurs of North America]]
[[Category:Troodontids]]
[[Category:Campanian life]]
[[Category:Fossil taxa described in 1856]]
[[Category:Taxa named by Joseph Leidy]]
[[Category:Paleontology in Montana]]
[[Category:Campanian genus first appearances]]
[[Category:Campanian genus extinctions]]

Caenagnathidae

2017-12-09T17:09:53Z

Wikkler: Both have been found to possibly not be caenagnathids. I'm putting question marks here.

{{automatic Taxobox
| name = Caenagnathids
| fossil_range = [[Late Cretaceous]], {{fossil range|91|65|}}
| image = Chirostenotes skull.jpg
| image_caption = Reconstructed skull of ''[[Anzu wyliei]]''
| authority = [[Charles Hazelius Sternberg|Sternberg]], 1940
| type_species = ''[[Caenagnathus collinsi]]''
| type_species_authority = Sternberg, 1940
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
*{{extinct}}''[[Beibeilong]]''
*{{extinct}}''[[Epichirostenotes]]''
*{{extinct}}''[[Gigantoraptor]]''?
*{{extinct}}''[[Hagryphus]]''
*{{extinct}}''[[Microvenator]]''?
*{{extinct}}''[[Ojoraptorsaurus]]''
*{{extinct}}'''Caenagnathinae'''
**{{extinct}}''[[Anzu wyliei|Anzu]]''
**{{extinct}}''[[Caenagnathus]]''
*{{extinct}}'''Elmisaurinae'''
**{{extinct}}''[[Apatoraptor]]''?
**{{extinct}}''[[Caenagnathasia]]''
**{{extinct}}''[[Chirostenotes]]''
**{{extinct}}''[[Elmisaurus]]''
**{{extinct}}''[[Leptorhynchos (dinosaur)|Leptorhynchos]]''
| synonyms =
* '''Elmisauridae''' [[Halszka Osmólska|Osmólska]], 1981
}}
'''Caenagnathidae''' is a family of bird-like [[maniraptora]]n [[theropod]] [[dinosaur]]s from the Late Cretaceous of North America and Asia. They are a member of the [[Oviraptorosauria]], and close relatives of the [[Oviraptoridae]].<ref>Osmólska, H., P. J. Currie, et al. (2004). Oviraptorosauria. The Dinosauria. D. B. Weishampel, P. Dodson and H. Osmolska. Berkeley, University of California Press: 165-183.</ref> Like other oviraptorosaurs, caenagnathids had specialized beaks,<ref name="Currie, P. J. 1993">{{cite journal | last1 = Currie | first1 = P. J. | last2 = Godfrey | first2 = S. J. | display-authors = 2 | last3 = et al | year = 1993 | title = New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia | url = | journal = Canadian Journal of Earth Sciences | volume = 30 | issue = 10-11| pages = 2255–2272 | doi=10.1139/e93-196}}</ref> long necks,<ref name="Sues, H. D. 1997">{{cite journal | last1 = Sues | first1 = H. D. | year = 1997 | title = On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from western North America | url = | journal = Journal of Vertebrate Paleontology | volume = 17 | issue = 4| pages = 698–716 | doi=10.1080/02724634.1997.10011018}}</ref> and short tails,<ref name = "barsboldetal2000">{{cite journal | last1 = Barsbold | first1 = R. | last2 = Osmolska | first2 = H. | last3 = Watabe | first3 = M. | last4 = Currie | first4 = P. J. | last5 = Tsogtbaatar | first5 = K. | year = 2000 | title = New oviraptorosaur (Dinosauria, Theropoda) from Mongolia: The first dinosaur with a pygostyle | url = http://www.app.pan.pl/archive/published/app45/app45-097.pdf | format=PDF| journal = Acta Palaeontologica Polonica | volume = 45 | issue = 2| pages = 97–106 }}</ref> and would have been covered in feathers. The relationships of caenagnathids were long a puzzle. The family was originally named by [[Charles Hazelius Sternberg]] in 1940 <ref name = "sternberg1940">{{cite journal | last1 = Sternberg | first1 = R.M. | year = 1940 | title = A toothless bird from the Cretaceous of Alberta | url = | journal = Journal of Paleontology | volume = 14 | issue = 1| pages = 81–85 }}</ref> as a family of flightless birds. The discovery of skeletons of the related oviraptorids revealed that they were in fact non-avian theropods,<ref>{{cite journal | last1 = Osmólska | first1 = H | year = 1976 | title = New light on the skull anatomy and systematic position of Oviraptor | url = | journal = Nature | volume = 262 | issue = | pages = 683–684 | doi=10.1038/262683a0}}</ref> and the discovery of more complete caenagnathid remains <ref name="Sues, H. D. 1997"/><ref name = "currierussell1988">{{cite journal | last1 = Currie | first1 = P.J. | last2 = Russell | first2 = D.A. | year = 1988 | title = Osteology and relationships of ''Chirostenotes pergracilis'' (Saurischia, Theropoda) from the Judith River Oldman Formation of Alberta | doi = 10.1139/e88-097 | journal = Canadian Journal of Earth Sciences | volume = 25 | issue = 3| pages = 972–986 }}</ref> revealed that ''Chirostenotes pergracilis'', originally named on the basis of a pair of hands, and ''"Ornithomimus" elegans'', named from a foot, were caenagnathids as well.

==Anatomy==
[[File:Oviraptorid Clean.png|thumb|left|''Anzu wyliei'' skeleton cast in the [[Rocky Mountain Dinosaur Resource Center]] in Woodland Park, Colorado, USA.]]
Overall, the anatomy of the caenagnathids is similar to that of the closely related Oviraptoridae, but there are a number of differences. In particular, caenagnathid jaws exhibited a distinct suite of specializations not seen in other oviraptorosaurs. Compared to the oviraptorids, the jaws tended to be relatively long and shallow, suggesting that the bite was not as powerful. The inside of the lower jaws also bore a complex series of ridges and toothlike processes, as well as a pair of horizontal, shelf-like structures. Furthermore, the jaws were unusual in being hollow and air filled, apparently being connected to the air sac system.<ref name="Currie, P. J. 1993"/> Caenagnathids also tended to be more lightly built than the oviraptorids. They had slender arms and long, gracile legs,<ref name="currierussell1988"/> although they lacked the extreme cursorial specializations seen in avimimids and ''Caudipteryx''.

==Etymology==
The name ''Caenagnathus'' (and hence Caenagnathidae) means "recent jaws"—when first discovered, it was thought that caenagnathids were close relatives of [[paleognathae|paleognath]] birds (such as the [[ostrich]]) based on features of the lower jaw. Since it would be unusual to find a recent group of birds in the Cretaceous, the name "recent jaws" was applied. Most paleontologists, however, now think that the birdlike features of the jaw were acquired [[convergent evolution|convergently]] with modern birds.<ref name = "cracraft1971">{{cite journal | last1 = Cracraft | first1 = J. | year = 1971 | title = Caenagnathiformes: Cretaceous birds convergent in jaw mechanism to dicynodont reptiles | url = | journal = Journal of Paleontology | volume = 45 | issue = | pages = 805–809 }}</ref><ref name = "barsboldetal1990">[[Rinchen Barsbold|Barsbold]], R., Maryańska, T., and Osmólska, H. (1990). "Oviraptorosauria." pg. 249-258 ''in'' Weishampel, Dodson, and Osmolska (eds.) ''The Dinosauria'', University of California Press (Berkeley).</ref>

==Evolution==
The earliest known caenagnathid is ''Caenagnathasia martinsoni'', from the Turonian Bissekty Formation of Uzbekistan.<ref name="Currie, P. J. 1993"/> The jaw of ''Caenagnathasia'' already has the specialized ridges and crushing surfaces seen in later forms. This suggests that caenagnathids originated well before the Turonian, but currently, there are no Early Cretaceous caenagnathid fossils. Caenagnathids later appeared in western North America, during the Campanian, suggesting that they may have originated in Asia, then migrated into western North America. Caenagnathids showed considerable variation in form. The tiny jaws of ''Caenagnathasia'' suggest a small animal, perhaps the size of a turkey. An unnamed species from the Hell Creek Formation <ref name="Currie, P. J. 1993"/> indicates a much larger animal, considerably larger than a human. If ''Gigantoraptor erlianensis'' is a caenagnathid, then it would represent far and away the largest member of the group, measuring up to 8 meters in length and weighing up to 1.4 tons. Their beaks also show considerable variation; that of ''Caenagnathasia'' is relatively short and deep, while that of ''Caenagnathus'' is long and shovel-shaped. This variation in size and beak shape suggests that caenagnathids evolved to exploit a range of ecological niches. Caenagnathids persisted up until the end of the Cretaceous period, as shown by the presence of at least two species in the late Maastrichtian Hell Creek Formation, before vanishing at the end of the Cretaceous along with all other non-avian dinosaurs.

==Classification==
The family Caenagnathidae, together with its sister group the [[Oviraptoridae]], comprises the superfamily [[Caenagnathoidea]]. In [[phylogenetic taxonomy]], the [[clade]] Caenagnathidae is defined as the most inclusive group containing ''[[Chirostenotes|Chirostenotes pergracilis]]'' but not ''[[Oviraptor|Oviraptor philoceratops]]''. While before 2010s only about two to six species were commonly recognized as belonging to the Caenagnathidae, currently that number may be much greater, with new discoveries and theories about older species that may inflate this number to up to ten. Much of this historical difference centers on the first caenagnathid to be described, ''Chirostenotes pergracilis''. Due to the poor preservation of most caenagnathid remains and resulting misidentifications, different bones and different specimens of ''Chirostenotes'' have historically been assigned to a number of different species. For example, the feet of one species, named ''Macrophalangia canadensis'',<ref name = "sternberg1932">{{cite journal | last1 = Sternberg | first1 = C. H. | year = 1932 | title = Two new theropod dinosaurs from the Belly River Formation of Alberta | url = | journal = The Canadian Field-Naturalist | volume = 46 | issue = | pages = 99–105 }}</ref> were known from the same region from which ''Chirostenotes pergracilis'' was recovered, but the discovery of a new specimen with both hands and feet preserved<ref name="currierussell1988"/> provided the support to combine them, while the later discovery of a partial skull with hands and feet <ref name="Sues, H. D. 1997"/> suggested that ''Chirostenotes'' and ''Caenagnathus'' were the same animal, and current studies of caenagnathid relationships continue to find them as closely related genera.<ref name=leptorhynchos>{{Cite journal | last1 = Longrich | first1 = N. R. | last2 = Barnes | first2 = K. | last3 = Clark | first3 = S. | last4 = Millar | first4 = L. | title = Caenagnathidae from the Upper Campanian Aguja Formation of West Texas, and a Revision of the Caenagnathinae | doi = 10.3374/014.054.0102 | journal = Bulletin of the Peabody Museum of Natural History | volume = 54 | pages = 23–49 | year = 2013 | pmid = | pmc = }}</ref>
[[File:Caenagnathidae.jpg|thumb|Caenagnathid skeletons to scale]]
Longrich and colleagues (2013) defined a subgroup of Caenagnathidae, the '''Caenagnathinae''', as all caenagnathids more closely related to ''Caenagnathus collinsi'' than to ''Caenagnathasia martinsoni'' or ''Elmisaurus rarus''.<ref name=leptorhynchos/> In 2015, the group '''Elmisaurinae''' was defined, including all species more closely related to ''Elmisaurus rarus'' than to ''Caenagnathus collinsi''.<ref>Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.</ref><ref name="currie15">{{cite journal|last=Currie|first=P.J.|authorlink=Philip J. Currie|last2=Funston|first2=G.F.|last3=Osmólska|first3=H.†|authorlink3=Halszka Osmólska|year=2015|title=New specimens of the crested theropod dinosaur ''Elmisaurus rarus'' from Mongolia|url=http://app.pan.pl/archive/published/app60/app001302014_acc.pdf|journal=Acta Palaeontologica Polonica|volume=XX|issue=XX|pages=XXX–XXX|doi=10.4202/app.00130.2014}}</ref>

The [[cladogram]] below follows an analysis by Funston & Currie in 2016.<ref name=apatoraptor1>{{cite journal |author1=Gregory F. Funston |author2=Philip J. Currie |year=2016 |title=A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae |journal=Journal of Vertebrate Paleontology |volume=Online edition |issue= |pages=e1160910 |doi=10.1080/02724634.2016.1160910 }}</ref>

{{clade| style=font-size:100%;line-height:100%
|label1=Caenagnathidae
|1={{clade
|1=''[[Microvenator celer]]''
|label2=unnamed
|2={{clade
|1=''[[Gigantoraptor erlianensis]]''
|label2=unnamed
|2={{clade
|1=''[[Hagryphus giganteus]]''
|2={{clade
|1={{clade
|1=''[[Epichirostenotes curriei]]''
|2=''[[Anzu wyliei]]'' }}
|2={{clade
|1=''[[Caenagnathus collinsi]]''
|label2='''Elmisaurinae'''
|2={{clade
|1=''[[Caenagnathasia martinsoni]]''
|2={{clade
|1=''[[Chirostenotes pergracilis]]''
|2={{clade
|1=''[[Leptorhynchos elegans]]''
|2={{clade
|1=''[[Apatoraptor pennatus]]''
|2=''[[Elmisaurus rarus]]''
}} }} }} }} }} }} }} }} }} }}

===Species===
Roughly a dozen caenagnathid species have been named, but it remains unclear how many are valid. Many species are known from fragmentary remains, such as jaws, hands, or feet, making comparisons between them difficult. ''Caenagnathus sternbergi'', for example, was described on the basis of a jaw bone. It has been interpreted as either the jaws of ''Chirostenotes pergracilis'' (described on the basis of a pair of hands) or ''Chirostenotes elegans''<ref name="Sues, H. D. 1997"/> (described on the basis of a foot), but because no complete skeleton is known, it is difficult to be certain which animal it belongs to. The relationships of other species remain in doubt. ''Gigantoraptor'' was originally interpreted as an oviraptorid, but may in fact represent a primitive caenagnathid.<ref name="Longrichetal10">{{cite journal |year=2010 |title=A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia |journal=Palaeontology |volume=53 |issue=5 |pages=945–960 |doi=10.1111/j.1475-4983.2010.00968.x |url=http://onlinelibrary.wiley.com/doi/10.1111/j.1475-4983.2010.00968.x/abstract|author1=Nicholas R. Longrich |author2=Philip J. Currie |author3=Dong Zhi-Ming }}</ref>

* ''[[Anzu wyliei]]'' - ([[Hell Creek Formation]], Montana, United States)<ref name="anzu">{{Cite journal | doi = 10.1371/journal.pone.0092022| title = A New Large-Bodied Oviraptorosaurian Theropod Dinosaur from the Latest Cretaceous of Western North America| journal = PLoS ONE| volume = 9| issue = 3| pages = e92022| year = 2014| last1 = Lamanna | first1 = M. C. | last2 = Sues | first2 = H. D. | last3 = Schachner | first3 = E. R. | last4 = Lyson | first4 = T. R. | pmid=24647078 | pmc=3960162}}</ref><ref name=triebold>Varricchio, D. J. (2001). Late Cretaceous Oviraptorosaur (Theropoda) dinosaurs from Montana. Mesozoic Vertebrate Life. D. H. Tanke and K. Carpenter. Bloomington, Indiana University Press: 42-57.</ref>
* [[Apatoraptor|''Apatoraptor pennatus'']] - ([[Horseshoe Canyon Formation]], Alberta)
* ''[[Caenagnathasia martinsoni]]'' - ([[Bissekty Formation]], Mongolia)
* ''[[Chirostenotes pergracilis]]'' - ([[Dinosaur Park Formation]], Alberta, Canada)
* ''[[Caenagnathus collinsi]]'' - ([[Dinosaur Park Formation]], Alberta, Canada)
* ''[[Elmisaurus rarus]]'' - ([[Nemegt Formation]], Mongolia)
* ''[[Epichirostenotes curriei]]'' - ([[Horseshoe Canyon Formation]], Alberta, Canada)
* ''[[Gigantoraptor erlianensis]]'' - ([[Iren Dabasu Formation]], Inner Mongolia, China)
* ''[[Hagryphus giganteus]]'' - ([[Kaiparowits Formation]], Utah, United States)
* ''[[Leptorhynchos elegans]]'' - ([[Dinosaur Park Formation]], Alberta, Canada)
* ''[[Leptorhynchos gaddisi]]'' - ([[Aguja Formation]], Texas, United States)<ref name=leptorhynchos/>
* ''[[Ojoraptorsaurus boerei]]'' - ([[Ojo Alamo Formation]], New Mexico, United States)

Caenagnathids are only known from the Late Cretaceous of North America and Asia. The earliest and most primitive known caenagnathid is ''[[Caenagnathasia|Caenagnathasia martinsoni]]'', from the Bissekty Formation of Uzbekistan.<ref name = "CurrieGodfreyNesov1994">{{cite journal | last1 = Currie | first1 = P.J. | last2 = Godfrey | first2 = S.J. | last3 = Nesov | first3 = L.A. | year = 1994 | title = New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia | url = | journal = Canadian Journal of Earth Sciences | volume = 30 | issue = | pages = 2255–2272 | doi=10.1139/e93-196}}</ref> The largest is the enormous ''Gigantoraptor erlianensis''.<ref name="Longrichetal10"/>

==See also==
{{Portal|Dinosaurs}}
* [[Timeline of oviraptorosaur research]]

==References==
{{Reflist}}

==External links==
* [http://qilong.8m.com/Caenagnathidae.html Overview of Caenagnathidae by Jaime Headden.]
* [https://web.archive.org/web/20060827031308/http://www.carnegiemnh.org/ditw/ovi/res_mount.htm Photo of the Triebold caenagnathid, on display at the Carnegie Museum].
* [https://web.archive.org/web/20070312081952/http://www.skeletaldrawing.com/psgallery/pages/chirostenotes.html Skeletal reconstruction of the Triebold specimens.]

{{Oviraptorosauria}}

[[Category:Oviraptorosaurs]]
[[Category:Caenagnathids]]

Coelurosauria

2017-12-09T03:01:18Z

Wikkler: Not possible for Coelurus to not be a coelurosaur.

{{Automatic taxobox |
| name = Coelurosaurians
| fossil_range = [[Late Jurassic]]–[[Holocene|Present]], {{Fossil range|165|0|earliest=196}}Possible [[Early Jurassic]] record
| image = Zuni Coelurosaur.jpg
| image_width = 250px
| image_caption = Reconstructed coelurosaur skeleton, [[Wyoming Dinosaur Center]]
| authority = [[Friedrich von Huene|von Huene]], 1914
| 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 =
*{{extinct}}''[[Aniksosaurus]]''<ref name="Holtz2008"/>
*{{extinct}}''[[Bagaraatan]]''<ref name="Holtz2008"/>
*{{extinct}}''[[Bicentenaria]]''
*{{extinct}}''[[Coelurus]]''
*{{extinct}}''[[Lourinhanosaurus]]''?<ref name=carranno2012>{{cite journal|last1=Carrano|first1=M.T.|last2=Benson|first2=R.B.J.|last3=Sampson|first3=S.D.|title=The phylogeny of Tetanurae (Dinosauria: Theropoda)|journal=Journal of Systematic Palaeontology|date=2012|volume=10|issue=2|pages=211–300|doi=10.1080/14772019.2011.630927}}</ref>
*{{extinct}}''[[Nedcolbertia]]''<ref name="Holtz2008"/>
*{{extinct}}''[[Phaedrolosaurus]]''<ref name="Holtz2008"/>
*{{extinct}}''[[Richardoestesia]]''<ref name="Holtz2008"/>
*{{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>
*{{extinct}}''[[Tanycolagreus]]''?
*{{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>
*{{extinct}}''[[Tugulusaurus]]''
*{{extinct}}''[[Xinjiangovenator]]''<ref name="Holtz2008"/>
*{{extinct}}''[[Zuolong]]''
*'''Tyrannoraptora''' [[Paul Sereno|Sereno]], 1999
**{{extinct}}''[[Aorun]]''
**{{extinct}}''[[Ornitholestes]]''
**{{extinct}}[[Compsognathidae]]
**{{extinct}}[[Tyrannosauroidea]]
**[[Maniraptoriformes]]
}}
'''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>

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==

===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]]).<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 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|Feathered dinosaurs}}
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>

===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 [[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==
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 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|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 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 [[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 />

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 (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>
{{clade| style=font-size:100%;line-height:80%
|label1='''Coelurosauria'''
|1={{clade
|1=†''[[Bicentenaria]]''
|2={{clade
|1=†''[[Zuolong]]''
|label2='''Tyrannoraptora'''
|2={{clade
|1=†[[Tyrannosauroidea]][[File:Rjpalmer tyrannosaurusrex (white background).jpg|130 px]]
|label2={{color|white|unnamed}}
|2={{clade
|1=†''[[Aorun]]''
|2={{clade
|1=†''[[Scipionyx]]''
|2={{clade
|1=†''[[Ornitholestes]]
|2={{clade
|1=†[[Compsognathidae]][[File:Sinosauropteryx mmartyniuk solosml (flipped).png|80px]]
|label2=[[Maniraptoriformes]]
|2={{clade
|1=†[[Ornithomimosauria]][[File:Hypothetical Deinocheirus (flipped).jpg|120 px]]
|2=[[Maniraptora]][[File:Deinonychus ewilloughby (flipped).png|80px]]
}} }} }} }} }} }} }} }} }}

==="Coelurosaurus"===
"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==
{{Portal|Dinosaurs}}
*[[Feathered dinosaurs]]
*[[Origin of birds]]
*[[List of fossil birds]]

==References==
{{Reflist|2|refs=
<ref name="currie-2005">Currie (2005) p. 368.</ref>
<ref name="csaharicus-endo-abs-19">"Abstract," Larsson (2001). Page 19.</ref>
<ref name=KP04>Padian (2004). Basal Avialae. Pages 210–231.</ref>}}

==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}}
* 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.
* {{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}}.
* 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==
{{Wiktionary|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]
* [https://web.archive.org/web/20101011112158/http://www.palaeos.com/Vertebrates/Units/340Theropoda/340.500.html Palaeos: Coelurosauria]
* [http://tolweb.org/Coelurosauria/15769 Tree of Life Web: Coelurosauria]

[[Category:Coelurosaurs| ]]
[[Category:Late Jurassic dinosaurs]]
[[Category:Cretaceous dinosaurs]]
[[Category:Cenozoic dinosaurs]]
[[Category:Extant Late Jurassic first appearances]]
[[Category:Fossil taxa described in 1914]]
[[Category:Taxa named by Friedrich von Huene]]

Tetanurae

2017-12-09T02:06:34Z

Wikkler: All three have been recovered as non-tetanurans in various analyses.

{{Automatic taxobox
| name = Tetanurans
| fossil_range = [[Early Jurassic]]–[[Holocene|Present]], {{Fossil range|201|0|earliest=210}}
| image = Monolophosaurus jiangi.jpg
| image_width = 250px
| image_caption = Skeleton of ''[[Monolophosaurus jiangi]]''
| authority = [[Jacques Gauthier|Gauthier]], 1986
| subdivision_ranks = Subgroups<ref name="mateus">{{Cite journal | doi = 10.1371/journal.pone.0088905| title = ''Torvosaurus gurneyi'' n. sp., the Largest Terrestrial Predator from Europe, and a Proposed Terminology of the Maxilla Anatomy in Nonavian Theropods| url = http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0088905| editor-last = Evans | editor-first = Alistair Robert| journal = PLoS ONE| volume = 9| issue = 3| pages = e88905| year = 2014| last1 = Hendrickx | first1 = C. | last2 = Mateus | first2 = O.V. | pmid = 24598585 | pmc=3943790| bibcode = 2014PLoSO...988905H}}</ref><ref name="carrano">{{Cite journal | last1 = Carrano | first1 = M. T. | last2 = Benson | first2 = R. B. J. | last3 = Sampson | first3 = S. D. | doi = 10.1080/14772019.2011.630927 | title = The phylogeny of Tetanurae (Dinosauria: Theropoda) | journal = Journal of Systematic Palaeontology | volume = 10 | issue = 2 | pages = 211–300| year = 2012 | pmid = | pmc = }}</ref>
| subdivision =
*†''[[Chilesaurus]]''?<ref>{{Cite journal | doi = 10.1038/nature14307| pmid = 25915021| title = An enigmatic plant-eating theropod from the Late Jurassic period of Chile| journal = Nature| volume = 522| issue = 7556| pages = 331| year = 2015| last1 = Novas | first1 = F. E. | last2 = Salgado | first2 = L. | last3 = Suárez | first3 = M. | last4 = Agnolín | first4 = F. L. | last5 = Ezcurra | first5 = M. N. D. | last6 = Chimento | first6 = N. S. R. | last7 = de la Cruz | first7 = R. | last8 = Isasi | first8 = M. P. | last9 = Vargas | first9 = A. O. | last10 = Rubilar-Rogers | first10 = D. | bibcode = 2015Natur.522..331N}}</ref>
*†''[[Sinosaurus]]''?
*†''[[Cryolophosaurus]]''?
*†''[[Chuandongocoelurus]]''
*†''[[Monolophosaurus]]''
*†''[[Cruxicheiros]]''?<ref name="benson">{{Cite journal | doi = 10.4202/app.2009.0083| title = A New Large-Bodied Theropod Dinosaur from the Middle Jurassic of Warwickshire, United Kingdom| journal = Acta Palaeontologica Polonica| volume = 55| pages = 35–42| year = 2010| last1 = Benson | first1 = R. B. J. | last2 = Radley | first2 = J. D. }}</ref>
*†''[[Kaijiangosaurus]]''?
*[[Orionides]]
**{{extinct}}[[Megalosauroidea]]
**[[Avetheropoda]]
***{{extinct}}[[Carnosauria]]
***[[Coelurosauria]]
| synonyms_ref =
| synonyms =
'''Avipoda''' Novas, 1992
}}
'''Tetanurae''' (/ˌtɛtəˈnjuːriː/ or "stiff tails") is a clade that includes most [[theropod]] [[dinosaur]]s, including [[tyrannosaurid]]s, [[megalosaurid]]s, [[ornithomimid]]s, [[allosaurid]]s, [[maniraptora]], and [[Aves]].<ref>{{Cite web|url=http://www.ucmp.berkeley.edu/diapsids/saurischia/tetanurae.html|title=Tetanurae|website=www.ucmp.berkeley.edu|access-date=2016-06-03}}</ref> Tetanurans are defined as all theropods more closely related to modern birds than to ''[[Ceratosaurus]]'' and contain the majority of predatory dinosaur diversity.<ref name=":0">{{Cite journal|last=Carrano|first=Matthew T.|last2=Benson|first2=Roger B. J.|last3=Sampson|first3=Scott D.|date=2012-06-01|title=The phylogeny of Tetanurae (Dinosauria: Theropoda)|url=https://dx.doi.org/10.1080/14772019.2011.630927|journal=Journal of Systematic Palaeontology|volume=10|issue=2|pages=211–300|doi=10.1080/14772019.2011.630927|issn=1477-2019}}</ref> Tetanurae and Ceratosauria, the other group of theropods, likely diverged during the late Triassic.<ref name=":1">{{Cite journal|last=Sereno P.C., Wilson J.A., Larsson, H.C.E., Dutheil D.B., & H. Sues.|date=1994|title=Early Cretaceous Dinosaurs from the Sahara|url=|journal=Science |volume=266|issue=5183|pages=267–71|doi=10.1126/science.266.5183.267|pmid=17771449|bibcode=1994Sci...266..267S}}</ref> Tetanurae first appeared in the fossil record by the Early Jurassic about 190 mya and by the Middle Jurassic had become globally distributed.<ref name=":0" />

The group was named by Jacques Gauthier in 1986 and originally had two main subgroups: Carnosauria and Coelurosauria, the clade containing birds and related dinosaurs such as compsognathids, tyrannosaurids, ornithomimosaurs, and maniraptorans.<ref>{{Cite journal|date=1986-01-01|title=Saurischian monophyly and the origin of birds|url=http://biostor.org/reference/110202|journal=Memoirs of the California Academy of Sciences|volume=8|issn=0885-4629}}</ref> The original Carnosauria was a polyphyletic group including any large carnivorous theropod.<ref name=":2">{{Cite web|url=http://palaeos.com/vertebrates/theropoda/tetanurae.html#Tetanurae.|title=Palaeos Vertebrates Theropoda: Basal Tetanurae|website=palaeos.com|access-date=2016-06-03}}</ref> Many of Gauthier’s carnosaurs, such as tyrannosaurids, have since been re-classified as coelurosaurs or primitive tetanurans.<ref name=":0" /> Carnosauria has been reclassified as a group containing allosaurids that split from the Coelurosauria at the Neotetanurae/Avetheropoda node.<ref name=":0" /> Members of Spinosauroidea are believed to represent basal tetanurans.<ref name=":0" />

Tetanuran evolution was characterized by parallel diversification of multiple lineages, repeatedly attaining large body size and similar locomotor morphology.<ref name=":0" /> ''Cryolophosaurus'' has been claimed as the first true member of the group, but subsequent studies have disagreed on whether it is a dilophosaurid or tetanuran.<ref name=":0" /><ref>{{Cite journal|author1=Smith N. D. |author2=Hammer W.R. |author3=P.J. Currie |last-author-amp=yes |date=2005|title=Osteology and phylogenetic relationships of Cryolophosaurus ellioti (Dinosauria: Theropoda): Implications for basal theropod evolution|url=|journal=Journal of Vertebrate Paleontology |volume=25 |issue=3 |pages=1|doi=10.1080/02724634.2005.10009942|pmid=}}</ref> Arcucci and Coria (2003) classified ''Zupaysaurus'' as an early tetanuran,<ref>{{Cite journal|last=Arcucci|first=Andrea B.|last2=Coria|first2=Rodolfo A.|date=2013-04-19|title=A new Triassic carnivorous dinosaur from Argentina|url=http://www.ameghiniana.org.ar/index.php/ameghiniana/article/viewArticle/955|journal=Ameghiniana|volume=40|issue=2|issn=1851-8044}}</ref> but it was later placed as a sister taxon to the clade containing dilophosaurids, ceratosaurs, and tetanurans.<ref>{{Cite journal|last=Hans-Dieter Sues, Sterling J. Nesbitt, David S. Berman and Amy C. Henrici|date=2011|title=A late-surviving basal theropod dinosaur from the latest Triassic of North America|url=|journal=Proceedings of the Royal Society B |volume=278 |issue=1723|year=1723 |pages=3459–3464|doi=10.1098/rspb.2011.0410|pmid=21490016|pmc=3177637}}</ref>

Shared tetanuran features include a ribcage indicating a sophisticated air-sac-ventilated lung system similar to that in modern birds.<ref name=":2" /><ref name=":4">{{Cite book|title=The Dinosauria|author1=Weishampel D.B., Dodson P. |author2=H. Osmólska |lastauthoramp=yes |publisher=University of California Press|year=|isbn=9780520941434|location=2004|pages=}}</ref> This character would have been accompanied by an advanced circulatory system.<ref name=":2" /><ref name=":4" /> Other tetanuran characterizing features include the absence of the fourth digit of the hand, placement of the maxillary teeth anterior to the orbit, a strap-like scapula, maxillary fenestrae, and stiffened tails.<ref name=":1" /><ref name=":2" /><ref name=":4" /> During the Late Jurassic and Early Cretaceous, large spinosaurids and allosaurids flourished but possibly died out before the end of the Cretaceous, perhaps due to competition from ceratosaurs and tyrannosaurids.<ref name=":5">{{Cite journal|last=Zanno|first=Lindsay E.|last2=Makovicky|first2=Peter J.|date=2013-11-22|title=Neovenatorid theropods are apex predators in the Late Cretaceous of North America|url=http://www.nature.com/ncomms/2013/131122/ncomms3827/full/ncomms3827.html|journal=Nature Communications|volume=4|pages=2827|doi=10.1038/ncomms3827|pmid=24264527|bibcode=2013NatCo...4E2827Z}}</ref> Coelurosaurs persisted until the end of the Mesozoic Era, when all except for crown clade avians died out.

==Morphology==
[[File:Monolophosaurus jiangi jmallon.jpg|thumb|left|Illustration of ''[[Monolophosaurus jiangi]]'']]

===Anatomy===
Tetanurans have two basic skull morphologies.<ref name=":0" /> The first skull type, typical in large theropods such as ''Allosaurus'', is common within ceratosaurs and may be primitive for tetanurans. In this type, the skull is about three times longer than tall, with a blunter snout and frequent elaborations such as horns or spikes along the lacrimals, nasals, and frontals.<ref name=":0" /> In the second skull type, the skull is lower and longer, with a less elaborated skull roof and a more elongated snout.<ref name=":0" /> Shared tetanuran features include the maxillary fenestra (an opening in the antorbital fossa), a pneumatic excavation in the jugal, and the position of the maxillary teeth anterior to the orbit.<ref name=":1" /> The posterior skull is little modified in tetanurans, except within Spinosauridae.

In the postcranial skeleton, tetanurans transition between the most primitive theropod morphologies in basal tetanurans towards more derived, bird-like states in coelurosaurs.<ref name=":0" /> Most tetanurans possess specialized wrist bones, the absence or reduction of the fourth digit of the hand, a strap-like scapula, stiffened tails, and a laminar astragalar ascending process.<ref name=":1" /><ref name=":2" /><ref name=":4" /> Advanced tetanurans would have possessed a sophisticated air-sac-ventilated lung system similar to birds, and an advanced circulatory system.<ref name=":2" /><ref name=":4" /> In megalosaurids and allosaurids, the orientation of the femoral head is anteromedial such as in ceratosaurs, but in avetheropods this orientation is fully medial.<ref name=":0" /> Tetanuran locomotor morphology is relatively generalized, with few variations between taxa.<ref name=":0" />

===Body Size===
Basal tetanurans were the first theropod clade to achieve truly giant body sizes, with both megalosauroid and allosuroid taxa weighing over 1 ton.<ref name=":0" /> Sequential temporal appearances of large body size in subsequent clades suggest a pattern of size-cycles, with the extinction of incumbent giant forms allowing for replacement with a new, more bird-like theropod group that then also evolved giant body size.<ref name=":0" /> It is however possible that more than one giant tetanuran existed at a time in the same paleoenvironment, perhaps with feeding habit variations. Within most dinosaur clades, body size tended to increase over time along a lineage according to Cope’s Rule.<ref>{{Cite journal|last=Hone|first=D. W. E.|last2=Keesey|first2=T. M.|last3=Pisani|first3=D.|last4=Purvis|first4=A.|date=2005-05-01|title=Macroevolutionary trends in the Dinosauria: Cope's rule|journal=Journal of Evolutionary Biology|volume=18|issue=3|pages=587–595|doi=10.1111/j.1420-9101.2004.00870.x|issn=1010-061X|pmid=15842488}}</ref> Coelurosaurian theropods are the notable exception to the pattern of body size increases.<ref name=":0" />

==History of Study==

===History of Classification===
Tetanurae was recognized and named by Gauthier in 1986.<ref name=":6">{{Cite journal|last=Gauthier, J. A.|date=1986-01-01|title=Saurischian monophyly and the origin of birds|url=http://biostor.org/reference/110202|journal=Memoirs of the California Academy of Sciences|volume=8|issn=0885-4629}}</ref> The earliest discovered tetanuran is the earliest named dinosaur, ''Megalosaurus''.<ref name=":0" /> For a century after the description of ''Megalosaurus'', most large carnivorous dinosaurs were serially arrayed into the family Megalosauridae within the order Theropoda.<ref name=":0" /> In 1914, Friedrich von Huene separated small, lightly built forms into the infraorder Coelurosauria and larger taxa into the infraorder Pachypodosauria.<ref name=":0" /> Later, he transferred large, carnivorous taxa to the new infraorder Carnosauria, which came to include all known large-bodied carnivores other than ''Ceratosaurus''.<ref name=":0" /> The size-based arrangement persisted until Gauthier, who redefined Carnosauria and Coelurosauria based on new cladistic analyses but retained the terms.<ref name=":6" /> Gauthier defined Coelurosauria as a taxon comprising birds and theropods closer to birds than to Carnosauria, and listed within Carnosauria several large-bodied theropod taxa but did not formally define the group.<ref name=":6" /> Many of these original carnosaurs have since been reclassified as coelurosaurs or primitive tetanurans, and Carnosauria has now been defined as ''Allosaurus'' and all Avetheropods closer to ''Allosaurus'' than to birds.<ref name=":7">{{Cite book|title=Encyclopedia of Dinosaurs|author1=Currie P.J. |author2=K. Padian |lastauthoramp=yes |publisher=Academic Press|year=1997|isbn=9780080494746|location=|pages=}}</ref>

Initial cladistics studies supported the arrangement of primitive megalosaurs as serial outgroups to a clade of allosaurids, followed by the Coelurosauria. Subsequent studies have discovered that many of these basal tetanurans formed a true clade, termed Megalosauroidea or alternatively Spinosauroidea.<ref name=":0" />

===Current Phylogeny===
Current phylogeny agrees on a monophyletic Tetanurae that includes a series of generally large-bodied basal taxa outside a monophyletic Coelurosauria.<ref name=":0" /> Coelophysoids are basal to Tetanurae, with Ceratosauria forming a sister taxa that diverged during the late Triassic.<ref name=":1" />

After their initial appearance, Tetanurae radiated into two main clades, Spinosauroidea or Megalosauroidea and Avetheropoda or Neotetanurae.<ref name=":7" /> Spinosauroidea are believed to represent basal Tetanurans. At the Neotetanurae/Avetheropoda node, allosaurids split from the Coelurosauria. Tyrannosauridae has been placed within Coelurosauria. The allosaurids and their closest relatives form a reconstituted Carnosauria.<ref name=":0" /> Debate persists about whether the allosaurids form a clade with spinosauroids/megalosauroids, and whether Allosauroidea belongs in Avetheropoda with Coelurosauria or forms a sister taxa to Megalosauroidea, and whether Megalosauroidea forms a valid clade.<ref name=":0" />

The [[cladogram]] presented below follows a phylogenetic analysis published by Zanno and Makovicky in 2013.<ref name=":5" />
{{clade | style=font-size:85%;line-height:80%
|label1=Tetanurae
|1={{clade
|1=†''[[Cryolophosaurus]]''[[File:Cryolophosaurus in Japan White Background.jpg|50 px]]
|2=†''[[Sinosaurus]]''[[File:Sinosaurus triassicus white background.JPG|50 px]]
|3={{clade
|1=†''[[Chuandongocoelurus]]''
|2=†''[[Monolophosaurus]]''[[File:Monolophosaurus jiangi White Background.jpg|50 px]]
|label3=[[Orionides]]
|3={{clade
|label1=†[[Megalosauroidea]]
|1={{clade
|1=†[[Piatnitzkysauridae]]
|label2=†[[Megalosauria]]
|2={{clade
|1=†[[Spinosauridae]][[File:Spinosaurus white background.jpg|50px]]
|2=†[[Megalosauridae]][[File:Complete skeleton of Torvosaurus white background.jpg|50px]]}} }}
|label2=[[Avetheropoda]]
|2={{clade
|1=[[Coelurosauria]][[File:FMNH Deinonychus white background.JPG|50 px]]
|label2=†[[Allosauroidea]]
|2={{clade
|1=†[[Metriacanthosauridae]]
|label2=†[[Allosauria]]
|2={{clade
|1=†[[Allosauridae]][[File:Allosaurus AMNH White Background.jpg|50px]]
|label2=†[[Carcharodontosauria]]
|2={{clade
|1=†[[Neovenatoridae]]
|2=†[[Carcharodontosauridae]][[File:Acrocanthosaurus white background.jpg|50px]]
}} }} }} }} }} }} }} }}

==Paleobiology==

===Biogeography===
The biogeographical history of non-avian Tetanurae spans over 110 million years and all continents.<ref name=":0" /> The presence of major lineages prior to the breakup of Pangaea implies wide dispersal of these clades, with later absences indicating regional extinctions or dispersal failure.<ref name=":0" /> The density of sampling is currently insufficient to provide a detailed analysis of biogeographical evolution for the Tetanurae.<ref name=":0" />

===Diversity===
Tetanurae and Ceratosauria likely diverged during the late Triassic, more than 200 mya. By the Early Jurassic, Tetanurae fossils appear in the fossil record and reached global distribution by the Middle Jurassic.<ref name=":0" /> In the Late Jurassic, the fossil record demonstrates widespread presence of multiple clades within both megalosauroids and avetheropods.<ref name=":0" /> The Megalosauroidea contained high diversity with two Jurassic clades, Piatnitzkysauridae and Megalosauridae, as well as the Cretaceous Spinosauridae.<ref name=":0" /> Tetanuran evolution appears to exhibit waves of diversification, although this may be due to uneven sampling.<ref name=":0" /> During the Late Jurassic and Early Cretaceous, large spinosaurids and allosaurids flourished, but the latter possibly died out before the end of the Cretaceous due to the [[Cenomanian-Turonian boundary event]], while spinosaurids are known from the Santonian. Soon afterwards the niche of terrestrial apex predator ceratosaurs and tyrannosaurid coelurosaurs, which dominated terminal Cretaceous terrestrial ecosystems.<ref name=":5" /> Coelurosaurs persisted through the end of the Mesozoic Era.<ref name=":5" /> Modern birds are the only living representatives of the Tetanurae.<ref name=":5" />

==References==
{{reflist}}

{{Portal|Dinosaurs}}

[[Category:Tetanurans| ]]
[[Category:Hettangian first appearances]]
[[Category:Extant Early Jurassic first appearances]]

Anchiornithidae

2017-12-08T23:43:03Z

Wikkler: Grammar correction

{{use dmy dates|date=December 2017}}
{{Automatic taxobox
| name = Anchiornithidae
| fossil_range = [[Late Jurassic]]–[[Early Cretaceous]] {{Fossil range|164|122}}
| image = Anchiornis huxleyi skel hartman 2017.png
| image_width = 250px
| image_caption = Skeletal restoration of ''[[Anchiornis huxleyi]]'' by Scott Hartman, 2017
| authority = [[Xu Xing (paleontologist)|Xu]] et al. 2016 ''sensu'' Foth & Rauhut, 2017
| type_species = {{extinct}}''[[Anchiornis huxleyi]]''
| type_species_authority = [[Xu Xing (paleontologist)|Xu]] ''et al.'', 2009
| subdivision_ranks = Genera
| subdivision =
*{{extinct}}''[[Anchiornis]]''
*{{extinct}}''[[Aurornis]]''
*{{extinct}}''[[Eosinopteryx]]''
*{{extinct}}''[[Ostromia]]''
*{{extinct}}''[[Pedopenna]]''
*{{extinct}}''[[Serikornis]]''
*{{extinct}}''[[Xiaotingia]]''
*{{extinct}}''[[Yixianosaurus]]''
| synonyms =
*'''Tetrapterygidae''' ([[Sankar Chatterjee|Chatterjee]], 2015)
}}
'''Anchiornithidae''' is a family of [[Paraves|eumaniraptoran]]s that are the sister taxon to the [[bird]] clade [[Avialae]].<ref name="desc">{{cite journal | last1 = Foth | first1 = C. | last2 = Rauhut | first2 = O. W. M. | title = Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs | journal = BMC Evolutionary Biology | volume = 17 | page = 236 | date = 2017 | doi = 10.1186/s12862-017-1076-y | url = https://link.springer.com/article/10.1186/s12862-017-1076-y}}</ref> In the past anchiornithids were classified at various position of the [[Maniraptora|maniraptoran]] tree, with some scientists classifying them as a subfamily of [[Troodontidae]]<ref name=Xiaotingia>{{cite journal |authors=Xing Xu, Hailu You, Kai Du and Fenglu Han |title=An ''Archaeopteryx''-like theropod from China and the origin of Avialae |url=http://www.ivpp.ac.cn/qt/papers/201403/P020140314389417822583.pdf |date=28 July 2011 |journal=Nature |volume=475 |pages=465–470 |doi=10.1038/nature10288 |issue=7357 |pmid=21796204}}</ref><ref name=leeetal2011>{{cite journal|author=Lee, M. S. Y. and Worthy, T. H. |title=Likelihood reinstates ''Archaeopteryx'' as a primitive bird|journal=Biology Letters|volume=8|issue=2|pages=299–303|doi=10.1098/rsbl.2011.0884|pmid=22031726|pmc=3297401|year=2011}}</ref><ref>{{Cite journal|author1=Stephen L. Brusatte |author2=Graeme T. Lloyd |author3=Steve C. Wang |author4=Mark A. Norell |year=2014 |title=Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition |journal=Current Biology |volume=24 |issue=20 |pages=2386–2392 |doi=10.1016/j.cub.2014.08.034 |pmid=25264248}}</ref><ref name=Shenetal2017>{{Cite journal|author1=Caizhi Shen |author2=Junchang Lü |author3=Sizhao Liu |author4=Martin Kundrát |author5=Stephen L. Brusatte |author6=Hailong Gao |year=2017 |title=A new troodontid dinosaur from the Lower Cretaceous Yixian Formation of Liaoning Province, China |journal=Acta Geologica Sinica (English Edition) |volume=91 |issue=3 |pages=763–780 |url=http://www.geojournals.cn/dzxben/ch/reader/view_abstract.aspx?file_no=2017endzxb03001&flag=1 }}</ref><ref name=Shenetal20172>{{Cite journal|author1=Cai-zhi Shen |author2=Bo Zhao |author3=Chun-ling Gao |author4=Jun-chang Lü |author5=Martin Kundrát |year=2017 |title=A New Troodontid Dinosaur (''Liaoningvenator curriei'' gen. et sp. nov.) from the Early Cretaceous Yixian Formation in Western Liaoning Province |journal=Acta Geoscientica Sinica |volume=38 |issue=3 |pages=359–371 |doi=10.3975/cagsb.2017.03.06 }}</ref>, [[Archaeopterygidae]]<ref name=Xiaotingia>{{cite journal |authors=Xing Xu, Hailu You, Kai Du and Fenglu Han |title=An ''Archaeopteryx''-like theropod from China and the origin of Avialae |url=http://www.ivpp.ac.cn/qt/papers/201403/P020140314389417822583.pdf |date=28 July 2011 |journal=Nature |volume=475 |pages=465–470 |doi=10.1038/nature10288 |issue=7357 |pmid=21796204}}</ref>, or an assemblage of dinosaurs that are an evolutionary grade within [[Avialae]].<ref name=wangetal2016>{{cite journal | last1 = Wang | first1 = M. | last2 = Wang | first2 = X. | last3 = Wang | first3 = Y. | last4 = Zhou | first4 = Z. | year = 2016 | title = A new basal bird from China with implications for morphological diversity in early birds | journal = Scientific Reports | volume = 6 | issue = | page = 19700 | doi = 10.1038/srep19700 | pmid=26806355 | pmc=4726217}}</ref>

In 2015 Chatterjee created '''Tetrapterygidae''' in the second edition of his book ''The Rise of Birds: 225 Million Years of Evolution'', where he included ''[[Xiaotingia]]'', ''[[Aurornis]]'', ''[[Anchiornis]]'', and even ''[[Microraptor]]''; together they were proposed to be the sister group of the [[Avialae]].<ref>Chatterjee, S. (2015). The rise of birds: 225 million years of evolution. Johns Hopkins University Press, 45–48.</ref> However this family is invalid as must include the genus ''Tetrapteryx'', which is the junior synonymn of ''[[Grus (genus)|Grus]]'' – therefore Tetrapterygidae is a junior synonym of [[Gruidae]].<ref>{{cite web |author=Matthew Martyniuk |date=May 24, 2015 |title=The Crane and the Microraptor |publisher=DinoGoss: A blog about stem-birds |url=http://dinogoss.blogspot.com/2015/05/the-crane-and-microraptor.html?utm_source=feedburner&utm_medium=feed&utm_campaign=Feed%3A+Dinogoss+%28DinoGoss%29 |archiveurl=https://web.archive.org/web/20150616185940/http://dinogoss.blogspot.com/2015/05/the-crane-and-microraptor.html?utm_source=feedburner&utm_medium=feed&utm_campaign=Feed%3A+Dinogoss+%28DinoGoss%29 |archivedate=16 June 2015}}</ref>

The clade was originally named as "'''Anchiornithinae'''" by Xu et al. (2016) and defined as for "the most inclusive clade including ''[[Anchiornis]]'' but not ''[[Archaeopteryx]]'', ''[[Junglefowl|Gallus]]'', ''[[Troodon]]'', ''[[Dromaeosaurus]]'', ''[[Unenlagia]]'', or ''[[Epidexipteryx]]''".<ref>Xu et al. (2016) An Updated Review of the Middle-Late Jurassic Yanliao Biota: Chronology, Taphonomy, Paleontology and Paleoecology. ''Acta Geologica Sinica'' Vol. 90 No. 6 pp.2229–2243.</ref>

In 2017 Foth and Rauhut in their re-evaluation of the Haarlem ''[[Archaeopteryx]]'' specimen (which they classified it in its own distinct genus ''[[Ostromia]]''<ref name="desc"/>) found that the anchiornithids are a distinct family closer to the ancestry of birds.<ref name="desc"/> They provided their own definition of Anchiornithidae as "as all maniraptoran [[theropod]]s that are more closely related to ''[[Anchiornis huxleyi]]'' than to ''[[Passer domesticus]]'', ''[[Archaeopteryx lithographica]]'', ''[[Dromaeosaurus albertensis]]'', ''[[Troodon formosus]]'', or ''[[Oviraptor philoceratops]]''."<ref name="desc"/>

Below is the phylogeny of the group based on the description of ''[[Halszkaraptor]]'' from Cau et al. (2017), who incorporate a large-scale phylogenetic matrix in their study and found support for Anchiornithidae being a distinctive family:<ref name="cauetal2017">{{cite journal|url=https://www.nature.com/articles/nature24679|title=Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs|first1=A.|last1=Cau|first2=V.|last2=Beyrand|first3=D.|last3=Voeten|first4=V.|last4=Fernandez|first5=P.|last5=Tafforeau|first6=K.|last6=Stein|first7=R.|last7=Barsbold|first8=K.|last8=Tsogtbaatar|first9=P.|last9=Currie|first10=P.|last10=Godefroit|year=2017|publisher=|journal=Nature}}</ref>

{{clade| style=font-size:85%; line-height:85%;
|label1=[[Avialae]]
|1={{clade
|label1='''Anchiornithidae'''
|1={{clade
|1={{clade
|1=''[[Yixianosaurus longimanus]]''
|2=''[[Xiaotingia zhengi]]'' }}
|2={{clade
|1=''[[Anchiornis huxleyi]]''
|2={{clade
|1=''[[Eosinopteryx brevipenna]]''
|2={{clade
|1=''[[Aurornis xui]]''
|2=''[[Serikornis sungei]]'' }} }} }} }}
|2={{clade
|1=''[[Archaeopteryx lithographica]]''
|2={{clade
|1=[[Scansoriopterygidae]]
|2={{clade
|1=''[[Rahonavis ostromi]]''
|2={{clade
|1=''[[Balaur bondoc]]''
|2={{clade
|1=[[Jeholornithidae]]
|2=[[Pygostylia]] }} }} }} }} }} }} }}

==References==
{{reflist}}

[[Category:Paravians| 01]]
[[Category:Late Jurassic dinosaurs]]
[[Category:Jurassic birds]]
[[Category:Coelurosaurs]]
[[Category:Theropods]]

Laevisuchus

2017-12-08T23:41:40Z

Wikkler: Typo correction

{{italic title}}{{automatic taxobox
| name = ''Laevisuchus''
| fossil_range = [[Late Cretaceous]], {{fossilrange|70|latest=66}}
| image = Laevisuchus indicus.jpg
| image_caption = Vertebra
| authority = [[Friedrich von Huene|Huene]] & [[Charles Alfred Matley|Matley]], 1933
| type_species = {{extinct}}'''''Laevisuchus indicus'''''
| type_species_authority = [[Friedrich von Huene|Huene]] & [[Charles Alfred Matley|Matley]], 1933
}}
'''''Laevisuchus''''' ({{IPAc-en|ˌ|l|ɛ|v|ɪ|ˈ|sj|uː|k|ə|s}} {{respell|LEV|i|SEW|kəs}}, "light crocodile") is a [[genus]] of [[abelisauroidea|abelisauroid]] [[Theropoda|theropod]] [[dinosaur]] from the [[Late Cretaceous]]. Its remains were discovered by [[Charles Alfred Matley]] near [[Jabalpur]] in [[Maastrichtian]] deposits in the [[Lameta Formation]] in [[India]], and were named and described by [[paleontologist]]s [[Friedrich von Huene]] and Matley in 1933.<ref>F. v. Huene and C. A. Matley, 1933, "The Cretaceous Saurischia and Ornithischia of the Central Provinces of India", ''Palaeontologica Indica (New Series), Memoirs of the Geological Survey of India'' '''21'''(1): 1-74</ref> The [[type species]] is '''''Laevisuchus indicus'''''. The generic name is derived from [[Latin]] ''laevis'', "light" and the Greek name for the Egyptian crocodile god, ''Soukhos''. The [[specific name (zoology)|specific name]] means "Indian" in [[Latin]]. It is known only from three cervical [[vertebrae]] ('''GSI K20/613, GSI K20/614 and GSI K27/696''') and a dorsal vertebra ('''GSI K27/588'''). A [[holotype]] was not assigned by Huene and Matley and a [[lectotype]] has never been chosen from the [[syntype]]s. All remains except GSI K27/696 were lost; GSI K20/613 was rediscovered in 2012.

==Description==
''Laevisuchus'' was a small bipedal carnivore. In 1998 [[David Lambert (author)|David Lambert]] estimated it was some {{convert|2|m|ft|spell=in}} long, {{convert|0.9|m|ft|abbr=off|sp=us}} high, and approximately {{convert|30|kg|lb|abbr=on}} in weight.<ref name="lambert1998">Lambert, D. (1998). ''The Wordsworth Book of Dinosaurs'', Britain: Mackays of Chatham PLC.</ref>

==Classification==
''Laevisuchus'' was originally classified by Huene as a [[coelurid]] due to the similarity of its vertebrae with those of "[[Aristosuchus]]". However, an analysis in 2004 has shown it to be an [[Abelisaur|abelisauroid]] because of its long epipophyses, a pair of foramina on the centrum, and low and triangular neural spines. The vertebrae specifically resemble those of [[noasaurid|noasaurids]] like ''[[Masiakasaurus]]'' and ''[[Noasaurus]]'' due to having more anteriorly placed neural spines and posteriorly reduced [[epipophyses]] <ref name="tykoskirowe2004">Tykoski, R.S. & Rowe, T. (2004). "Ceratosauria". In: Weishampel, D.B., Dodson, P., & Osmolska, H. (Eds.) ''The Dinosauria'' (2nd edition). Berkeley: University of California Press. Pp. 47–70 {{ISBN|0-520-24209-2}}</ref><ref>F. E. Novas, F. L. Agnolin, and S. Bandyopadhyay, 2004, "Cretaceous theropods from India: a review of specimens described by Huene and Matley (1933)", ''Revista del Museo Argentino de Ciencias Naturales, nuevo serie'' '''6'''(1): 67-103</ref>

==See also==
{{Portal|Dinosaurs}}
* [[Timeline of ceratosaur research]]

==References==
{{Reflist}}

==External links==
*[http://dinosaurier-info.de/animals/dinosaurs/pages_l/laevisuchus.php Dinosaurier-Info] {{de icon}}

{{Ceratosauria}}

[[Category:Late Cretaceous dinosaurs]]
[[Category:Abelisaurs]]
[[Category:Dinosaurs of India and Madagascar]]
[[Category:Fossil taxa described in 1933]]
[[Category:Taxa named by Friedrich von Huene]]
[[Category:Taxa named by Charles Alfred Matley]]

{{theropod-stub}}

Halszka Osmólska

2017-12-06T22:52:34Z

Wikkler: I spelled the name wrong.

{{Infobox person
|name=Halszka Osmólska
|image=Halszka Osmólska.jpg
|birth_date={{birth date|1930|9|15}}
|birth_place=[[Poznań]], [[Poland]]
|death_date={{death date and age|2008|3|31|1930|9|15}}
|death_place=[[Poland]]
|occupation=[[Paleontologist]]
}}

'''Halszka Osmólska''' (September 15, 1930 – March 31, 2008) was a [[Poles|Polish]] [[paleontologist]] who had specialized in [[Mongolia]]n [[dinosaur]]s.<ref name=DL90>{{cite book |last=Lambert |first=David |author2=the Diagram Group |title=The Dinosaur Data Book |year=1990 |publisher=Avon Books |location=New York |isbn=0-380-75896-2 |chapter=Dinosaurologists |pages=281}}</ref>

She was born in [[Poznań]]. A member of the 1965 and 1970 Polish–Mongolian expeditions to the [[Gobi Desert]],<ref name=EHC00>{{cite book |last=Colbert |first=Edwin H. |authorlink=Edwin H. Colbert |editor=Benton, Michael J. |editor2=Shishkin, Mikhail A. |editor3=Unwin, David M. |editor4=Kurochkin, Evgenii N. |title=The Age of Dinosaurs in Russia and Mongolia |year=2000 |publisher=Cambridge University Press |location=Cambridge |isbn=0-521-55476-4 |pages=211–234 |chapter=Asiatic dinosaur rush }}</ref> she described many finds from these rocks, often with [[Teresa Maryańska]]. Among the dinosaurs she described are: ''[[Elmisaurus]]'' (and [[Elmisauridae]] (1981), ''[[Hulsanpes]]'' (1982), ''[[Borogovia]]'' (1987), and ''[[Bagaraatan]]'' (1996); with Maryańska, ''[[Homalocephale]]'', ''[[Prenocephale]]'', and ''[[Tylocephale]]'' (and [[Pachycephalosauria]]) (1974), ''[[Bagaceratops]]'' (1975), and ''[[Barsboldia]]'' (1981); with Maryańska and [[Altangerel Perle]], ''[[Goyocephale]]'' (1982); with [[Ewa Roniewicz]], ''[[Deinocheirus]]'' (1967); with Roniewicz and [[Rinchen Barsbold]], ''[[Gallimimus]]'' (1972);<ref name=DL90/> with Kurzanov, ''[[Tochisaurus]]'' (1991); and with several other authors, ''[[Nomingia]]'' (2000). Her other work included discussions of the [[paleobiology]] of [[hadrosaurid]]s, and co-editing the two editions of ''[[The Dinosauria]]''.<ref name=dinosauria1>{{cite book |editor1=Weishampel, David B. |editor2=Osmólska, Halszka |editor3=Dodson, Peter |title=The Dinosauria |edition=1st |year=1990 |publisher=University of California Press |location=Berkeley |isbn=0-520-06727-4 }}</ref><ref name=dinosauria2>{{cite book |editor1=Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |edition=2nd |year=2004 |publisher=University of California Press |location=Berkeley |isbn=0-520-24209-2 }}</ref> As of 2004, she was affiliated with the [[Instytut Paleobiologii]] of the [[Polska Akademia Nauk]].<ref name=dinosauria2b>{{cite book |editor1=Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |edition=2nd |year=2004 |publisher=University of California Press |location=Berkeley |isbn=0-520-24209-2 |pages=776 }}</ref> She is recognized for her work in the names of the Mongolian [[oviraptoridae|oviraptorid]] ''[[Citipati (dinosaur)|Citipati osmolskae]]'', the [[People's Republic of China|Chinese]] [[dromaeosauridae|dromaeosaurid]] ''[[Velociraptor|Velociraptor osmolskae]]'', the [[Mongolia|Mongolian]] dromaeosaurid ''[[Halszkaraptor|Halszkaraptor escuilliei]]'', the [[archosauriformes|archosauriform]] reptile ''[[Osmolskina|Osmolskina czatkowicensis]]'', and the Polish [[Pliocene]] [[Lagomorpha|lagomorph]] ''[[Prolagus osmolskae]]''.<ref>Fostowicz-Frelik, Ł. 2010. A new species of Pliocene Prolagus (Lagomorpha, Ochotonidae) from Poland is the northernmost record of the genus. Journal of Vertebrate Paleontology 30 (2): 609-612.</ref>

Osmólska was, in recognition of her scientific work, a recipient of a number of awards including the [[Polish Cross of Merit]].<ref>Moody, R., & Geological Society of London. (2010). ''Dinosaurs and other extinct saurians: A historical perspective''. London: Geological Society.</ref>

==Selected publications==
* H. Osmólska and E. Roniewicz (1970). Deinocheiridae, a new family of theropod dinosaurs. ''Palaeontologica Polonica'' '''21''':5-19.
* H. Osmólska, E. Roniewicz, and R. Barsbold (1972). A new dinosaur, ''Gallimimus bullatus'' n. gen., n. sp. (Ornithomimidae) from the Upper Cretaceous of Mongolia. ''Palaeontologia Polonica'' '''27''':103-143.
* H. Osmólska (1972). Preliminary note on a crocodilian from the Upper Cretaceous of Mongolia. ''Palaeontologia Polonica'' '''27''':43-47.
* T. Maryańska and H. Osmólska (1974). Pachycephalosauria, a new suborder of ornithischian dinosaurs. ''Palaeontologia Polonica'' '''30''':45-102.
* T. Maryańska and H. Osmólska (1975). Protoceratopsidae (Dinosauria) of Asia. ''Palaeontologica Polonica'' '''33''':133-181.
* H. Osmólska (1976). New light on the skull anatomy and systematic position of ''Oviraptor''. ''Nature'' '''262''':683-684.
* H. Osmólska (1981). Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. ''Palaeontologica Polonica'' '''42''':79-95.
* T. Maryańska and H. Osmólska (1981). First lambeosaurine dinosaur from the Nemegt Formation, Upper Cretaceous, Mongolia. ''Acta Palaeontologica Polonica'' '''26'''(3-1):243-255.
* T. Maryańska and H. Osmólska (1981). Cranial anatomy of ''Saurolophus angustirostris'' with comments on the Asian Hadrosauridae (Dinosauria). ''Palaeontologia Polonica'' '''42''':5-24.
* H. Osmólska (1982). ''Hulsanpes perlei'' n.g. n.sp. (Deinonychosauria, Saurischia, Dinosauria) from the Upper Cretaceous Barun Goyot Formation of Mongolia. ''Neues Jahrbuch für Geologie und Paläontologie Monatshefte'' '''1982'''(7):440-448.
* A. Perle, T. Maryańska, and H. Osmólska (1982). ''Goyocephale lattimorei'' gen. et sp. n., a new flat-headed pachycephalosaur (Ornithischia, Dinosauria) from the Upper Cretaceous of Mongolia. ''Acta Palaeontologica Polonica'' '''27'''(1-4):115-127.
* T. Maryańska and H. Osmólska (1984). Postcranial anatomy of ''Saurolophus angustirostris'' with comments on other hadrosaurs. ''Palaeontologia Polonica'' '''46''':119-141.
* T. Maryańska and H. Osmólska (1985). On ornithischian phylogeny. ''Acta Palaeontologica Polonica'' '''30'''(3-4):137-149.
* H. Osmólska (1987). ''Borogovia gracilicrus'' gen. et sp. n., a new troodontid dinosaur from the Late Cretaceous of Mongolia. ''Acta Palaeontologica Polonica'' '''32'''(1-2):133-150.
* R. Barsbold, H. Osmólska, and S.M. Kurzanov (1987). On a new troodontid (Dinosauria, Theropoda) from the Early Cretaceous of Mongolia. ''Acta Palaeontologica Polonica'' '''32'''(1-2):121-132.
* S. M. Kurzanov and H. Osmólska (1991). ''Tochisaurus nemegtensis'' gen. et sp. n., a new troodontid dinosaur (Dinosauria, Theropoda) from Mongolia. ''Acta Palaeontologica Polonica'' '''36'''(1):69-76.
* H. Osmólska (1996). An unusual theropod dinosaur from the Late Cretaceous Nemegt Formation of Mongolia. ''Acta Palaeontologica Polonica'' '''41'''(1):1-38.
* R. Barsbold and H. Osmólska (1999). The skull of ''Velociraptor'' (Theropoda) from the Late Cretaceous of Mongolia. ''Acta Palaeontologica Polonica'' '''44'''(2):189-219.
* R. Barsbold, H. Osmólska, M. Watabe, P.J. Currie, and K. Tsogtbaatar (2000). A new oviraptorosaur (Dinosauria, Theropoda) from Mongolia: the first dinosaur with a pygostyle. ''Acta Palaeontologica Polonica'' '''45'''(2):97-106.
* T. Maryańska, H. Osmólska, and M. Wolsan (2002). Avialan status for Oviraptorosauria. ''Acta Palaeontologica Polonica'' '''47'''(1):97-116.
* H. Osmólska, P.J. Currie, and R. Barsbold (2004). Oviraptorosauria. In: D.B. Weishampel, P. Dodson, and H. Osmólska (eds.), ''The Dinosauria'' (second edition). University of California Press, Berkeley 165-183.

==References==
{{reflist}}

==Further reading==
*Borsuk-Białynicka, M. and Maryańska, T. "Halszka Osmólska (1930-2008) In Memoriam." ''Acta Palaeontologica Polonica'' 53, 2, 206.cp.
*Dodson, Peter (2008) "Polish Women in the Gobi – In Loving Memory of Halszka Osmólska (1930-2008)." ''American Paleontologist'' 16.3: 30

{{Authority control}}

{{DEFAULTSORT:Osmolska, Halszka}}
[[Category:Polish paleontologists]]
[[Category:Polish female geologists]]
[[Category:Women paleontologists]]
[[Category:1930 births]]
[[Category:2008 deaths]]
[[Category:Taxa named by Halszka Osmólska| ]]
[[Category:People from Poznań]]
[[Category:20th-century women scientists]]

Halszka Osmólska

2017-12-06T22:51:50Z

Wikkler: A new species of dromaeosaurid has been named after her.

{{Infobox person
|name=Halszka Osmólska
|image=Halszka Osmólska.jpg
|birth_date={{birth date|1930|9|15}}
|birth_place=[[Poznań]], [[Poland]]
|death_date={{death date and age|2008|3|31|1930|9|15}}
|death_place=[[Poland]]
|occupation=[[Paleontologist]]
}}

'''Halszka Osmólska''' (September 15, 1930 – March 31, 2008) was a [[Poles|Polish]] [[paleontologist]] who had specialized in [[Mongolia]]n [[dinosaur]]s.<ref name=DL90>{{cite book |last=Lambert |first=David |author2=the Diagram Group |title=The Dinosaur Data Book |year=1990 |publisher=Avon Books |location=New York |isbn=0-380-75896-2 |chapter=Dinosaurologists |pages=281}}</ref>

She was born in [[Poznań]]. A member of the 1965 and 1970 Polish–Mongolian expeditions to the [[Gobi Desert]],<ref name=EHC00>{{cite book |last=Colbert |first=Edwin H. |authorlink=Edwin H. Colbert |editor=Benton, Michael J. |editor2=Shishkin, Mikhail A. |editor3=Unwin, David M. |editor4=Kurochkin, Evgenii N. |title=The Age of Dinosaurs in Russia and Mongolia |year=2000 |publisher=Cambridge University Press |location=Cambridge |isbn=0-521-55476-4 |pages=211–234 |chapter=Asiatic dinosaur rush }}</ref> she described many finds from these rocks, often with [[Teresa Maryańska]]. Among the dinosaurs she described are: ''[[Elmisaurus]]'' (and [[Elmisauridae]] (1981), ''[[Hulsanpes]]'' (1982), ''[[Borogovia]]'' (1987), and ''[[Bagaraatan]]'' (1996); with Maryańska, ''[[Homalocephale]]'', ''[[Prenocephale]]'', and ''[[Tylocephale]]'' (and [[Pachycephalosauria]]) (1974), ''[[Bagaceratops]]'' (1975), and ''[[Barsboldia]]'' (1981); with Maryańska and [[Altangerel Perle]], ''[[Goyocephale]]'' (1982); with [[Ewa Roniewicz]], ''[[Deinocheirus]]'' (1967); with Roniewicz and [[Rinchen Barsbold]], ''[[Gallimimus]]'' (1972);<ref name=DL90/> with Kurzanov, ''[[Tochisaurus]]'' (1991); and with several other authors, ''[[Nomingia]]'' (2000). Her other work included discussions of the [[paleobiology]] of [[hadrosaurid]]s, and co-editing the two editions of ''[[The Dinosauria]]''.<ref name=dinosauria1>{{cite book |editor1=Weishampel, David B. |editor2=Osmólska, Halszka |editor3=Dodson, Peter |title=The Dinosauria |edition=1st |year=1990 |publisher=University of California Press |location=Berkeley |isbn=0-520-06727-4 }}</ref><ref name=dinosauria2>{{cite book |editor1=Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |edition=2nd |year=2004 |publisher=University of California Press |location=Berkeley |isbn=0-520-24209-2 }}</ref> As of 2004, she was affiliated with the [[Instytut Paleobiologii]] of the [[Polska Akademia Nauk]].<ref name=dinosauria2b>{{cite book |editor1=Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |edition=2nd |year=2004 |publisher=University of California Press |location=Berkeley |isbn=0-520-24209-2 |pages=776 }}</ref> She is recognized for her work in the names of the Mongolian [[oviraptoridae|oviraptorid]] ''[[Citipati (dinosaur)|Citipati osmolskae]]'', the [[People's Republic of China|Chinese]] [[dromaeosauridae|dromaeosaurid]] ''[[Velociraptor|Velociraptor osmolskae]]'', the [[Mongolia|Mongolian]] dromaeosaurid ''[[Halzskaraptor|Halzskaraptor escuilliei]]'', the [[archosauriformes|archosauriform]] reptile ''[[Osmolskina|Osmolskina czatkowicensis]]'', and the Polish [[Pliocene]] [[Lagomorpha|lagomorph]] ''[[Prolagus osmolskae]]''.<ref>Fostowicz-Frelik, Ł. 2010. A new species of Pliocene Prolagus (Lagomorpha, Ochotonidae) from Poland is the northernmost record of the genus. Journal of Vertebrate Paleontology 30 (2): 609-612.</ref>

Osmólska was, in recognition of her scientific work, a recipient of a number of awards including the [[Polish Cross of Merit]].<ref>Moody, R., & Geological Society of London. (2010). ''Dinosaurs and other extinct saurians: A historical perspective''. London: Geological Society.</ref>

==Selected publications==
* H. Osmólska and E. Roniewicz (1970). Deinocheiridae, a new family of theropod dinosaurs. ''Palaeontologica Polonica'' '''21''':5-19.
* H. Osmólska, E. Roniewicz, and R. Barsbold (1972). A new dinosaur, ''Gallimimus bullatus'' n. gen., n. sp. (Ornithomimidae) from the Upper Cretaceous of Mongolia. ''Palaeontologia Polonica'' '''27''':103-143.
* H. Osmólska (1972). Preliminary note on a crocodilian from the Upper Cretaceous of Mongolia. ''Palaeontologia Polonica'' '''27''':43-47.
* T. Maryańska and H. Osmólska (1974). Pachycephalosauria, a new suborder of ornithischian dinosaurs. ''Palaeontologia Polonica'' '''30''':45-102.
* T. Maryańska and H. Osmólska (1975). Protoceratopsidae (Dinosauria) of Asia. ''Palaeontologica Polonica'' '''33''':133-181.
* H. Osmólska (1976). New light on the skull anatomy and systematic position of ''Oviraptor''. ''Nature'' '''262''':683-684.
* H. Osmólska (1981). Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. ''Palaeontologica Polonica'' '''42''':79-95.
* T. Maryańska and H. Osmólska (1981). First lambeosaurine dinosaur from the Nemegt Formation, Upper Cretaceous, Mongolia. ''Acta Palaeontologica Polonica'' '''26'''(3-1):243-255.
* T. Maryańska and H. Osmólska (1981). Cranial anatomy of ''Saurolophus angustirostris'' with comments on the Asian Hadrosauridae (Dinosauria). ''Palaeontologia Polonica'' '''42''':5-24.
* H. Osmólska (1982). ''Hulsanpes perlei'' n.g. n.sp. (Deinonychosauria, Saurischia, Dinosauria) from the Upper Cretaceous Barun Goyot Formation of Mongolia. ''Neues Jahrbuch für Geologie und Paläontologie Monatshefte'' '''1982'''(7):440-448.
* A. Perle, T. Maryańska, and H. Osmólska (1982). ''Goyocephale lattimorei'' gen. et sp. n., a new flat-headed pachycephalosaur (Ornithischia, Dinosauria) from the Upper Cretaceous of Mongolia. ''Acta Palaeontologica Polonica'' '''27'''(1-4):115-127.
* T. Maryańska and H. Osmólska (1984). Postcranial anatomy of ''Saurolophus angustirostris'' with comments on other hadrosaurs. ''Palaeontologia Polonica'' '''46''':119-141.
* T. Maryańska and H. Osmólska (1985). On ornithischian phylogeny. ''Acta Palaeontologica Polonica'' '''30'''(3-4):137-149.
* H. Osmólska (1987). ''Borogovia gracilicrus'' gen. et sp. n., a new troodontid dinosaur from the Late Cretaceous of Mongolia. ''Acta Palaeontologica Polonica'' '''32'''(1-2):133-150.
* R. Barsbold, H. Osmólska, and S.M. Kurzanov (1987). On a new troodontid (Dinosauria, Theropoda) from the Early Cretaceous of Mongolia. ''Acta Palaeontologica Polonica'' '''32'''(1-2):121-132.
* S. M. Kurzanov and H. Osmólska (1991). ''Tochisaurus nemegtensis'' gen. et sp. n., a new troodontid dinosaur (Dinosauria, Theropoda) from Mongolia. ''Acta Palaeontologica Polonica'' '''36'''(1):69-76.
* H. Osmólska (1996). An unusual theropod dinosaur from the Late Cretaceous Nemegt Formation of Mongolia. ''Acta Palaeontologica Polonica'' '''41'''(1):1-38.
* R. Barsbold and H. Osmólska (1999). The skull of ''Velociraptor'' (Theropoda) from the Late Cretaceous of Mongolia. ''Acta Palaeontologica Polonica'' '''44'''(2):189-219.
* R. Barsbold, H. Osmólska, M. Watabe, P.J. Currie, and K. Tsogtbaatar (2000). A new oviraptorosaur (Dinosauria, Theropoda) from Mongolia: the first dinosaur with a pygostyle. ''Acta Palaeontologica Polonica'' '''45'''(2):97-106.
* T. Maryańska, H. Osmólska, and M. Wolsan (2002). Avialan status for Oviraptorosauria. ''Acta Palaeontologica Polonica'' '''47'''(1):97-116.
* H. Osmólska, P.J. Currie, and R. Barsbold (2004). Oviraptorosauria. In: D.B. Weishampel, P. Dodson, and H. Osmólska (eds.), ''The Dinosauria'' (second edition). University of California Press, Berkeley 165-183.

==References==
{{reflist}}

==Further reading==
*Borsuk-Białynicka, M. and Maryańska, T. "Halszka Osmólska (1930-2008) In Memoriam." ''Acta Palaeontologica Polonica'' 53, 2, 206.cp.
*Dodson, Peter (2008) "Polish Women in the Gobi – In Loving Memory of Halszka Osmólska (1930-2008)." ''American Paleontologist'' 16.3: 30

{{Authority control}}

{{DEFAULTSORT:Osmolska, Halszka}}
[[Category:Polish paleontologists]]
[[Category:Polish female geologists]]
[[Category:Women paleontologists]]
[[Category:1930 births]]
[[Category:2008 deaths]]
[[Category:Taxa named by Halszka Osmólska| ]]
[[Category:People from Poznań]]
[[Category:20th-century women scientists]]

Camptosaurus

2017-12-06T00:10:17Z

Wikkler: /* Classification */ It's six... not five.

:''Not to be confused with ''[[Camposaurus]]''.
{{Speciesbox
| fossil_range= {{Fossil range|Late Jurassic}}
| image = Camptosaurus dispar skeleton.jpg
| image_caption = Reconstructed skeleton in Japan
| taxon = Camptosaurus dispar
| greatgrandparent_authority = [[Othniel Charles Marsh|Marsh]], 1885
| grandparent_authority = Marsh, 1885
| parent_authority = Marsh, 1885
| authority = (Marsh, 1879 [originally ''Camptonotus''])
| display_parents = 4
| synonyms = {{collapsible list|bullets = true|title=List
|''Brachyrophus altarkansanus'' [[Edward Drinker Cope|Cope]], 1878
|''Symphyrophus musculosus'' Cope, 1878
|''Camptonotus dispar'' Marsh, 1879 ([[Camptonotus|preoccupied]])
|''Camptosaurus medius'' Marsh, 1894
|''Camptosaurus nanus'' Marsh, 1894
|''Camptosaurus browni'' [[Charles W. Gilmore|Gilmore]], 1909
}}}}

'''''Camptosaurus''''' ({{IPAc-en|ˌ|k|æ|m|p|t|oʊ-|ˈ|s|ɔːr|ə|s}} {{respell|KAMP|to|SAWR|əs}}) is a [[genus]] of plant-eating, beaked [[ornithischian]] [[dinosaur]]s of the [[Late Jurassic]] [[Period (geology)|period]] of western [[North America]] and possibly Europe. The name means 'flexible lizard' ([[Ancient Greek|Greek]] καμπτος/''kamptos'' meaning 'bent' and σαυρος/''sauros'' meaning 'lizard').

==Description==
[[File:Camptosaurus.jpg|left|thumb|Restoration]]
''Camptosaurus'' is a relatively heavily built form, with robust hindlimbs and broad feet, still having four toes.<ref name="GaltonPowell"/> Due to the separate status of ''Uteodon'' it has become problematic which material from the Morrison belongs to ''Camptosaurus''. The specimens with certainty belonging to ''Camptosaurus dispar'', from Quarry 13, have been recovered from very deep layers, probably dating to the [[Callovian]]-[[Oxfordian (stage)|Oxfordian]].<ref name=CarpenterWilson/> The largest fragments from later strata indicate adult individuals more than {{convert|7.9|m|ft}} long, and {{convert|2|m|ft}} at the hips.<ref name=BRE03>{{cite book |last=Erickson |first=Bruce R. |authorlink=Bruce Erickson (paleontologist)|title=Dinosaurs of the Science Museum of Minnesota |year=2003 |publisher=The Science Museum of Minnesota |location=St. Paul, Minnesota |isbn=978-0-911338-54-6 |page=33 }}</ref> The Quarry 13 individuals are smaller though. They have been described as reaching 6 meters (19.7 feet) in length and 785 – 874 kg in weight.<ref name="foster-Camptosaurus">Foster, J. (2007). "''Camptosaurus dispar''." ''Jurassic West: The Dinosaurs of the Morrison Formation and Their World''. Indiana University Press. p. 219-221.</ref> In 2010 [[Gregory S. Paul]] gave an even lower estimate: a length of five metres and a weight of half a tonne.<ref>Paul, G.S., 2010, ''The Princeton Field Guide to Dinosaurs'', Princeton University Press p. 284</ref>

Earlier reconstructions, such as those by Marsh and Gilmore, were based on the skull of ''Theiophytalia'' and display an incorrect, more rectangular profile. The skull was in fact triangular with a pointed snout, equipped with a beak. Its teeth were more tightly packed in the jaw compared to other Morrison euornithopods.<ref name="foster-Camptosaurus"/> Museum curator John Foster describes them as having "thick median ridges on their lateral sides and denticles along their edges," these features were similar to, but "more fully developed" than those in ''[[Dryosaurus]]''.<ref name="foster-Camptosaurus"/> ''Camptosaurus'' teeth frequently exhibit extensive wear, which indicates that individuals in the genus had a diet of relatively tough vegetation.<ref name="foster-Camptosaurus"/>

==History of discovery==
[[File:Marsh Camptosaurus.jpg|thumb|left|Historical skeletal restoration by O.C. Marsh, with skull based on remains now referred to ''Theiophytalia'']]
On September 4, 1879 [[William Harlow Reed]] in [[Albany County, Wyoming]] found the remains of a small [[Euornithopoda|euornithopod]]. That same year Professor [[Othniel Charles Marsh]] described and named the find as ''[[Camptonotus]]'', or "flexible back", from Greek κάμπτω, "to bend" and νῶτον, "back", in reference to the presumed flexibility of the [[sacral vertebrae]]. The [[holotype]] was YPM 1877, a partial skeleton.<ref>{{cite journal | last1 = Marsh | first1 = O.C. | year = 1879 | title = Notice of new Jurassic reptiles | url = | journal = American Journal of Science and Arts | volume = 18 | issue = | pages = 501–505 }}</ref> The genus was renamed ''Camptosaurus'' by him in 1885 because the original name was already in use for a [[cricket (insect)|cricket]].<ref>{{cite journal | last1 = Marsh | first1 = O.C. | year = 1885 | title = Names of extinct reptiles | url = | journal = American Journal of Science | volume = 29 | issue = | page = 169 }}</ref> In 1879, Marsh named ''C. dispar'' ([[type species]] of the genus) for material he received from his collectors at ''Quarry 13'' near [[Como Bluff]], [[Wyoming]] in the [[Morrison Formation]] and ''C. amplus'' based on the holotype YPM 1879, a foot found by [[Arthur Lakes]] at ''Quarry 1A''. The foot was later shown to have belonged to ''[[Allosaurus]]''.<ref name="GaltonPowell">{{Cite journal|author1=Galton, P.M. |author2=Powell, H.P. |year=1980 |title=The ornithischian dinosaur Camptosaurus prestwichii from the Upper Jurassic of England |journal=Palaeontology |volume=23 |issue= |pages=411–443 |doi= }}</ref><ref name="Bakker">{{cite book |author=Bakker, R.T. |year=1998 |chapter=Dinosaur mid-life crisis: the Jurassic-Cretaceous transition in Wyoming and Colorado |editor=Lucas, S.G. |editor2=Kirkland, J.I. |editor3=Estep, J.W. |title=Lower and Middle Cretaceous Terrestrial Ecosystems |publisher=New Mexico Museum of Natural History and Science Bulletin |volume=14 |pages=67–77}}</ref> Throughout the 1880 and 1890s, he continued to receive specimens from Quarry 13 and in 1894 named two additional species: ''C. medius'' and ''C. nanus'', based in part on size.<ref>{{cite journal | last1 = Marsh | first1 = O.C. | year = 1894 | title = The typical Ornithopoda of the American Jurassic | url = | journal = American Journal of Science |series =3 | volume = 48 | issue = | pages = 85–90 }}</ref> [[Charles W. Gilmore]] named two additional species, ''C. browni'' and ''C. depressus'' in his 1909<ref name=Gilmore09>{{Cite journal|author=Gilmore, C.W. |year=1909 |title=Osteology of the Jurassic reptile ''Camptosaurus'', with a revision of the species of the genus, and descriptions of two new species |journal=Proceedings of the United States National Museum |volume=36 |pages=197–332 |doi=10.5479/si.00963801.36-1666.197}}</ref> redescription of the Marsh specimens. In the Morrison Formation, ''Camptosaurus'' fossils are present in stratigraphic zones 2-6.<ref name="foster-appendix">Foster, J. (2007). "Appendix." ''Jurassic West: The Dinosaurs of the Morrison Formation and Their World''. Indiana University Press. pp. 327-329.</ref>
[[File:Camptosaurus nanum - AMNH - DSC06302.JPG|thumb|Outdated mount of a ''C. nanus'' skeleton at the AMNH, now thought to be a growth stage of ''C. dispar'']]
Then in 1980, [[Peter Galton]] and H.P. Powell in their redescription of ''C. prestwichi'' (see following), considered ''C. nanus'', ''C. medius'' and ''C. browni'' to be different growth stages or different gender of the larger ''C. dispar'', and therefore only ''C. dispar'' was a valid species.<ref name="GaltonPowell"/> They also considered a skull, YPM 1887, in 1886 referred to ''C. amplus'' by Marsh, later confirmed by Gilmore, to belong to ''C. dispar'' as well. Gilmore had used this skull to describe the skull of ''Camptosaurus'', but the specimen was recently shown by Brill and Carpenter not to belong to ''Camptosaurus''. In 2007, they put it into its own genus and species, ''[[Theiophytalia kerri]]''.<ref name=BrillCarpenter>{{cite book |author1=Brill, K. |author2=K. Carpenter |year=2007 |chapter=A description of a new ornithopod from the Lytle Member of the Purgatoire Formation (Lower Cretaceous) and a reassessment of the skull of ''Camptosaurus'' |editor=Carpenter, Kenneth |title=Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs |publisher=Indiana University Press |location=Bloomington and Indianapolis |pages=49–67 |isbn=0-253-34817-X}}</ref>
[[File:9162 - Milano, Museo storia naturale - Camptosaurus dispar - Foto Giovanni Dall'Orto 22-Apr-2007.jpg|thumb|left|''Camptosaurus'' skeleton mounted in outdated quadrupedal posture, and with ''[[Theiophytalia]]'' skull, [[Natural History Museum of Milan]]]]
''Camptosaurus depressus'' was recovered from the [[Lakota Formation]] near the town of [[Hot Springs, South Dakota]]. It was described by [[Charles Gilmore]] in 1909 based on the holotype and only known specimen USNM 4753, a fragmentary postcranium, by the "narrowness or depressed nature of the ilia".<ref name=Gilmore09/> Carpenter and Wilson (2008) referred this species to ''[[Planicoxa]]'', as ''P. depressa'', on the basis of similarities between its ilium and the holotype ilium of ''Planicoxa venenica''.<ref name=CarpenterWilson>{{cite journal |author1=Carpenter, K. |author2=Wilson, Y. |year=2008 |title=A new species of ''Camptosaurus'' (Ornithopoda: Dinosauria) from the Morrison Formation (Upper Jurassic) of Dinosaur National Moument, Utah, and a biomechanical analysis of its forelimb |journal=Annals of the Carnegie Museum |volume=76 |issue= |pages=227–263 |doi=10.2992/0097-4463(2008)76[227:ansoco]2.0.co;2}}</ref> However, McDonald and colleagues (2010), and McDonald (2011) found that the horizontal postacetabular process of ''C. depressus'' is more likely a product of distortion.<ref name=iguanacolossus>{{cite journal | last1 = McDonald | first1 = A.T. | last2 = Kirkland | first2 = J.I. | last3 = DeBlieux | first3 = D.D. | last4 = Madsen | first4 = S.K. | last5 = Cavin | first5 = J. | last6 = Milner | first6 = A.R.C. | last7 = Panzarin | first7 = L. | year = 2010 | title = New Basal Iguanodonts from the Cedar Mountain Formation of Utah and the Evolution of Thumb-Spiked Dinosaurs | url = | journal = PLoS ONE | volume = 5| issue = 11| page = e14075 | doi = 10.1371/journal.pone.0014075 | pmid = 21124919 | pmc=2989904}}</ref> Therefore, McDonald put it into its own genus, ''[[Osmakasaurus]]''.<ref name="McDonald">{{Cite journal|author=Andrew T. McDonald |year=2011 |title=The taxonomy of species assigned to ''Camptosaurus'' (Dinosauria: Ornithopoda) |url=http://www.mapress.com/zootaxa/2011/f/z02783p068f.pdf |journal=Zootaxa |volume=2783 |issue= |pages=52–68 |doi= }}</ref> An additional species, ''Camptosaurus aphanoecetes'', was named by Carpenter and Wilson in 2008 for specimens from [[Dinosaur National Monument]].<ref name=CarpenterWilson/> It differs from ''C. dispar'' in the lower jaw, shorter neck vertebrae, and straighter [[ischium]] ending in a small "foot" among other features. An analysis by Andrew McDonald and colleagues in 2010 suggested that like ''C. aphanoecetes'' is actually more closely related to more advanced iguanodonts ([[Styracosterna]]).<ref name=iguanacolossus/> It has been moved to the new genus ''[[Uteodon]]''.<ref name="McDonald"/>
[[File:Camptosaurus sp salt lake city.jpg|thumb|Cast of a skull from [[Bone Cabin Quarry]] West, Wyoming]]
While Marsh was describing ''Camptosaurus'' species in North America, numerous species from Europe were also referred to the genus in the late 19th and early 20th centuries: ''C. inkeyi'', ''C. hoggii'', ''C. leedsi'', ''C. prestwichi'', and ''C. valdensis''. ''C. inkeyi'' (Nopcsa, 1900) consists of fragmentary material, a dentary and articular from Upper Cretaceous rocks of the [[Haţeg Basin]] in [[Romania]]. It is almost certainly a rhabdodontid and is no longer considered valid (''[[nomen dubium]]'').<ref name="SachsHornung">{{Cite journal|author1=Sachs, S. |author2=Hornung, J.J. |year=2006 |title=Juvenile ornithopod (Dinosauria: Rhabdodontidae) remains from the Upper Cretaceous (Lower Campanian, Gosau Group) of Muthmannsdorf (Lower Austria) |journal=Geobios |volume=39 |issue= |pages=415–425 |doi= 10.1016/j.geobios.2005.01.003}}</ref> ''C. valdensis'' is a [[nomen dubium|dubious]] dryosaurid,<ref name=PMG09/> based on the holotype and only known specimen NHMUK R167, a poorly preserved left femur lacking the distal end. It is thus difficult to compare it to other dryosaurids, including the contemporary ''[[Valdosaurus|Valdosaurus canaliculatus]]''.<ref name="McDonald"/> ''C. leedsi'' is probably a valid dryosaurid that has been moved to the new genus ''[[Callovosaurus]]''.<ref name=JROetal07>{{cite book |last1=Ruiz-Omeñaca |first1=José Ignacio |last2=Pereda Suberbiola |first2=Xabier |last3=Galton |first3=Peter M. |year=2007 |chapter=''Callovosaurus leedsi'', the earliest dryosaurid dinosaur (Ornithischia: Euornithopoda) from the Middle Jurassic of England |editor-last=Carpenter |editor-first=Kenneth |title=Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs |publisher=Indiana University Press |location=Bloomington and Indianapolis |pages=3–16 |isbn=0-253-34817-X}}</ref> ''C. hoggii'' was originally named ''[[Iguanodon]] hoggii'' by [[Richard Owen]] in 1874 and was moved to ''Camptosaurus'' by Norman and Barrett in 2002.<ref>Norman D.B. and Barrett, P.M. 2002. Ornithischian dinosaurs from the Lower Cretaceous (Berriasian) of England. Palaeontology 68:161-189</ref> It has since been transferred to the genus ''[[Owenodon]]''.<ref name=PMG09>{{cite journal |last=Galton |first=Peter M. |authorlink=Peter Galton |year=2009 |title=Notes on Neocomian (Late Cretaceous) ornithopod dinosaurs from England - ''Hypsilophodon'', ''Valdosaurus'', "Camptosaurus", "Iguanodon" – and referred specimens from Romania and elsewhere |journal=Revue de Paléobiologie |volume=28 |issue=1 |pages=211–273}}</ref>

The remaining European species ''Camptosaurus prestwichii'' was recovered from Chawley Brick Pits, [[Cumnor Hurst]] in [[Oxfordshire]] in England. The fossil was found when a [[tramway (industrial)|tramway]] was driven into the side of a hill. It was described by Hulke in 1880 as ''Iguanodon prestwichii'', and then placed in its own genus ''[[Cumnoria]]'' by Seeley in 1888, but was soon sunk into ''Camptosaurus'' by Lydekker in 1889.<ref>{{cite book|title=Fossil Reptiles of Great Britain|year=1995|vauthors=Benton MJ, Spencer PS |publisher=Chapman & Hall|isbn=0-412-62040-5}}</ref> However, Naish & Martill (2008), McDonald and colleagues (2010), and McDonald (2011) found that Seeley's original generic distinction was valid. ''Cumnoria'' has been recovered as a [[styracosterna]]n, more closely related to advanced iguanodonts than to ''Camptosaurus dispar'', similar to the case of ''Uteodon''.<ref name=iguanacolossus/><ref name="McDonald"/>

==Classification==
Marsh in 1885 assigned ''Camptosaurus'' to a family of its own: the Camptosauridae. Alternatively some authors considered it an early member of the [[Iguanodontidae]].

Modern [[phylogenetics]] has made ''Camptosaurus'' by definition part of the [[clade]] [[Ankylopollexia]], of which group it would then be a basal member.
This would mean that the genus is closely related to the ancestor of later [[iguanodontid]] and [[hadrosaurid]] dinosaurs and was more [[Synapomorphy|derived]] than contemporaries such as ''[[Dryosaurus]]'', ''[[Drinker (dinosaur)|Drinker]]'', and ''[[Othnielosaurus]]''.<ref name="foster-Camptosaurus"/>

In the 2010 and 2011 [[cladistic]] analyses of McDonald and colleagues, ''Camptosaurus'' was placed as follows:<ref name="McDonald">{{Cite journal|author=Andrew T. McDonald |year=2011 |title=The taxonomy of species assigned to ''Camptosaurus'' (Dinosauria: Ornithopoda) |url=http://www.mapress.com/zootaxa/2011/f/z02783p068f.pdf |journal=Zootaxa |volume=2783 |issue= |pages=52–68 |doi= }}</ref><ref name=hippoiguana>{{Cite journal|author=McDonald, A.T.|author2=Kirkland, J.I.|author3=DeBlieux, D.D.|author4=Madsen, S.K.|author5=Cavin, J.|author6=Milner, A.R.C.|author7=Panzarin, L.|last-author-amp=yes |year=2010 |title=New Basal Iguanodonts from the Cedar Mountain Formation of Utah and the Evolution of Thumb-Spiked Dinosaurs |journal=PLoS ONE |volume=5 |issue=11 |pages=e14075 |doi=10.1371/journal.pone.0014075 |pmid=21124919 |pmc=2989904}}</ref>
[[File:Ornithopods jconway.jpg|thumb|Restoration of six [[ornithopod]] dinosaurs, including ''Camptosaurus'' at the far left]]
{{clade| style=font-size:90%; line-height:90%
|label1=[[Ankylopollexia]]
|1={{clade
|1='''''Camptosaurus'''''
|label2=[[Styracosterna]]
|2={{clade
|1=''[[Uteodon]]''
|2={{clade
|1={{clade
|1=''[[Hippodraco]]''
|2=''[[Theiophytalia]]''}}
|2={{clade
|1=''[[Iguanacolossus]]''
|2={{clade
|1=''[[Lanzhousaurus]]''
|2={{clade
|1=''[[Kukufeldia]]''
|2={{clade
|1=''[[Barilium]]''
|2=[[Hadrosauriformes]] }} }} }} }} }} }} }} }}

==Paleobiology==
[[File:Camptoscale.svg|thumb|Size compared to a human]]
Based on studies of other iguanodonts (clade [[Iguanodontia]]), scientists believe they may have been able to achieve running speeds of 25 km per hour (15 mph).<ref name="foster-Camptosaurus"/> A tiny 9 inch fossilized embryo, referred to ''Camptosaurus'', was retrieved from Morrison Formation strata at Dinosaur National Monument in Utah.<ref name="foster-Camptosaurus"/>

==See also==
* [[List of dinosaurs]]
* [[Paleobiota of the Morrison Formation]]

==References==
{{Portal|Dinosaurs}}
{{Reflist}}
{{commons category}}

[[Category:Iguanodonts]]
[[Category:Late Jurassic dinosaurs of North America]]
[[Category:Dinosaurs of the Morrison Formation]]
[[Category:Fossil taxa described in 1879]]
[[Category:Fossil taxa described in 1885]]
[[Category:Taxa named by Othniel Charles Marsh]]
[[Category:Paleontology in Wyoming]]

Ornithischia

2017-12-04T00:26:57Z

Wikkler: Replaced a generic "Archosauromorpha" template with no links to Ornithischia or any ornithischians with the "Ornithodira" template, which does.

{{Automatic taxobox
| name = Ornithischians
| fossil_range = [[Early Jurassic]]–[[Late Cretaceous]], {{fossilrange|200|66|earliest=228}} (Possible Late Triassic record)
| image = 조반류.png
| image_caption = Diversity of ornithischian dinosaurs.
| authority = [[Harry Govier Seeley|Seeley]], 1888
| subdivision_ranks = Subgroups
| subdivision =
* {{extinct}}''[[Chilesaurus]]''?
* {{extinct}}''[[Eocursor]]''
* {{extinct}}''[[Fabrosaurus]]''
* {{extinct}}''[[Taveirosaurus]]''
* {{extinct}}''[[Trimucrodon]]''
* {{extinct}}'''[[Heterodontosauridae]]'''
* {{extinct}}'''[[Genasauria]]'''
| synonyms =
* Predentata<ref name=ferigolo&langer2006>{{Cite journal | doi = 10.1080/08912960600845767| title = A Late Triassic dinosauriform from south Brazil and the origin of the ornithischian predentary bone| journal = Historical Biology| volume = 19| pages = 23–33| year = 2007| last1 = Ferigolo | first1 = J. | last2 = Langer | first2 = M. C. }}</ref> [[Othniel Charles Marsh|Marsh]], 1894
}}

'''Ornithischia''' ({{IPAc-en|ɔːr|n|ᵻ|ˈ|θ|ɪ|s|k|i|ə}} {{respell|or|ni|THISS|kee|ə}}) is an extinct [[clade]] of mainly [[Herbivore|herbivorous]] [[dinosaur]]s<nowiki/> characterized by a pelvic structure similar to that of [[bird]]s.<ref name=":0">{{Cite book|title=Dinosaurs: A Concise Natural History|last=Fastovsky|first=David E.|last2=Weishampel|first2=David B.|publisher=Cambridge University Press|year=2012|isbn=1107276462|location=Cambridge|pages=}}</ref> The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the [[Ancient Greek|Greek]] stem ''{{transl|el|ornith-}}'' ({{lang|el|ὀρνιθ-}}), meaning "of a bird", and ''{{transl|el|ischion}}'' ({{lang|el|ἴσχιον}}), plural ''{{transl|el|ischia}}'', meaning "hip joint". Birds, are however, unrelated, instead being [[theropod]] dinosaurs.<ref name=":0" />

Ornithischians with well known anatomical adaptations include the [[ceratopsian]]s or "horn-faced" dinosaurs (e.g. ''[[Triceratops]]''), armored dinosaurs ([[Thyreophora]]) such as [[stegosaurs]] and [[ankylosaurs]], [[Pachycephalosauria|pachycephalosaurids]] and the [[ornithopod]]s.<ref name=":0" /> There is strong evidence that certain groups of ornithischians lived in herds,<ref name=":0" /><ref name=":1">{{Cite journal|last=Qi|first=Zhao|last2=Barrett|first2=Paul M.|last3=Eberth|first3=David A.|date=2007-09-01|title=Social Behaviour and Mass Mortality in the Basal Ceratopsian Dinosaur Psittacosaurus (early Cretaceous, People's Republic of China)|url=http://onlinelibrary.wiley.com/doi/10.1111/j.1475-4983.2007.00709.x/abstract|journal=Palaeontology|language=en|volume=50|issue=5|pages=1023–1029|doi=10.1111/j.1475-4983.2007.00709.x|issn=1475-4983}}</ref> often segregated by age group, with juveniles forming their own flocks separate from adults.<ref name="Nest">{{Cite journal | last1 = Zhao | first1 = Q. | doi = 10.4202/app.2012.0128 | title = Juvenile-only clusters and behaviour of the Early Cretaceous dinosaur ''Psittacosaurus'' | journal = Acta Palaeontologica Polonica | year = 2013 | pmid = | pmc = }}</ref> Some were at least partially covered in filamentous (hair- or feather- like) pelts, and there is much debate over whether these filaments found in specimens of ''[[Tianyulong]]'',<ref name="ButlerEtAl" /> ''[[Psittacosaurus]]'',<ref>{{Cite journal|last=Mayr|first=Gerald|last2=Peters|first2=Stefan D.|last3=Plodowski|first3=Gerhard|last4=Vogel|first4=Olaf|date=2002-08-01|title=Bristle-like integumentary structures at the tail of the horned dinosaur Psittacosaurus|url=https://link.springer.com/article/10.1007/s00114-002-0339-6|journal=Naturwissenschaften|language=en|volume=89|issue=8|pages=361–365|doi=10.1007/s00114-002-0339-6|issn=0028-1042|pmid=12435037}}</ref> and ''[[Kulindadromeus]]'' may have been primitive [[feather]]s.<ref name="Godefroit2014">{{cite journal | last1 = Godefroit | first1 = P. | last2 = Sinitsa | first2 = S.M. | last3 = Dhouailly | first3 = D. | last4 = Bolotsky | first4 = Y.L. | last5 = Sizov | first5 = A.V. | last6 = McNamara | first6 = M.E. | last7 = Benton | first7 = M.J. | last8 = Spagna | first8 = P. | year = 2014 | title = A Jurassic ornithischian dinosaur from Siberia with both feathers and scales | url = http://palaeo.gly.bris.ac.uk/Benton/reprints/2014Kulinda.pdf | format = PDF | journal = Science | volume = 345 | issue = 6195| pages = 451–455 | doi = 10.1126/science.1253351 }}</ref>

Currently, the exact placement of Ornithischia within the dinosaur lineage is a contentious issue. Traditionally, the group is considered sister to [[Saurischia]] (which contains Theropoda and [[Sauropodomorpha]]).<ref name="seeley1888">{{cite journal | last1 = Seeley | first1 = H.G. | year = 1888 | title = On the classification of the fossil animals commonly named Dinosauria | url = | journal = Proceedings of the Royal Society of London | volume = 43 | issue = | pages = 165–171 | doi=10.1098/rspl.1887.0117}}</ref> In the alternative evolutionary hypothesis for dinosaurs that was proposed by Baron, Norman & Barrett in the journal ''[[Nature (journal)|Nature]]'' in 2017, Ornithischia as well as Theropoda were grouped together in the clade [[Ornithoscelida]].<ref>{{cite journal | last1 = Baron | first1 = M.G. | last2 = Norman | first2 = D.B. | last3 = Barrett | first3 = P.M. | year = 2017 | title = A new hypothesis of dinosaur relationships and early dinosaur evolution | url = | journal = Nature | volume = 543 | issue = | pages = 501–506 | doi = 10.1038/nature21700 }}</ref> The work by Baron and colleagues was recently challenged by an international consortium of early dinosaur experts led by Max Langer. However, the data that supported the more traditional placement of Ornithischia, as sister-taxon of Saurischia, was found not to be statistically significant from the evidence that supported the Ornithoscelida hypothesis, in both the study by Langer et al. and the reply to the study by Baron et al..<ref>{{Cite journal|author1=Max C. Langer |author2=Martín D. Ezcurra |author3=Oliver W. M. Rauhut |author4=Michael J. Benton |author5=Fabien Knoll |author6=Blair W. McPhee |author7=Fernando E. Novas |author8=Diego Pol |author9=Stephen L. Brusatte |year=2017 |title=Untangling the dinosaur family tree |journal=Nature |volume=551 |issue=7678 |pages=E1–E3 |doi=10.1038/nature24011 }}</ref><ref>{{Cite journal|author1=Matthew G. Baron |author2=David B. Norman |author3=Paul M. Barrett |year=2017 |title=Baron et al. reply |journal=Nature |volume=551 |issue=7678 |pages=E4–E5 |doi=10.1038/nature24012 }}</ref> A further 2017 study found some support for the previously abandoned [[Phytodinosauria]] model, which classifies ornithischians together with sauropodomorphs.<ref name=LP17>{{Cite journal|author1=Luke A. Parry |author2=Matthew G. Baron |author3=Jakob Vinther |year=2017 |title=Multiple optimality criteria support Ornithoscelida |journal=Royal Society Open Science |volume=4 |issue=10 |pages=170833 |doi=10.1098/rsos.170833 }}</ref>

== Description ==
In 1887, [[Harry Seeley]] divided Dinosauria into two clades: Ornithischia and Saurischia. Ornithischia is a strongly supported clade with an abundance of diagnostic characters (common traits).<ref name=":0" /> The two most notable traits are a "bird-like" hip and beak-like predentary structure though they shared other features as well.<ref name=":0" />

=== "Bird-hip" ===
The ornithischian pelvis is "opisthopubic", meaning that the pubis points down and back ([[Anterior|posterior]]) parallel with the ischium (Figure 1a).<ref name=":0" /> Additionally, the pelvis has a forward-pointing process to support the abdomen.<ref name=":0" /> This results in a four-pronged pelvic structure. In contrast to this, the saurischian pelvis is "propubic", meaning the pubis points toward the head ([[anterior]]), as in ancestral reptiles (Figure 1b).<ref name=":0" />

The opisthopubic pelvis independently evolved at least three times in dinosaurs (in ornithischians, birds and [[therizinosaur]]oids).<ref name=":3">{{Cite book|url=https://books.google.com/books?id=7t9M5TsmjOUC|title=Encyclopedia of Dinosaurs|last=Currie|first=Philip J.|last2=Padian|first2=Kevin|date=1997-10-06|publisher=Academic Press|isbn=9780080494746|pages=537–538|language=en}}</ref> Some argue that the opisthopubic pelvis evolved a fourth time in the clade [[Dromaeosauridae]], but this is controversial, as other authors argue that dromeosaurids are mesopubic.<ref name=":3" />

=== Predentary ===
Ornithischians share a unique bone called the [[predentary]] (Figure 2).<ref name=":0" /> This unpaired bone is found at the front of the lower jaws and presumably aided ornithischians in cropping vegetation. In 2017 Baron & Barrett suggested that ''[[Chilesaurus]]'' may represent an early diverging ornithischian that had not yet acquired the predentary of all other ornithischians.<ref>{{cite journal|last1=Baron |first1=Matthew G. |last2=Barrett |first2=Paul M. |title=A dinosaur missing-link? ''Chilesaurus'' and the early evolution of ornithischian dinosaurs |volume=13 |number=8 |year=2017 |doi=10.1098/rsbl.2017.0220 |publisher=[[The Royal Society]] |url=http://rsbl.royalsocietypublishing.org/content/13/8/20170220 |journal=[[Biology Letters]]}}</ref>

{|
![[File:Ornithischia pelvis structure.svg|thumb|left|Figure 1a - Ornithischian propisthopubic [[pelvis|pelvic]] structure (left side)|276x276px]]
![[File:Saurischia pelvis structure.svg|left|thumb|245x245px|Figure 1b - Saurischian propubic pelvic structure (left side)]]
![[File:Heterodontosaurus skull reconstruction sereno 2012 edited.jpg|thumb|440x440px|Figure 2 - Heterodontosaurus skull with palpebral bone (red), antorbital fenestra (yellow) and predentary (green) colored.]]
|}

=== Other characteristics ===

* Ornithischians had paired [[premaxilla]]ry bones that were toothless and roughened at the tip of the snout (presumably due to the attachment of a keratinous beak).<ref name=":0" />
* Ornithischians developed a narrow "eyebrow", or palpebral bone, across the outside of the eye socket.<ref name=":0" />
* Ornithischians had reduced, or even closed-off, [[antorbital fenestra]]e (the fenestra in front of the eye socket).<ref name=":0" />
* Ornithischian jaw joints were lowered below the level of the teeth, bringing the teeth into simultaneous occlusion.<ref name=":0" />
* Ornithischians had "leaf-shaped" cheek teeth.<ref name=":0" />
* Ornithischian backbones were stiffened near the pelvis by the ossification of tendons above the sacrum. Additionally, ornithischians had at least five sacral vertebrae attaching to the pelvis.<ref name=":0" />

==Classification==
Ornithischia is a branch-based taxon defined as all dinosaurs more closely related to ''[[Triceratops horridus]]'' Marsh, 1889 than to either ''[[Passer domesticus]]'' (Linnaeus, 1758) or ''[[Saltasaurus loricatus]]'' Bonaparte & Powell, 1980.<ref name="Butleretal2008">{{cite journal|last=Butler|first=Richard |author2=Upchurch, Paul |author3=Norman, David |date=2008|title=The phylogeny of ornithischian dinosaurs|journal=[[Journal of Systematic Palaeontology]]|volume=6|issue=1|pages=1–40|doi=10.1017/S1477201907002271}}</ref> [[Genasauria]] comprises the [[clades]] [[Thyreophora]] and [[Neornithischia]]. Thyreophora includes [[Stegosauria]] (like the armored ''[[Stegosaurus]]'') and [[Ankylosauria]] (like ''[[Ankylosaurus]]''). Neornithischia comprises several basal taxa, [[Marginocephalia]] ([[Ceratopsia]] and [[Pachycephalosauria]]), and [[Ornithopoda]] (including [[hadrosaur|duck-bills]] (hadrosaurs), such as ''[[Edmontosaurus]])''. Cerapoda is a relatively recent concept (Sereno, 1986).

The [[cladogram]] below follows a 2009 analysis by Zheng and colleagues. All tested members of [[Heterodontosauridae]] form a [[polytomy]].<ref name="zheng2009">{{cite journal|last=Zheng|first=Xiao-Ting|author2=You, Hai-Lu |author3=Xu, Xing |author4= Dong, Zhi-Ming |date=19 March 2009|title=An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures|journal=[[Nature (journal)|Nature]]|volume=458|issue=7236|pages=333–336|doi=10.1038/nature07856|pmid=19295609}}</ref>

{{clade| style=font-size:85%;line-height:80%
|label1='''Ornithischia'''
|1={{clade
|1=''[[Pisanosaurus]]'' [[File:Pisanosaurus.jpg|50px]]
|2={{clade
|1=[[Heterodontosauridae]] [[File:Fruitadens.jpg|50px]]
|label2=[[Genasauria]]
|2={{clade
|label1=[[Thyreophora]]
|1={{clade
|1=''[[Lesothosaurus]]''
|2={{clade
|1=''[[Scutellosaurus]]'' [[File:Scutellosaurus.jpg|50px]]
|2={{clade
|1=''[[Emausaurus]]''
|2={{clade
|1=''[[Scelidosaurus]]'' [[File:Scelidosaurus harrisonii.png|50px]]
|2={{clade
|1='''[[Stegosauria]]''' [[File:Stegosaurus BW.jpg|50px]]
|2='''[[Ankylosauria]]''' [[File:Edmontonia dinosaur.png|50px]] }} }} }} }} }}
|label2=[[Neornithischia]]
|2={{clade
|1=''[[Stormbergia]]''
|2={{clade
|1=''[[Agilisaurus]]'' [[File:Agilisaurus2.jpg|50px]]
|2={{clade
|1=''[[Hexinlusaurus]]''
|label2=[[Cerapoda]]
|2={{clade
|1=''[[Othnielia]]''
|2=''[[Hypsilophodon]]'' [[File:Hypsilophodon.jpg|50px]]
|3=''[[Jeholosaurus]]''
|4=''[[Yandusaurus]]''
|5={{clade
|1=''[[Orodromeus]]'' [[File:Orodromeus (pencil 2013).png|50px]]
|2=''[[Zephyrosaurus]]'' }}
|6='''[[Ornithopoda]]''' [[File:Parasaurolophuspic steveoc.jpg|50px]]
|label7=[[Marginocephalia]]
|7={{clade
|1='''[[Pachycephalosauria]]''' [[File:Pachycephalosauria jmallon.jpg|50px]]
|2='''[[Ceratopsia]]''' [[File:Psittacosaurus mongoliensis whole BW.jpg|50px]][[File:Triceratops BW.jpg|50px]] }} }} }} }} }} }} }} }} }}

Cladogram after Butler ''et al.'', 2011. [[Ornithopoda]] includes ''[[Hypsilophodon]]'', ''[[Jeholosaurus]]'' and others.<ref name=ButlerEtAl>{{Cite journal | author = Richard J. Butler, Jin Liyong, Chen Jun, [[Pascal Godefroit]] | date = May 2011 | title = The postcranial osteology and phylogenetic position of the small ornithischian dinosaur ''Changchunsaurus parvus'' from the Quantou Formation (Cretaceous: Aptian–Cenomanian) of Jilin Province, north-eastern China | journal = Palaeontology | volume = 54 | issue = 3 | pages = 667–683 | doi = 10.1111/j.1475-4983.2011.01046.x }}</ref>

{{clade| style=font-size:85%;line-height:80%
|label1='''Ornithischia'''
|1={{clade
|1=''[[Pisanosaurus]]'' [[File:Pisanosaurus.jpg|50px]]
|2={{clade
|1=[[Heterodontosauridae]] [[File:Fruitadens.jpg|50px]]
|2={{clade
|1=''[[Eocursor]]''
|label2=[[Genasauria]]
|2={{clade
|1=''[[Lesothosaurus]]''
|label2=[[Thyreophora]]
|2={{clade
|1=''[[Scutellosaurus]]'' [[File:Scutellosaurus.jpg|50px]]
|2={{clade
|1=''[[Emausaurus]]''
|2={{clade
|1=''[[Scelidosaurus]]'' [[File:Scelidosaurus harrisonii.png|50px]]
|2={{clade
|1='''[[Stegosauria]]''' [[File:Stegosaurus BW.jpg|50px]]
|2='''[[Ankylosauria]]''' [[File:Edmontonia dinosaur.png|50px]] }} }} }} }}
|label3=[[Neornithischia]]
|3={{clade
|1=''[[Stormbergia]]''
|2={{clade
|1=''[[Agilisaurus]]'' [[File:Agilisaurus2.jpg|50px]]
|2={{clade
|1=''[[Hexinlusaurus]]''
|2={{clade
|1=''[[Othnielosaurus]]''
|label2=[[Cerapoda]]
|2={{clade
|1='''[[Ornithopoda]]''' [[File:Parasaurolophuspic steveoc.jpg|50px]]
|label2=[[Marginocephalia]]
|2={{clade
|1='''[[Pachycephalosauria]]''' [[File:Pachycephalosauria jmallon.jpg|50px]]
|2='''[[Ceratopsia]]''' [[File:Psittacosaurus mongoliensis whole BW.jpg|50px]][[File:Triceratops BW.jpg|50px]] }} }} }} }} }} }} }} }} }} }} }}

== Palaeoecology ==
Ornithischians shifted from bipedal to quadrupedal posture at least three times in their evolutionary history and it has been shown primitive members may have been capable of both forms of movement.<ref>{{cite journal|url=http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0007331|title=Dynamic Locomotor Capabilities Revealed by Early Dinosaur Trackmakers from Southern Africa|author=Jeffrey A. Wilson|author2=Claudia A. Marsicano|author3=Roger M. H. Smith|date=6 October 2009|publisher=PLOS ONE|doi=10.1371/journal.pone.0007331|volume=4|journal=PLoS ONE|pages=e7331}}</ref>

Most ornithischians were herbivorous.<ref name=":0" /> In fact, most of the unifying characters of Ornithischia are thought to be related to this herbivory.<ref name=":0" /> For example, the shift to an opisthopubic pelvis is thought to be related to the development of a large stomach or stomachs and gut which would allow ornithischians to digest plant matter better.<ref name=":0" /> The smallest known Ornithischians are ''[[Fruitadens]] haagarorum''.<ref name=":4">{{Cite journal|last=Butler|first=Richard J.|last2=Galton|first2=Peter M.|last3=Porro|first3=Laura B.|last4=Chiappe|first4=Luis M.|last5=Henderson|first5=Donald M.|last6=Erickson|first6=Gregory M.|date=2010-02-07|title=Lower limits of ornithischian dinosaur body size inferred from a new Upper Jurassic heterodontosaurid from North America|url=http://rspb.royalsocietypublishing.org/content/277/1680/375|journal=Proceedings of the Royal Society of London B: Biological Sciences|language=en|volume=277|issue=1680|pages=375–381|doi=10.1098/rspb.2009.1494|issn=0962-8452|pmc=2842649|pmid=19846460}}</ref> The largest ''Fruitadens'' individuals reached just 65–75 cm. Previously, only carnivorous, saurischian theropods were known to reach such small sizes.<ref name=":4" /> At the other end of the spectrum, the largest known ornithischians reach about 15 meters (smaller than the largest saurischians).<ref>{{Cite journal|last=Yannan|first=Ji|last2=Xuri|first2=Wang|last3=Yongqing|first3=Liu|last4=Qiang|first4=Ji|date=2011-02-01|title=Systematics, Behavior and Living Environment of Shantungosaurus Giganteus (Dinosauria: Hadrosauridae)|url=http://onlinelibrary.wiley.com/doi/10.1111/j.1755-6724.2011.00378.x/abstract|journal=Acta Geologica Sinica - English Edition|language=en|volume=85|issue=1|pages=58–65|doi=10.1111/j.1755-6724.2011.00378.x|issn=1755-6724}}</ref>

However, not all ornithischians were strictly herbivorous. Some groups, like the [[heterodontosauridae|heterodontosaurids]], were likely [[omnivore]]s.<ref>{{cite journal | last1 = Barrett | first1 = P. M. | last2 = Rayfield | first2 = E. J. | year = 2006 | title = Ecological and evolutionary implications of dinosaur feeding behaviour | url = | journal = Trends in Ecology & Evolution | volume = 21 | issue = 4| pages = 217–224 | doi=10.1016/j.tree.2006.01.002}}</ref> At least one species of [[ankylosauria]]n, ''[[Liaoningosaurus paradoxus]]'', appears to have been at least partially [[carnivore|carnivorous]], with hooked claws, fork-like teeth, and stomach contents suggesting that it may have fed on fish.<ref name=liaoningosaurus_fish>{{cite journal | last1 = Ji | first1 = Q. | last2 = Wu | first2 = X. | last3 = Cheng | first3 = Y. | last4 = Ten | first4 = F. | last5 = Wang | first5 = X. | last6 = Ji | first6 = Y. | year = 2016 | title = Fish-hunting ankylosaurs (Dinosauria, Ornithischia) from the Cretaceous of China | url = | journal = Journal of Geology | volume = 40 | issue = | page = 2 }}</ref>

There is strong evidence that some ornithischians lived in herds.<ref name=":0" /><ref name=":1" /> This evidence consists of multiple [[Bone bed|bones beds]] with large numbers of the same species in different age classes who died simultaneously.<ref name=":0" /><ref name=":1" />

==References==
{{Reflist}}
*{{cite journal | last1 = Butler | first1 = R.J. | year = 2005 | title = The 'fabrosaurid' ornithischian dinosaurs of the Upper Elliot Formation (Lower Jurassic) of South Africa and Lesotho | url = | journal = Zoological Journal of the Linnean Society | volume = 145 | issue = 2| pages = 175–218 | doi=10.1111/j.1096-3642.2005.00182.x}}
*{{cite journal | last1 = Sereno | first1 = P.C. | year = 1986 | title = Phylogeny of the bird-hipped dinosaurs (order Ornithischia) | url = | journal = National Geographic Research | volume = 2 | issue = 2| pages = 234–256 }}

==External links==
{{Wikispecies|Ornithischia}}
{{Portal|Dinosaurs}}
* [http://www.palaeos.com/Vertebrates/Units/320Ornithischia/100.html#Ornithischia Ornithischia], from Palæos. (cladogram, characteristics)

{{Ornithodira|O.}}

{{taxonbar}}

[[Category:Ornithischians| ]]
[[Category:Carnian first appearances]]
[[Category:Maastrichtian extinctions]]
[[Category:Taxa named by Harry Seeley]]
[[Category:Fossil taxa described in 1888]]

Wikipedia:Administrator intervention against vandalism

2017-11-29T07:10:16Z

Wikkler:

<noinclude>{{#ifeq:{{PROTECTIONLEVEL:edit}}|autoconfirmed|{{pp|1=vandalism|action=edit|small=yes|expiry=}}}}<noinclude>[[Category:Wikipedia noticeboards]]{{/Header}}{{Floating link|Administrator instructions|Administrator instructions}}
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===User-reported===

*{{vandal|NickT19}} – actions evidently indicate a vandalism-only account. [[User:Tornado chaser|Tornado chaser]] ([[User talk:Tornado chaser|talk]]) 02:37, 29 November 2017 (UTC)
::{{AIV|n}} Another admin can block if they want but user had no warnings until I added one [[User:NeilN|NeilN]] [[User talk:NeilN|talk to me]] 02:46, 29 November 2017 (UTC)
*{{vandal|DannyHerrera98}} – Continues to disrupt articles with unsupported changes and fancruft after final warning, does not seem to listen to notices. [[User:SNUGGUMS|Snuggums]] ([[User talk:SNUGGUMS|talk]] / [[Special:Contributions/SNUGGUMS|edits]]) 03:26, 29 November 2017 (UTC)
*{{IPvandal|112.201.212.17}} – Persistent vandalism, Obvious Block evasion as IP Sock of [[User:Brianmagallano]]. [[User:LG-Gunther|LG-Gunther]] : [[User_talk:LG-Gunther| Talk ]] 03:56, 29 November 2017 (UTC) 
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*{{IPvandal|124.6.238.131}} – On [[Muhammad Hamidullah Khan]] ({{diff|Muhammad Hamidullah Khan|811964532||diff}}): vandalism after final warning. This user is changing large number of disputed information about Bangladesh but all of his edit summery says "Fixed typo". [[User:Mar11|Mar11]] ([[User talk:Mar11|talk]]) 04:35, 29 November 2017 (UTC)
* {{Vandal|Ilyasien_seven10}} Persistent Vandalism and harassment. Saying harshly in my user talks, continuously. [[User:Tommy1933|Tommy1933]] ([[User talk:Tommy1933|talk]]) 04:54, 29 November 2017 (UTC)
* {{vandal|Drlove2017}} Vandalizing a template in a similar way to the already blocked accounts, [[User:Kristen16]] and [[User:Destroyer100]]. I suspect that these are the same person since they're embedding the same image into assorted templates. Reverting the changes made to the templates seems to have no effect on the articles they are on... That [[User:Wikkler|Wikkler]] guy that nobody likes 07:04, 29 November 2017 (UTC)
:*[[User:Beth2017]] is another associated user. That [[User:Wikkler|Wikkler]] guy that nobody likes 07:10, 29 November 2017 (UTC)

Template:Cleanup section

2017-11-29T07:09:14Z

Wikkler: Undid revision 812683298 by Beth2017 (talk) Please.

{{ {{{|safesubst:}}}#invoke:Unsubst||date=__DATE__ |$B=
{{cleanup
|1=section
|reason={{{reason|<noinclude> </noinclude>}}}
|date={{{date|}}}
|small={{{small|left}}}
|talksection={{{talksection|}}}
|nocat={{{nocat|}}}
|demospace={{{demospace|<noinclude>main</noinclude>}}}
}}
}}<noinclude>
{{documentation}}
</noinclude>

Wikipedia:Administrator intervention against vandalism

2017-11-29T07:05:15Z

Wikkler: /* User-reported */

Wikipedia:Administrator intervention against vandalism

2017-11-29T07:04:56Z

Wikkler: /* User-reported */

<noinclude>{{#ifeq:{{PROTECTIONLEVEL:edit}}|autoconfirmed|{{pp|1=vandalism|action=edit|small=yes|expiry=}}}}<noinclude>[[Category:Wikipedia noticeboards]]{{/Header}}{{Floating link|Administrator instructions|Administrator instructions}}
__NEWSECTIONLINK__

{{adminbacklog}}</noinclude>


==Reports==
{{Wikipedia:Administrator intervention against vandalism/TB2}}

===User-reported===

*{{vandal|NickT19}} – actions evidently indicate a vandalism-only account. [[User:Tornado chaser|Tornado chaser]] ([[User talk:Tornado chaser|talk]]) 02:37, 29 November 2017 (UTC)
::{{AIV|n}} Another admin can block if they want but user had no warnings until I added one [[User:NeilN|NeilN]] [[User talk:NeilN|talk to me]] 02:46, 29 November 2017 (UTC)
*{{vandal|DannyHerrera98}} – Continues to disrupt articles with unsupported changes and fancruft after final warning, does not seem to listen to notices. [[User:SNUGGUMS|Snuggums]] ([[User talk:SNUGGUMS|talk]] / [[Special:Contributions/SNUGGUMS|edits]]) 03:26, 29 November 2017 (UTC)
*{{IPvandal|112.201.212.17}} – Persistent vandalism, Obvious Block evasion as IP Sock of [[User:Brianmagallano]]. [[User:LG-Gunther|LG-Gunther]] : [[User_talk:LG-Gunther| Talk ]] 03:56, 29 November 2017 (UTC) 
:*'''Note''': User is in the category: [[:Category:Suspected Wikipedia sockpuppets|Suspected Wikipedia sockpuppets]]. [[User:HBC AIV helperbot5|HBC AIV helperbot5]] ([[User talk:HBC AIV helperbot5|talk]]) 04:23, 29 November 2017 (UTC)
*{{IPvandal|71.191.190.229}} After filibustering previous attempts ([https://en.wikipedia.org/w/index.php?title=Wikipedia:Administrator_intervention_against_vandalism&diff=prev&oldid=812494446 this], which became stale for lack of his editing), this editor has resurfaced with continued willful insertion of incorrect information into articles. [[Special:Contributions/135.23.202.24|135.23.202.24]] ([[User talk:135.23.202.24|talk]]) 04:16, 29 November 2017 (UTC)
*{{IPvandal|124.6.238.131}} – On [[Muhammad Hamidullah Khan]] ({{diff|Muhammad Hamidullah Khan|811964532||diff}}): vandalism after final warning. This user is changing large number of disputed information about Bangladesh but all of his edit summery says "Fixed typo". [[User:Mar11|Mar11]] ([[User talk:Mar11|talk]]) 04:35, 29 November 2017 (UTC)
* {{Vandal|Ilyasien_seven10}} Persistent Vandalism and harassment. Saying harshly in my user talks, continuously. [[User:Tommy1933|Tommy1933]] ([[User talk:Tommy1933|talk]]) 04:54, 29 November 2017 (UTC)
* {vandal|Drlove2017]] Vandalizing a template in a similar way to the already blocked accounts, [[User:Kristen16]] and [[User:Destroyer100]]. I suspect that these are the same person since they're embedding the same image into assorted templates. Reverting the changes made to the templates seems to have no effect on the articles they are on... That [[User:Wikkler|Wikkler]] guy that nobody likes 07:04, 29 November 2017 (UTC)

Template:Use Oxford spelling

2017-11-29T06:55:50Z

Wikkler: Undid revision 812680244 by Drlove2017 (talk) Stop putting this picture in templates please.

{{ {{{|safesubst:}}}#invoke:Unsubst||date=__DATE__ |$B=
{{DMCA|Use British English Oxford spelling|from|{{{date|}}}}}
}}<noinclude>
{{Documentation}}
</noinclude>

Template:Graphical timeline

2017-11-29T06:25:13Z

Wikkler: Undid revision 812678323 by Destroyer100 (talk) No.

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2017-11-29T06:22:17Z

Wikkler: Undid revision 812677928 by Destroyer100 (talk) How about not having vandalism on this template?

Horned screamer

2017-11-24T20:20:53Z

Wikkler: /* Description */

{{speciesbox
| name = Horned screamer
| image = Anhima cornuta -near Manu Wildlife Center, Manu National Park, Peru -three-8.jpg
| image_caption = Three in Manu National Park, Peru [[File:Horned Screamer (Anhima cornuta) (W1CDR0000523 BD3).ogg|thumb|left|Horned Screamer calls, recorded in Amacayacu National Park, Colombia]]
| status = LC
| status_system = IUCN3.1
| status_ref = <ref>{{IUCN|id=22679723 |title=''Anhima cornuta'' |assessor=BirdLife International |assessor-link=BirdLife International |version=2013.2 |year=2012 |accessdate=26 November 2013}}</ref>
| genus = Anhima
| parent_authority = [[Mathurin Jacques Brisson|Brisson]], 1760
| species = cornuta
| authority = ([[Carl Linnaeus|Linnaeus]], 1766)
| range_map = Horned Screamer Range.png
| range_map_caption = Distribution map
}}

The '''horned screamer ''' (''Anhima cornuta'') is a member of a small family of [[bird]]s, the Anhimidae, which occurs in wetlands of tropical [[South America]]. There are three [[screamer]] species, the other two being the [[southern screamer]] and the [[northern screamer]] in the genus ''Chauna''. They are related to the [[duck]]s, [[goose|geese]] and [[swan]]s, which are in the family [[Anatidae]], but have bills looking more like those of [[Galliformes|game birds]].

==Description==
The horned screamer is a massive {{convert|84|–|95|cm|in|round=0.5|abbr=on}} long, {{convert|3.5|kg|lb|abbr=on}} bird, with a small [[chicken|chicken-like]] bill. The upperparts, head, and breast are black, with white speckles on the crown, throat and wing coverts. There is a long spiny structure projecting forward from the crown. This structure is unique among birds and is not derived from a feather but is a cornified structure that is loosely attached to the skull and grows continuously while often breaking at its tip. <ref>{{cite journal|journal=American Zoologist|volume=40|pages=461–477|year=2000| title=The Integumentary Morphology of Modern Birds—An Overview|first=Peter R.|last=Stettenheim|doi= 10.1093/icb/40.4.461}}</ref> This gives this species its name. It has very long and lanky legs and three large toes in each. The belly and under wing coverts are white. It has two sharp spurs on its wings and feet which are only partially webbed.

The horned screamer's call, as its name suggests, is a very loud echoing sound. It is called"El Clon-Clon" in Ecuador because of this peculiar feature{{Citation needed}}.

==Distribution, habitat and behavior==
The horned screamer is found in lowlands from [[Colombia]], [[Venezuela]], [[Brazil]], [[Bolivia]], [[Peru]], [[Ecuador]], [[Suriname]], [[French Guiana]], and [[Guyana]].<ref name="IUCN">IUCN (2009)</ref><ref name="Clements">Clements, J. (2007)</ref> It is now extinct in [[Trinidad and Tobago]].<ref name="IUCN"/> Despite having declined locally, it remains widespread and is fairly common overall. Its range in Brazil appears to have expanded in recent years.

It lives in well-vegetated marshes and feeds on water plants. Its nest is a large pile of floating vegetation anchored in shallow water. Three olive-brown eggs are laid, and the young, like those of most [[Anseriformes]], can run as soon as they are hatched.

==As a symbol==
The horned screamer is the official bird of the [[Arauca Department|Department of Arauca]] and the [[Arauca, Arauca|Municipality of Arauca]] in Colombia and it is also a symbol in the National Reserve of Churute in Ecuador. The department and its capital are named after the bird, which is called ''Arauco'' in [[Spanish language|Spanish]].

The bird appears in the arms of [[Tietê, São Paulo|Tietê]], Brazil.<ref name="camaratiete.com">[http://www.camaratiete.sp.gov.br/default.asp?c=2&s=2 camaratiete.com]</ref>

==Footnotes==
{{reflist}}

==References==
*Clements, James, (2007) [http://www.cornellpress.cornell.edu/cup_detail.taf?ti_id=4566 ''The Clements Checklist of the Birds of the World''], Cornell University Press, Ithaca
*Hilty, Steven, L. (2003) ''Birds of Venezuela'', {{ISBN|0-7136-6418-5}}

==External links==
{{Commons category|Anhima cornuta}}
*[http://ibc.lynxeds.com/species/horned-screamer-anhima-cornuta Horned Screamer] at Internet Bird Collection.

{{taxonbar}}

{{DEFAULTSORT:screamer, hooded}}
[[Category:Anhimidae|horned screamer]]
[[Category:Birds of Colombia]]
[[Category:Birds of Venezuela]]
[[Category:Birds of the Guianas]]
[[Category:Birds of Marajó]]
[[Category:Birds of the Amazon Basin]]
[[Category:Birds of Brazil]]
[[Category:Birds of Ecuador]]
[[Category:Birds described in 1766|horned screamer]]

Stegosauria

2017-11-07T10:42:16Z

Wikkler: You can't add something like this and not give any sources.

{{automatic taxobox
| name = Stegosaurians
| fossil_range = [[Middle Jurassic]] - [[Early Cretaceous]], {{fossilrange|earliest=Aalenian|165|125|latest=Maastrichtian}}Possible [[Aalenian]] and [[Maastrichtian]] records in the form of fossil tracks<ref name=NJGPA2017>{{cite journal |author=Peter M. Galton |year=2017 |title=Purported earliest bones of a plated dinosaur (Ornithischia: Stegosauria): a "dermal tail spine" and a centrum from the Aalenian-Bajocian (Middle Jurassic) of England, with comments on other early thyreophorans |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |volume=285 |issue=1 |pages=1–10 |doi=10.1127/njgpa/2017/0667 }}</ref><ref name=NJGPA20172>{{cite journal |author1=Peter M. Galton |author2=Krishnan Ayyasami |year=2017 |title=Purported latest bone of a plated dinosaur (Ornithischia: Stegosauria), a "dermal plate" from the Maastrichtian (Upper Cretaceous) of southern India |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |volume=285 |issue=1 |pages=91–96 |doi=10.1127/njgpa/2017/0671 }}</ref>
| image = Journal.pone.0138352.g001A.jpg
| image_caption = Mounted skeleton of ''[[Stegosaurus stenops]]'', [[Natural History Museum]], [[London]]
| authority = [[Othniel Charles Marsh|Marsh]], 1880
| type_species =
| type_species_authority =
| subdivision_ranks =Subgroups
| subdivision =
*{{extinct}}''[[Amargastegos]]''?
*{{extinct}}''[[Craterosaurus]]''?
*{{extinct}}''[[Gigantspinosaurus]]''
*{{extinct}}''[[Jiangjunosaurus]]''
*{{extinct}}''[[Paranthodon]]''
*{{extinct}}''[[Regnosaurus]]''?
*{{extinct}}'''Huayangosauridae'''
**{{extinct}}''[[Chungkingosaurus]]''
**{{extinct}}''[[Huayangosaurus]]''
*'''[[Stegosauridae]]'''
}}

'''Stegosauria''' is a group of [[Herbivore|herbivorous]] [[ornithischian]] [[dinosaur]]s that lived during the [[Jurassic]] and early [[Cretaceous]] [[Period (geology)|periods]]. Stegosaurian fossils have been found mostly in the [[Northern Hemisphere]], predominantly in what is now [[North America]], [[Europe]] and China, though one species (''[[Kentrosaurus aethiopicus]]'') is known to have lived in [[Africa]]. Their geographical origins are unclear; the earliest unequivocal stegosaurian, ''[[Huayangosaurus taibaii]]'', lived in China.

Stegosaurians were armored dinosaurs ([[thyreophora]]ns). Originally, they did not differ much from more primitive members of that group, being small, low-slung, running animals protected by armored [[scute]]s. An early evolutionary innovation was the development of tail spikes, or "[[thagomizer]]s", as defensive weapons. Later species, belonging to a subgroup called the [[Stegosauridae]], became larger, and developed long hindlimbs that no longer allowed them to run. This increased the importance of active defence by the thagomizer, which could ward off even large predators because the tail was in a higher position, pointing horizontally to the rear from the broad pelvis. Stegosaurids had complex arrays of spikes and plates running along their backs, hips and tails. Their necks became longer and their small heads became narrow, able to selectively bite off the best parts of [[cycad]]s with their beaks. When these plant types declined in diversity, so did the stegosaurians, which became extinct during the first half of the [[Cretaceous]] period.

The first stegosaurian finds in the early 19th century were fragmentary. Better fossil material, of the genus ''[[Dacentrurus]]'', was discovered in 1874 in England. Soon after, in 1877, the first nearly-complete skeleton was discovered in the United States. Professor [[Othniel Charles Marsh]] that year classified such specimens in the new genus ''[[Stegosaurus]]'', from which the group acquired its name, and which is still by far the most famous stegosaurian. During the latter half of the twentieth century, many important Chinese finds were made, representing about half of the presently known diversity of stegosaurians.

==Description==

===Skull===
Stegosaurians had characteristic small, long, flat, narrow heads and a horn-covered beak or [[beak#Anatomy|rhamphotheca]],<ref name=dinosauria2steg1>{{cite book|last=Galton|first=Peter|title=Dinosauria|year=2004|publisher=University of California Press|location=Berkeley|page=361|edition=2nd|author2=Paul Upchurch |editor=David B. Weishampel |editor2=Peter Dodson |editor3=Halszka Osmólska|chapter=16: Stegosauria}}</ref> which covered the front of the snout (two premaxillaries) and lower jaw (a single predentary) bones. Similar structures are seen in [[turtle]]s and [[bird]]s. Apart from ''[[Huayangosaurus]]'', stegosaurians subsequently lost all premaxillary teeth within the upper beak. ''Huayangosaurus'' still had seven per side.<ref>[[Paul Sereno|Sereno, P]] & [[Dong Zhiming|Z-M Dong]] (1992). The skull of the basal stegosaurian ''Huayangosaurus taibaii'' and a cladistic diagnosis of Stegosauria. ''Journal of Vertebrate Paleontology'' 51: 318-343</ref> The upper and lower jaws are equipped with rows of small teeth. Later species have a vertical bone plate covering the outer side of the lower jaw teeth. The structure of the upper jaw, with a low ridge above, and running parallel to, the tooth row, indicates the presence of a fleshy cheek. In stegosaurians, the typical [[archosauria]]n skull opening, the [[antorbital fenestra]] in front of the eye socket, is small, sometimes reduced to a narrow horizontal slit.

===Postcranial skeleton===
All stegosaurians are [[quadruped]]al, with hoof-like toes on all four limbs. All stegosaurians after ''Huayangosaurus'' have forelimbs much shorter than their hindlimbs. Their hindlimbs are long and straight, designed to carry the weight of the animal while stepping. The [[Knee#Articular bodies|condyles]] of the lower thighbone are short from the front to the rear. This would have limited the supported rotation of the knee joint, making running impossible. ''Huayangosaurus'' had a thighbone like a running animal. The upper leg was always longer than the lower leg.

''Huayangosaurus'' had relatively long and slender arms. The forelimbs of later forms are very robust, with a massive [[humerus]] and [[ulna]]. The [[Carpus|wrist bones]] were reinforced by a fusion into two blocks, an ulnar and a radial. The front feet of stegosaurians are commonly depicted in art and in museum displays with fingers splayed out and slanted downward. However, in this position, most bones in the hand would be disarticulated. In reality, the hand bones of stegosaurians were arranged into vertical columns, with the main fingers, orientated outwards, forming a tube-like structure. This is similar to the hands of [[Sauropoda|sauropod dinosaurs]], and is also supported by evidence from stegosaurian footprints and fossils found in a lifelike pose.<ref name=senter2010>{{cite journal|last=Senter|first=P.|year=2010|title=Evidence for a sauropod-like metacarpal configuration in stegosaurian dinosaurs|journal=Acta Palaeontologica Polonica|volume=55|issue=3|pages=427–432|doi=10.4202/app.2009.1105}}</ref>
[[File:DMSN dinosaurs.jpg|{{largethumb}}|''Stegosaurus'' mount showing to a good effect the high neck posture, the throat ossicles and the robust shoulder girdle and forelimbs]]
The long hindlimbs elevated the tail base, such that the tail pointed out behind the animal almost horizontally from that high position. While walking, the tail would not have sloped downwards as this would have impeded the function of the tail base retractor muscles, to pull the thighbones backwards. However, it has been suggested by [[Bob Bakker|Robert Thomas Bakker]] that stegosaurians could rear on their hind legs to reach higher layers of plants, the tail then being used as a "third leg". The mobility of the tail was increased by a reduction or absence of ossified tendons, that with many Ornithischia stiffen the hip region. ''Huayangosaurus'' still possessed them. In species that had short forelimbs, the relatively short torso towards the front curved strongly downwards. The dorsal vertebrae typically were very high, with very tall [[neural arch]]es and transverse processes pointing obliquely upwards to almost the level of the neural spine top. Stegosaurian back vertebrae can easily be identified by this unique configuration. The tall neural arches often house deep [[neural canal]]s; enlarged canals in the sacral vertebrae have given rise to the incorrect notion of a "second brain". Despite the downwards curvature of the rump, the neck base was not very low and the head was held a considerable distance off the ground. The neck was flexible and moderately long. ''Huayangosaurus'' still had the probably original number of nine cervical vertebrae; ''[[Miragaia (dinosaur)|Miragaia]]'' has an elongated neck with seventeen.<ref name=OMetal09/>

The stegosaurian [[shoulder girdle]] was very robust. In ''Huayangosaurus'', the [[acromion]], a process on the lower front edge of the [[shoulderblade]], was moderately developed; the [[coracoid]] was about as wide as the lower end of the [[scapula]], with which it formed the [[shoulder joint]]. Later forms tend to have a strongly expanded acromion, while the coracoid, largely attached to the acromion, no longer extends to the rear lower corner of the scapula. Ossified sternal plates have never been found with Stegosauria and perhaps the [[sternum]] was completely absent.

The stegosaurian pelvis was originally moderately large, as shown by ''Huayangosaurus''. Later species, however, convergent to the Ankylosauria developed very broad pelves, in which the [[iliac bone]]s formed wide horizontal plates with flaring front blades to allow for an enormous belly-gut. The ilia were attached to the sacral vertebrae via a sacral yoke formed by fused sacral ribs. ''Huayangosaurus'' still had rather long and obliquely oriented [[ischium|ischia]] and [[pubic bone]]s. In more derived species, these became more horizontal and shorter to the rear, while the front prepubic process lengthened.

===Osteoderms===
Like all [[Thyreophora]], stegosaurians were protected by bony [[scute]]s that were not part of the skeleton proper but skin ossifications instead: the so-called [[osteoderm]]s. ''Huayangosaurus'' had several types. On its neck, back, and tail were two rows of paired small vertical plates and spikes. On the rear of the tail, pairs of spikes were present forming the so-called "[[thagomizer]]", a defensive weapon. The very tail end bore a small club. Each flank had a row of smaller osteoderms, culminating in a long shoulder spine in front, curving to the rear. Later forms show very variable configurations, combining plates of various shape and size on the neck and front torso with spikes more to the rear of the animal. They seem to have lost the tail club and the flank rows are apparently absent also, with the exception of the shoulder spine, still shown by ''[[Kentrosaurus]]'' and extremely developed, as its name indicates, in ''[[Gigantspinosaurus]]''. As far as is known, all forms possessed some sort of thagomizer, though these are rarely preserved articulated allowing to establish the exact arrangement. A fossil of ''[[Chungkingosaurus]]'' sp. has been reported with three pairs of spikes pointing outwards and a fourth pair pointing to the rear.<ref>Z. Dong, S. Zhou, and Y. Zhang, 1983, "[Dinosaurs from the Jurassic of Sichuan]". ''Palaeontologica Sinica, New Series C'', '''162'''(23): 1-136</ref> The most derived species, like ''[[Stegosaurus]]'', ''[[Hesperosaurus]]'' and ''[[Wuerhosaurus]]'', have very large and flat back plates. To discern them from the smaller plates, which are intermediate to spines in having a thickened central section, these latter are sometimes called 'splates'. ''Stegosaurus'' plates are so large that it has been suggested that they were not arranged in paired but alternated rows or even formed a single overlapping midline row. With ''Stegosaurus'' fossils also ossicles have been found in the throat region, bony skin discs that protected the lower neck. Apart from protection, suggested functions of the osteoderms include display, species recognition and [[thermoregulation]].<ref>{{cite book|title=The Evolution and Extinction of the Dinosaurs (2nd Edition)|year=2005|chapter=Stegosauria:Hot Plates|editor1=Fastovsky D.E. |editor2=Weishampel D.B. |author1=Fastovsky D.E. |author2=Weishampel D.B. |pages=107–130|publisher=Cambridge University Press|isbn=0-521-81172-4}}</ref>

==Paleobiology==

===Trace fossils===
Stegosaurian tracks were first recognized in 1996 from a hindprint-only trackway discovered at the [[Cleveland-Lloyd Dinosaur Quarry|Cleveland-Lloyd quarry]], which is located near Price, Utah.<ref name="foster-stegosaur-tracks">"Walk and Don't Look Back: The Footprints; Stegosaurs" in Foster, J. (2007). Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. pg. 238</ref> Two years later, a new ichnogenus called ''[[Stegopodus]]'' was erected for another set of stegosaurian tracks which were found near [[Arches National Park]], also in Utah.<ref name="foster-stegosaur-tracks"/> Unlike the first, this trackway preserved traces of the forefeet. Fossil remains indicate that stegosaurians have five digits on the forefeet and three weight-bearing digits on the hind feet.<ref name="foster-stegosaur-tracks"/> From this, scientists were able to predict the appearance of stegosaurian tracks in 1990, six years in advance of the first actual discovery of [[Morrison Formation|Morrison]] stegosaurian tracks.<ref name="foster-stegosaur-tracks"/> More trackways have been found since the erection of ''Stegopodus''. None, however, have preserved traces of the front feet and stegosaurian traces remain rare.<ref name="foster-stegosaur-tracks"/>

==Evolutionary history==
[[File:Huayangosaurus BW.jpg|{{largethumb}}|''Huayangosaurus'' is the oldest and most basal stegosaurian of which good material is known, giving an impression of the build of the earliest members of the group]]
Like the spikes and shields of [[ankylosaur]]s, the [[bony plate]]s and spines of stegosaurians evolved from the low-keeled osteoderms characteristic of basal thyreophorans.<ref name="Norman">Norman, David (2001). "''Scelidosaurus'', the earliest complete dinosaur" in ''The Armored Dinosaurs'', pp 3-24. Bloomington: Indiana University Press. {{ISBN|0-253-33964-2}}.</ref> One such described genus, ''[[Scelidosaurus]]'', is proposed to be morphologically close to the [[last common ancestor]] of the clade uniting stegosaurians and ankylosaurians, the [[Eurypoda]].<ref name=CompleteDinosaur>{{cite book|last=Galton|first=Peter|title=The Complete Dinosaur|year=1997|publisher=Indiana University Press|location=Bloomington|isbn=9780253213136|url=https://books.google.com/?id=FOViD-lDPy0C&pg=PA291|editor=James O. Farlow |editor2=M. K. Brett-Surman|chapter=21: ''Stegosaurs''}}</ref> Footprints attributed to the [[ichnotaxon]] ''[[Deltapodus]] brodricki'' from the [[Middle Jurassic]] ([[Aalenian]]) of [[England]] represent the oldest probable record of stegosaurians reported so far.<ref name=NJGPA2017 /> The perhaps most basal known stegosaurian, the four metres long ''Huayangosaurus'', is still close to ''Scelidosaurus'' in build, with a higher and shorter skull, a short neck, a low torso, long slender forelimbs, short hindlimbs, large condyles on the thighbone, a narrow pelvis, long ischial and pubic shafts and a relatively long tail. Its small tail club might be a eurypodan [[synapomorphy]]. ''Huayangosaurus'' lived during the [[Bathonian]] stage of the [[Middle Jurassic]], about 166 million years ago.<ref>Marco Romano (2017). Disparity vs. diversity in Stegosauria (Dinosauria, Ornithischia): cranial and post-cranial sub-dataset provide different signals. ''Historical Biology.'' doi: http://dx.doi.org/10.1080/08912963.2017.1397655</ref>

A few million years later, during the [[Callovian]]-[[Oxfordian (stage)|Oxfordian]], from China much larger species are known, with long, "graviportal", i.e., a type adapted for moving only in a slow manner on land due to a high body weight, hindlimbs: ''[[Chungkingosaurus]]'', ''[[Chialingosaurus]]'', ''[[Tuojiangosaurus]]'' and ''[[Gigantspinosaurus]]''. Most of these are considered members of the derived [[Stegosauridae]]. ''[[Lexovisaurus]]'' and ''[[Loricatosaurus]]'', stegosaurid finds from England and France of approximately equivalent age to the Chinese specimens, are likely the same [[taxon]]. During the [[Late Jurassic]], stegosaurids seem to have experienced their greatest radiation. In Europe, ''[[Dacentrurus]]'' and the closely related ''[[Miragaia (dinosaur)|Miragaia]]'' were present. While older finds had been limited to the northern continents, in this phase [[Gondwana]] was colonised also as shown by ''[[Kentrosaurus]]'' living in [[Africa]]. No unequivocal stegosaurian fossils have been reported from [[South-America]], India, [[Madagascar]], Australia or [[Antarctica]], though. A Late Jurassic Chinese stegosaurian is ''[[Jiangjunosaurus]]''. The most derived Jurassic stegosaurians are known from [[North-America]]: ''Stegosaurus'' (perhaps several species thereof) and the somewhat older ''[[Hesperosaurus]]''. ''Stegosaurus'' was quite large (some specimens indicate a length of at least seven metres), had high plates, no shoulder spine, and a short, deep rump.

From the [[Early Cretaceous]], far fewer finds are known and it seems that the group had declined in diversity. Some fragmentary fossils have been described, such as ''[[Craterosaurus]]'' from England and ''[[Paranthodon]]'' from [[South Africa]]. The only more substantial discoveries are those of ''[[Wuerhosaurus]]'', the exact age of which is highly uncertain.<ref name="Holtz2011">{{Cite book | last1 = Holtz | first1 = Thomas R., Jr. | last2 = Rey | first2 = Luis V. | title = Dinosaurs: the most complete, up-to-date encyclopedia for dinosaur lovers of all ages | url = http://www.geol.umd.edu/~tholtz/dinoappendix/HoltzappendixWinter2011.pdf |year = 2007 | publisher = Random House | location = New York | isbn = 978-0-375-82419-7 }}</ref> In the fall of 2016, the teeth and tail spikes of stegosaurs were found in a [[Russia]]n ravine that was once part of an Early Cretaceous river. The remains of these stegosaurs and other dinosaurs in the area were all miniature, implying that either large amounts of baby dinosaurs spent the first stage in their lives here, or that the area was home to [[Insular dwarfism|dwarf]] dinosaurs. Several Russian scientists also theorize that the remains of these stegosaurs, as well as the remains of allosaurids in the same area, could imply that the area could have once been a "refugium", were these Jurassic dinosaurs managed to survive into the Cretaceous period.<ref>{{Cite web|url=http://siberiantimes.com/science/casestudy/news/n0730-dinosaur-kindergarten-found-washed-up-on-banks-of-ancient-river-scientists-believe/|title=Dinosaur 'kindergarten' found washed up on banks of ancient river, scientists believe|website=siberiantimes.com|access-date=2016-09-14}}</ref>

It has often been suggested that the stegasaurid decline was part of a Jurassic/Cretaceous transition, a [[faunal turnover]] caused by a [[floral turnover]], the new group of the [[angiosperms]] becoming the dominant plants, which in turn led to new groups of herbivores.<ref>Bakker, R.T., 1998, "Dinosaur mid-life crisis: the Jurassic-Cretaceous transition in Wyoming and Colorado", In: S.G. Lucas, J.I. Kirkland, & J.W. Estep (eds.) ''Lower and Middle Cretaceous Terrestrial Ecosystems; New Mexico Museum of Natural History and Science Bulletin'', '''14''': 67-77</ref> Although in general the case for such a causal relation is poorly supported by the data, stegosaurians are an exception in that their decline coincides with that of the [[Cycadophyta]].<ref>Butler, R.J., Barrett, P.M., Kenrick, P. and Penn, M.G., 2009, "Diversity patterns amongst herbivorous dinosaurs and plants during the Cretaceous: implications for hypotheses of dinosaur/angiosperm co-evolution", ''Journal of Evolutionary Biology'', '''22''': 446–459</ref>

Though Late Cretaceous stegosaurian fossils have been reported, these have mostly turned out to be misidentified. A well-known example is ''[[Dravidosaurus]]'', known from [[Coniacian]] fossils found in India. Though originally thought to be stegosaurian, in 1991 these badly-eroded fossils were suggested to instead have been based on [[plesiosauria]]n pelvis and hindlimb material,<ref>Chatterjee, S., and Rudra, D.K., 1996, "KT events in India: impact, rifting, volcanism and dinosaur extinction," in: Novas & Molnar, eds., ''Proceedings of the Gondwanan Dinosaur Symposium, Brisbane, Memoirs of the Queensland Museum'', '''39'''(3): iv + 489–731 : 489-532</ref> and none of the fossils are demonstrably stegosaurian.<ref>Wilson, J. A., Barrett, P. M., & Carrano, M. T. (2011). An associated partial skeleton of ''Jainosaurus cf. septentrionalis'' (Dinosauria: Sauropoda) from the Late Cretaceous of Chhota Simla, central India. ''Palaeontology'', 54(5), 981-998.</ref> The reinterpretation of ''Dravidosaurus'' as a plesiosaur wasn't accepted by [[Peter Galton|Galton]] and Upchurch (2004), who stated that the skull and plates of ''Dravidosaurus'' are certainly not plesiosaurian, and noted the need to redescribe the fossil material of ''Dravidosaurus''.<ref>{{cite book |author1=Peter M. Galton |author2=Paul Upchurch |editor1=[[David B. Weishampel]] |editor2=Peter Dodson |editor3=[[Halszka Osmólska]] |title=The Dinosauria |edition=2nd |year=2004|publisher=University of California Press |place=Berkeley |isbn=0-520-24209-2 |pages=343–362 |chapter=Stegosauria }}</ref> Purported stegosaurian dermal plate was reported from the latest Cretaceous ([[Maastrichtian]]) [[Kallamedu Formation]] (southern India); however, Galton & Ayyasami (2017) interpreted the specimen as a bone of a sauropod dinosaur. Nevertheless, the authors considered the survival of stegosaurians into the Maastrichtian to be possible, noting the presence of the stegosaurian ichnotaxon ''Deltapodus'' in the Maastrichtian [[Lameta Formation]] (western India).<ref name=NJGPA20172 />

==Classification==
[[File:FieldMuseum5 Chicago.jpg|thumb|250px|right|A fossil melee involving a stegosaurian (''[[Tuojiangosaurus]]'') and a mid-sized [[theropod]] (''[[Monolophosaurus]]''), [[Field Museum]] in [[Chicago]]]]
The Stegosauria was originally named as an order within [[Reptilia]] by [[Othniel Charles Marsh|O.C. Marsh]] in 1877,<ref name="marsh1877">Marsh, O.C. (1877). "New order of extinct Reptilia (Stegosauria) from the Jurassic of the Rocky Mountains." ''American Journal of Science'', '''14'''(ser.3):513-514.</ref> although today they are generally treated as an infraorder or suborder — or more often an unranked [[clade]] — within the [[Thyreophora]], the [[Armour (zoology)|armored]] dinosaurs. It includes in modern usage the families [[Huayangosauridae]] and [[Stegosauridae]], named in 1982 and 1880 respectively.

The Huayangosauridae were an early group of stegosaurians that lived during the early to middle [[Jurassic]] Period. They were smaller than later stegosaurians and had shorter and higher skulls. Huayangosauridae is undefined. Currently, the only unequivocal genus included is the [[type genus]] ''[[Huayangosaurus]]'' of China. The poorly known remains of ''[[Regnosaurus]]'' from the early Cretaceous of England, however, indicate that it too could be a member — or at least a basal stegosaurian. They consist of a lower jaw that is very similar to that of the former genus.

The vast majority of stegosaurian dinosaurs thus far recovered belong to the Stegosauridae, which lived in the later part of the Jurassic and early Cretaceous, and which were defined by [[Paul Sereno]] as all stegosaurians more closely related to ''Stegosaurus'' than to ''Huayangosaurus''.<ref>Sereno, P.C., 1998, "A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria", ''Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen'' '''210''': 41-83</ref> They include per definition the well-known ''[[Stegosaurus]]''. This group is widespread, with members across the Northern Hemisphere, [[Africa]] and possibly [[South America]].<ref name=Pereda-Suberbiola2013>{{cite journal|last=Pereda-Suberbiola|first=Xabier|author2=Galton, Peter M. |author3=Mallison, Heinrich |author4= Novas, Fernando |title=A plated dinosaur (Ornithischia, Stegosauria) from the Early Cretaceous of Argentina, South America: an evaluation|journal=Alcheringa: An Australasian Journal of Palaeontology|year=2013|volume=37|issue=1|pages=65–78|doi=10.1080/03115518.2012.702531|url=http://www.tandfonline.com/doi/abs/10.1080/03115518.2012.702531#preview}}</ref>

The first exact clade definition of Stegosauria was given by [[Peter Malcolm Galton]] in 1997: all thyreophoran [[Ornithischia]] more closely related to ''Stegosaurus'' than to ''[[Ankylosaurus]]''.<ref>Galton, P.M., 1997, "Stegosauria", pp. 701-703 in: P.J. Currie and K. Padian (eds.), ''Encyclopedia of Dinosaurs'', Academic Press, San Diego</ref> Thus defined, the Stegosauria are by definition the [[sister group]] of the [[Ankylosauria]] within the [[Eurypoda]].

===Phylogeny===
[[Kenneth Carpenter]] of the [[Denver Museum of Nature and Science]] published a preliminary phyletic tree<ref name="carpenter2001">Carpenter, K., Miles, C.A., and Cloward, K. (2001). "New Primitive Stegosaur from the Morrison Formation, Wyoming", in Carpenter, Kenneth(ed) ''The Armored Dinosaurs''. Indiana University Press. {{ISBN|0-253-33964-2}}, 55–75.</ref> of stegosaurians, in the 2001 description of ''Hesperosaurus''. An updated phylogeny was published by Mateus ''et al.'' (2009), which is shown below.<ref name=OMetal09>{{Cite journal |last=Mateus |first=Octávio |author2=Maidment, Susannah C.R. |author3= Christiansen, Nicolai A. |title=A new long-necked 'sauropod-mimic' stegosaur and the evolution of the plated dinosaurs |journal=Proceedings of the Royal Society B: Biological Sciences |year=2009 |volume=276 |url=http://rspb.royalsocietypublishing.org/content/early/2009/02/21/rspb.2008.1909.full.pdf+html |format=pdf |doi=10.1098/rspb.2008.1909 |pmid=19324778 |issue=1663 |pages=1815–21 |pmc=2674496}}</ref>
{{clade|style=font-size:90%;line-height:90%
|label1='''Stegosauria'''
|1={{clade
|1=''[[Tuojiangosaurus]]''
|2=''[[Paranthodon]]''
|3=''[[Gigantspinosaurus]]''
|label4=[[Huayangosauridae]]
|4={{clade
|1=''[[Huayangosaurus]]''
|2=''[[Chungkingosaurus]]'' }}
|label5=[[Stegosauridae]]
|5={{clade
|1=''[[Chialingosaurus]]''
|2={{clade
|1=''[[Kentrosaurus]]''
|2={{clade
|1=''[[Loricatosaurus]]''
|2={{clade
|label1=[[Dacentrurinae]]
|1={{clade
|1=''[[Dacentrurus]]''
|2=''[[Miragaia (dinosaur)|Miragaia]]'' }}
|label2=[[Stegosaurinae]]
|2={{clade
|1=''[[Stegosaurus]]''
|2={{clade
|1=''[[Wuerhosaurus]]''
|2=''[[Hesperosaurus]]''
}} }} }} }} }} }} }} }}

===Undescribed species===
To date, several genera from China bearing names have been proposed but not formally described, including "[[Changdusaurus]]".<ref name=Maidment>{{cite journal| last= Maidment| first= Susannah C.R.| authorlink = |author2=Guangbiao Wei | title= A review of the Late Jurassic stegosaurs (Dinosauria, Stegosauria) from the People's Republic of China| journal = Geological Magazine| volume = 143| issue = 5| pages = 621–634| publisher = | location = | year = 2006| url = http://geolmag.geoscienceworld.org/cgi/content/abstract/143/5/621| doi = 10.1017/S0016756806002500|}}</ref> Until formal descriptions are published, these genera are regarded as ''[[nomen nudum|nomina nuda]]''. ''[[Yingshanosaurus]]'', for a long time considered a ''nomen nudum'', was described in 1994.<ref>Zhu Songlin, 1994, "记四川盆地营山县一剑龙化石 [Record of a fossil stegosaur from Yingshan in the Sichuan Basin]", ''Sichuan Cultural Relics'', '''1994'''(S1): 8-14</ref>

==Discovery==
{{see also|Timeline of stegosaur research}}

The first known discovery of a possible stegosaurian was probably made in the early nineteenth century in England. It consisted of a lower jaw fragment and was in 1848 named ''[[Regnosaurus]]''. In 1845, in the area of the present state of [[South Africa]], remains were discovered that much later would be named ''[[Paranthodon]]''. In 1874, from England, other remains were named ''[[Craterosaurus]]'' was named. All three taxa were based on fragmentary material and were not recognised as possible stegosaurians until the twentieth century. They gave no reason to suspect the existence of a new distinctive group of dinosaurs.

In 1874, extensive remains, the first partial stegosaurian skeleton known,<ref name="Maidment2010"/> were uncovered in England of what was clearly a large herbivore equipped with spikes. They were named ''Omosaurus'' by [[Richard Owen]] in 1875. Later, this name was shown to be preoccupied by the [[phytosaur]] ''[[Omosaurus]]'' and the stegosaurian was renamed ''[[Dacentrurus]]''. Other English nineteenth century and early twentieth century finds would be assigned to ''Omosaurus''; later they would, together with French fossils, be partly renamed ''[[Lexovisaurus]]'' and ''[[Loricatosaurus]]''. None of these specimens was complete though, and even together they could not have provided a good understanding of stegosaurian build. Owen e.g., initially assumed that the spikes were placed on the wrists. However, very soon after the discovery of ''Omosaurus'', American finds would fully compensate for this.
[[File:Position of bones in skeleton sketched by Arthur Lakes.jpg|{{largethumb}}|''Stegosaurus'' bones illustrated by Arthur Lakes in 1879]]
In 1877, [[Arthur Lakes]], a fossil hunter working for Professor [[Othniel Charles Marsh]], in [[Wyoming]] excavated a fossil that Marsh the same year named ''[[Stegosaurus]]''. At first, Marsh still entertained some incorrect notions about its morphology. He assumed that the plates formed a flat skin cover — hence the name, meaning "roof saurian" — and that the animal was bipedal with the spikes sticking out sideways from the rear of the skull. A succession of additional discoveries from the [[Como Bluff]] sites allowed a quick update of the presumed build. In 1882, Marsh was able to publish the first skeletal reconstruction of a stegosaur. Hereby, stegosaurians became much better known to the general public. The American finds at the time represented the bulk of known stegosaurian fossils, with about twenty skeletons collected.<ref name="Maidment2010">Maidment, S.C.R., 2010, "Stegosauria: A review of the body fossil record and phylogenetic relationships", ''Swiss Journal of Geosciences'', '''103''': 199-210</ref>

The next important discovery was made when a German expedition to the [[Tendaguru]], then part of [[German East Africa]], from 1909 to 1912 excavated over a thousand bones of ''[[Kentrosaurus]]''. The finds increased the known variability of the group, ''Kentrosaurus'' being rather small and having long rows of spikes on the hip and tail.

After 1912, Western researchers for a long time failed to identify any new stegosaurians, scientific interest in dinosaurs as whole being rather limited during the middle of the twentieth century. From the 1950s onwards, the geology of China was systematically surveyed in detail and infrastructural works led to a vast increase of digging activities in that country. This resulted in a new wave of Chinese stegosaurian discoveries, starting with ''[[Chialingosaurus]]'' in 1957. Chinese finds of the 1970s and 1980s included ''[[Wuerhosaurus]]'', ''[[Tuojiangosaurus]]'', ''[[Chungkingosaurus]]'', ''[[Huayangosaurus]]'', ''[[Yingshanosaurus]]'' and ''[[Gigantspinosaurus]]''. This increased the age range of good fossil stegosaurian material, as they represented the first relatively complete skeletons from the Middle Jurassic and the Early Cretaceous. Especially important was ''Huayangosaurus'', which provided unique information about the early evolution of the group.

Towards the end of the twentieth century, the so-called [[Dinosaur Renaissance]] took place in which a vast increase in scientific attention was given to the Dinosauria. As part of this development, the rate of dinosaurian discoveries quickly picked up. However, this has not resulted in a peak of stegosaurian finds, partly because most new sites are from the Cretaceous, when stegosaurian diversity had declined. In 2007, ''[[Jiangjunosaurus]]'' was reported, the first Chinese dinosaur named since 1994. Nevertheless, European and North-American sites have become productive again during the 1990s, ''[[Miragaia (dinosaur)|Miragaia]]'' having been found in [[Portugal]] and a number of relatively complete ''[[Hesperosaurus]]'' skeletons having been excavated in Wyoming. Apart from the fossils per se, important new insights have been gained by applying the method of [[cladistics]], allowing for the first time to exactly calculate stegosaurian evolutionary relationships.

The following timeline shows the date of descriptions for valid stegosaurian genera beginning in 1824, when the first non-avian dinosaur, ''[[Megalosaurus]]'', was formally described. The fossils themselves were found earlier; in the case of ''Loricatosaurus'' e.g. there is a gap of 107 years between the discovery and the naming of the genus.

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color:1900s bar:NAM5 at:1992 mark:(line,black) text:[[Gigantspinosaurus]]
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color:1900s bar:NAM1 at:2007 mark:(line,black) text:[[Jiangjunosaurus]]
color:1900s bar:NAM3 at:2007 mark:(line,black) text:[[Loricatosaurus]]
color:1900s bar:NAM1 at:1874 mark:(line,black) text:[[Craterosaurus]]
color:1900s bar:NAM1 at:1977 mark:(line,black) text:[[Tuojiangosaurus]]
color:1900s bar:NAM3 at:1877 mark:(line,black) text:[[Stegosaurus]]
color:1800s bar:NAM4 at:1887 mark:(line,black) text:[[Lexovisaurus]]
color:1900s bar:NAM1 at:1915 mark:(line,black) text:[[Kentrosaurus]]
color:1900s bar:NAM4 at:1973 mark:(line,black) text:[[Wuerhosaurus]]
color:1800s bar:NAM2 at:1929 mark:(line,black) text:[[Paranthodon]]
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</timeline>

==See also==
{{Portal|Dinosaurs}}

==References==
{{reflist}}

==External links==
* [http://palaeos.com/vertebrates/ornithischia/stegosauria.html Stegosauria on Palaeos.com]
* http://www.kheper.net/evolution/dinosauria/Stegosauria.htm

{{Thyreophora|S.}}

{{taxonbar}}

[[Category:Stegosaurs| ]]
[[Category:Bajocian first appearances]]
[[Category:Early Cretaceous extinctions]]
[[Category:Fossil taxa described in 1877]]
[[Category:Taxa named by Othniel Charles Marsh]]

Coeluridae

2017-11-07T05:01:04Z

Wikkler: /* List of types names */ No modern analysis has recovered these as coelurids.

{{Automatic taxobox
| name = Coelurids
| fossil_range = [[Late Jurassic]], {{Fossil range|153|150}}
| image = Tanycolagreus.jpg
| image_width = 250px
| image_caption = ''Tanycolagreus'' fossil cast, [[North American Museum of Ancient Life]].
| authority = [[Othniel Charles Marsh|Marsh]], 1881
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
*''[[Coelurus]]'' ([[Type (zoology)|type]])
*''[[Tanycolagreus]]''?
}}

'''Coeluridae''' is a historically [[polyphyletic|unnatural group]] of generally small, [[Carnivore|carnivorous]] [[dinosaur]]s from the late [[Jurassic]] [[Period (geology)|Period]]. For many years, any small Jurassic or Cretaceous [[theropoda|theropod]] that did not belong to one of the more specialized families recognized at the time was classified with the coelurids, creating a confusing array of 'coelurid' theropods that were not closely related. Although they have been traditionally included in this family, there is no evidence that any of these primitive [[coelurosaur]]s form a [[clade|natural group]] with ''[[Coelurus]]'', the namesake of Coeluridae, to the exclusion of other traditional coelurosaur groups.

==Classification==
Before the use of phylogenetic analyses, Coeluridae and [[Coelurosauria]] were [[wastebasket taxon|taxonomic wastebaskets]] used for small theropods that did not belong to other groups; thus, they accumulated many [[nomen dubium|dubious]] genera.<ref name=GSP88>{{cite book|last=Paul |first=G.S. |year=1988 |title=Predatory Dinosaurs of the World |place=New York |publisher=Simon and Schuster |pages=248–250|isbn=0-671-61946-2 }}</ref><ref name=DBN90>{{cite book |last=Norman |first=David B. |authorlink=David B. Norman |year=1990 |chapter=Problematic theropoda: "coelurosaurs" |editor=Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |publisher=University of California Press |location=Berkeley |pages=280–305 |isbn=0-520-06727-4}}</ref> As late as the 1980s, popular books recognized over a dozen "coelurids", including such disparate forms as the [[Noasauridae|noasaurid]] ''[[Laevisuchus]]'' and the [[oviraptorosauria]]n ''[[Microvenator]]'', and considered them descendants of the [[coelophysidae|coelophysids]].<ref name=DL83>{{cite book |last=Lambert |first=David |author2=the Diagram Group |title=A Field Guide to Dinosaurs |year=1983 |publisher=Avon Books |location=New York |isbn=0-380-83519-3 |chapter= Coelurids |pages=44–47}}</ref> A wastebasket Coeluridae lingered into the early 1990s in some sources<ref name=LG93>{{cite book|title=The Dinosaur Society Dinosaur Encyclopedia|year=1993|author1=Lessem, Don |author2=Glut, Donald F.|publisher=Random House, Inc. |isbn=0-679-41770-2}}</ref> (and appears in at least one 2006 source)<ref>{{cite book|title=The Illustrated Encyclopedia of the Prehistoric World|year=2006|author=Palmer, Douglas|publisher=Chartwell Books, Inc|location=New Jersey|chapter=Ruling Reptiles: Dinosaurs and Their Kin|pages=232–313|isbn=0-7858-2086-8}}</ref> but since then it has only been recognized in a much reduced form.<ref name=OR03/><ref name=PS07/>

In 2003, O.W.M. Rauhut, using a [[cladistic]] analysis, found Coeluridae to include ''[[Coelurus]]'' ([[Late Jurassic]], North America), ''[[Compsognathus]]'' (Late Jurassic, Europe), ''[[Sinosauropteryx]]'' ([[Early Cretaceous]], Asia) and an unnamed ''[[Compsognathus]]''-like form (Early Cretaceous, South America; this dinosaur has since been placed in the new genus ''[[Mirischia]]''). Rauhut considered coelurids to be a [[monophyletic]] group of basal [[coelurosauria|coelurosaurs]], characterized by evolutionary reversals in some aspects of the vertebrae to the more primitive theropod condition.<ref name=OR03>{{cite journal |last=Rauhut |first=Oliver W.M. |year=2003 |title=The interrelationships and evolution of basal theropod dinosaurs |journal=Special Papers in Palaeontology |volume=69 |pages=1–213}}</ref> However, he and other authors have not since found this result.<ref name=PS07>{{cite_journal |last=Senter |first=Phil |year=2007 |title=A new look at the phylogeny of Coelurosauria (Dinosauria, Theropoda) |journal=Journal of Systematic Palaeontology |volume=5 |issue=4 |pages=429–463 |doi=10.1017/S1477201907002143}}</ref><ref name=HMC04>{{cite book|last=Holtz |first=Thomas R., Jr. |author2=Molnar, Ralph E. |author3= Currie, Philip J. |year=2004 |editor=Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |edition=2nd |publisher=University of California Press |location=Berkeley |isbn=0-520-24209-2 |pages=71–110 }}</ref><ref name=RX05>{{cite journal |last=Rauhut |first=Oliver W.M. |author2=Xu, Xing |year=2005 |title=The small theropod dinosaurs ''Tugulusaurus'' and ''Phaedrolosaurus'' from the Early Cretaceous of Xinjiang, China |journal=Journal of Vertebrate Paleontology |volume=25 |issue=1 |pages=107–118 |doi=10.1671/0272-4634(2005)025[0107:TSTDTA]2.0.CO;2}}</ref> Phil Senter proposed in 2007 that ''Coelurus'' and ''Tanycolagreus'' were the only coelurids, and were actually [[tyrannosauroidea|tyrannosauroids]].<ref name=PS07/>

Coeluridae received a formal phylogenetic definition in 2015, when it was defined as all species more closely related to ''Coelurus fragilis'' than to ''Proceratosaurus bradleyi'', ''Tyrannosaurus rex'', ''Allosaurus fragilis'', ''Compsognathus longipes'', ''Ornithomimus edmontonicus'', or ''Deinonychus antirrhopus'' by Hendrickx, Hartman and Mateus.<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> It remains unclear whether or not this group contains any species other than ''Coelurus'' itself, and while ''Tanycolagreus'' is often included, support for this relationship has been weak in most of the studies that recovered it.<ref name=tyrannosauroid_phylogeny_2016>Brusatte, S. L., & Carr, T. D. (2016). [http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4735739/ The phylogeny and evolutionary history of tyrannosauroid dinosaurs.] ''Scientific Reports'', 6.</ref>

Below is a cladogram placing ''Coelurus'' in Coelurosauria by Cau ''et al.'' in 2015.<ref name=cauetal2015>{{cite journal|doi=10.7717/peerj.1032|pmid=26157616|pmc=4476167|title=The phylogenetic affinities of the bizarre Late Cretaceous Romanian theropod ''Balaur'' bondoc (Dinosauria, Maniraptora): Dromaeosaurid or flightless bird?|journal=PeerJ|volume=3|pages=e1032|year=2015|last1=Cau|first1=Andrea|last2=Brougham|first2=Tom|last3=Naish|first3=Darren}}</ref>

{{clade| style=font-size:100%; line-height:100%
|label1=[[Coelurosauria]]
|1={{clade
|1=''[[Zuolong]]''
|2={{clade
|1=''[[Tugulusaurus]]''
|label2=[[Tyrannoraptora]]
|2={{clade
|1={{clade
|label1=[[Tyrannosauroidea]]
|1={{clade
|1='''''Coelurus'''''
|2=''[[Tanycolagreus]]''
|3={{clade
|1=''[[Guanlong]]''
|2={{clade
|1=''[[Dilong (dinosaur)|Dilong]]''
|2={{clade
|1=''[[Yutyrannus]]''
|2=''[[Tyrannosaurus]]'' }} }} }} }} }}
|2={{clade
|1=''[[Sinocalliopteryx]]''
|2={{clade
|1=[[Compsognathidae]]
|2=[[Maniraptoriformes]]
}} }} }} }} }} }}
==List of types names==
'''Family Coeluridae'''

'''Subamily Coleurinae'''

*''[[Coelurus fragilis]]''

*''[[Kakuru kujani]]''{{citation needed}}

*''[[Tanycolagreus topwilsoni ]]''

'''Subfamily [[Ornitholestinae]]'''

*''[[Ornitholestes hermanni]]''{{citation needed}}

==References==
{{Reflist}}

{{Portal|Dinosaurs}}

{{Tyrannosauroidea}}
{{taxonbar}}

[[Category:Coelurosaurs]]
[[Category:Kimmeridgian first appearances]]
[[Category:Late Jurassic taxonomic families]]
[[Category:Late Jurassic extinctions]]
[[Category:Taxa named by Othniel Charles Marsh]]

Pikaia

2017-11-05T16:20:17Z

Wikkler:

{{About|the extinct chordate|the anime series|Pikaia!}}
{{italic title}}{{speciesbox
| name = ''Pikaia''
| fossil_range = {{Fossil range|Middle Cambrian}}
| image = Pikaia gracilens B.jpg
| image_caption = Life reconstruction of ''Pikaia gracilens''
| parent_authority = [[Charles Doolittle Walcott|Walcott]], 1911
| genus = Pikaia
| authority = Walcott, 1911
| species = gracilens
}}
'''''Pikaia gracilens''''' is an extinct [[cephalochordata|cephalochordate]] animal known from the Middle [[Cambrian]] [[Burgess Shale]] of [[British Columbia]]. Sixteen specimens are known from the Greater [[Phyllopod bed]], where they comprised 0.03% of the community.<ref>[http://burgess-shale.rom.on.ca/en/fossil-gallery/view-species.php?id=101 "''Pikaia gracilens''"] ''Burgess Shale Fossil Gallery''. Virtual Museum of Canada. 2011.</ref> It resembled the [[lancelet]] and perhaps swam much like an [[eel]].

== Description ==
[[File:Pikaia NT small.jpg|thumb|left|Life reconstruction of ''Pikaia gracilens'']]

''Pikaia'' is a primitive chordate that lacked a well-defined head and averaged about {{convert|1+1/2|in|cm}} in length. Because it was once thought to be closely related to the ancestor of all [[vertebrate]]s; as such, it receives otherwise-inordinate attention among the multitude of animal fossils found in the famous [[Burgess Shale]] in the mountains of [[British Columbia]], Canada. ''Pikaia'' had a pair of large, antenna-like tentacles on its head, and a series of short appendages, which may be linked to gill slits, on either side of its head. In these ways, it differs from the living lancelet. The "[[tentacle]]s" on its head may be comparable to those in the present-day [[hagfish]], a jawless chordate.{{Citation needed|date=November 2007}}

Although primitive, ''Pikaia'' shows the essential prerequisites for [[vertebrates]]. When alive, ''Pikaia'' was a compressed, leaf-shaped animal with an expanded tail fin; the flattened body is divided into pairs of [[myomere|segmented muscle block]]s, seen as faint vertical lines. The muscles lie on either side of a flexible structure resembling a rod that runs from the tip of the head to the tip of the tail.<ref name=autogenerated1>Palmer, D., (2000). ''The Atlas of the Prehistoric World''. London: Marshall Publishing Ltd. p66-67.</ref> It likely swam by throwing its body into a series of S-shaped, zigzag curves, similar to the movement of eels; fish inherited the same swimming movement, but they generally have stiffer backbones. These adaptations may have allowed ''Pikaia'' to filter particles from the water as it swam along.{{Citation needed|date=November 2008}} ''Pikaia'' was probably a slow swimmer, since it lacked the fast-twitch fibers that are associated with rapid swimming in modern chordates.<ref name="Lacalli2012">{{Cite journal | last1 = Lacalli | first1 = T. | title = The Middle Cambrian fossil Pikaia and the evolution of chordate swimming | doi = 10.1186/2041-9139-3-12 | journal = EvoDevo | volume = 3 | issue = 1 | pages = 12 | year = 2012 | pmid = 22695332| pmc =3390900 }}</ref>

Conway Morris and Caron (2012) published an exhaustive description based on all 114 of the known fossil specimens; they discovered new and unexpected characteristics {{which|date=December 2016}} that they recognized as primitive features of the first chordate animals. On the basis of these findings, they constructed a new scenario for chordate evolution.<ref name="conway2012">Conway Morris, S.; Caron, J. B. (2012) "''Pikaia gracilens'' Walcott, a stem-group chordate from the Middle Cambrian of British Columbia". ''Biological Reviews'' 87: 480-512.</ref> Subsequently, Mallatt and Holland reconsidered Conway Morris and Caron's description, and concluded that many of the newly recognized characters are unique, already-divergent specializations that would not be helpful for establishing ''Pikaia'' as a basal chordate.<ref name="mallatt2013"> Mallatt, J.; Holland, N. D. (2013) "''Pikaia gracilens'' Walcott: stem chordate, or already specialized in the Cambrian?" ''Journal of Experimental Zoology'', Part B, Molecular and Developmental Evolution 320B: 247-271.</ref>

==Discovery==
[[Image:Burgess scale2.png|thumb|left|Scale diagram of various Burgess Shale invertebrates, ''P. gracilens'' in yellow]]
''P. gracilens'' was discovered by [[Charles Doolittle Walcott|Charles Walcott]] and first described by him in 1911. It was named after [[Pika Peak]], a mountain in [[Alberta]], [[Canada]]. Based on the obvious and regular segmentation of the body, Walcott classified it as a [[Polychaeta|polychaete]] worm.

[[Image:Pikaia Smithsonian.JPG|thumb|right|Fossil specimen on display at the [[National Museum of Natural History|Smithsonian]] in Washington, D.C.]]

During his re-examination of the Burgess Shale fauna in 1979, [[Paleontology|paleontologist]] [[Simon Conway Morris]] placed ''P. gracilens'' among the [[chordate]]s, making it perhaps the oldest known ancestor of modern [[vertebrate]]s. He did this because it seemed to have a very primitive, proto-[[notochord]], however, the status of ''Pikaia'' as a chordate is not universally accepted; its preservational mode suggests that it had [[cuticle]], which is uncharacteristic of the vertebrates <ref name=Butterfield1990>{{citation
| last = Butterfield | first = N. J.
| year = 1990
| title = Organic preservation of non-mineralizing organisms and the taphonomy of the Burgess Shale
| journal = Paleobiology
| volume = 16
| issue = 3
| pages = 272–286
| jstor = 2400788
| publisher = Paleontological Society}}</ref> (although characteristic of other [[cephalochordate]]s); further, its tentacles are unknown from other vertebrate lineages.<ref name=Butterfield1990/> The presence of earlier chordates among the [[Chengjiang]], including ''[[Haikouichthys]]'' and ''[[Myllokunmingia]]'', appears to show that cuticle is not necessary for preservation, overruling the taphonomic argument,<ref name=Morris2008>{{citation
| last = Conway Morris | first = S.
| year = 2008
| title = A Redescription of a Rare Chordate,'' Metaspriggina walcotti'' Simonetta and Insom, from the Burgess Shale (Middle Cambrian), British Columbia, Canada
| journal = Journal of Paleontology
| volume = 82
| issue = 2
| pages = 424–430
| doi=10.1666/06-130.1
| url=http://jpaleontol.geoscienceworld.org/cgi/content/extract/82/2/424 | accessdate=2009-04-28
}}</ref> but the presence of tentacles remains intriguing, and the organism cannot be assigned conclusively, even to the vertebrate [[stem group]]. Its anatomy closely resembles the modern creature ''[[Branchiostoma]]''.<ref name=Donoghue2005>{{citation
| last1 = Donoghue | first1 = P. C. J.
| last2 = Purnell | first2 = M. A.
| year = 2005
| title = Genome duplication, extinction and vertebrate evolution
| journal = Trends in Ecology & Evolution
| volume = 20
| issue = 6
| pages = 312–319
| url = http://palaeo.gly.bris.ac.uk/Publs/donoghue/Donoghue_and_Purnell_2005.pdf
|format=PDF
| doi = 10.1016/j.tree.2005.04.008
| pmid = 16701387}}</ref>

==Evolution==
Much debate on whether ''Pikaia'' is a vertebrate ancestor, its worm-like appearance notwithstanding, exists in scientific circles. It looks like a worm that has been flattened sideways. The fossils compressed within the Burgess Shale show chordate features such as traces of an elongate [[notochord]], dorsal nerve cord, and blocks of muscles ([[myotome]]s) down either side of the body – all critical features for the evolution of the vertebrates.

The notochord, a flexible rod-like structure that runs along the back of the animal, lengthens and stiffens the body so that it can be flexed from side to side by the muscle blocks for swimming. In the fish and all subsequent vertebrates, the notochord forms the backbone (or vertebral column). The backbone strengthens the body, supports strut-like limbs, and protects the vital dorsal nerve cord, while at the same time allowing the body to bend.

A ''Pikaia'' lookalike, the [[lancelet]] ''[[Branchiostoma]]'', still exists today. With a notochord and paired muscle blocks, the lancelet and ''Pikaia'' belong to the chordate group of animals from which the vertebrates descended. Molecular studies have refuted earlier hypotheses that lancelets might be the closest living relative to the vertebrates, instead favoring [[tunicates]] in this position.<ref>Delsuc et al. (2008): "Additional Molecular Support for the New Chordate Phylogeny". - ''Genesis'', 46(11): 592-604 [http://hal.archives-ouvertes.fr/docs/00/33/84/11/PDF/Delsuc-Genesis08-HAL.pdf PDF]</ref> Other living and fossil groups, such as [[acorn worms]] and [[graptolite]], are more primitive; called the [[hemichordates]], they have only a notochord-like structure at an early stage of their lives.

The presence of a creature as complex as ''Pikaia'' some 530 million years ago reinforces the controversial view that the diversification of life must have extended back well before [[Cambrian]] times - perhaps deep into the [[Precambrian]].<ref name=autogenerated1 />

===Development of the head===
The first sign of head development, [[cephalization]], is seen in chordates such as ''Pikaia'' and ''Branchiostoma''. It is thought that development of a head structure resulted from a long body shape, a swimming habit, and a mouth at the end that came into contact with the environment first, as the animal swam forward. The search for food required ways of continually testing what lay ahead so it is thought that anatomical structures for seeing, feeling, and smelling developed around the mouth. The information these structures gathered was processed by a swelling of the nerve cord (''efflorescence'') – the precursor of the brain. Altogether, these front-end structures formed the distinct part of the vertebrate body known as the head.<ref name=autogenerated1 />

== See also ==
{{portal|Paleontology}}
*''[[Metaspriggina]]''

== References ==
{{reflist}}

== Further reading ==
* Bishop, A., Woolley, A. and Hamilton, W. (1999) ''Minerals, Rocks and Fossils''. London: Phillip's
* Lacalli T. (2012) "The Middle Cambrian fossil ''Pikaia'' and the evolution of chordate swimming". ''[[EvoDevo (journal)|EvoDevo]]'' '''3''': 12. {{doi|10.1186/2041-9139-3-12}}
*[[Simon Conway Morris|Conway Morris, Simon]]. 1998. ''The Crucible of Creation: The Burgess Shale and the Rise of Animals''. Oxford University Press, New York, New York.
*[[Stephen Jay Gould|Gould, Stephen Jay]]. 1989. ''Wonderful Life: The [[Burgess Shale]] and the Nature of History''. W. W. Norton, New York, NY.
* Norman, D. (1994) ''Prehistoric Life: the Rise of the Vertebrates'', London: Boxtree
* Sheldon, P., Palmer D., Spicer, B. (2001). ''Fossils and the History of Life''. Aberystwyth: Cambrian Printers/The Open University. p. 41-42.

== External links ==
{{commons category|Pikaia}}
* {{Burgess Shale species | 101}}
* [http://www.ucm.es/info/zoo/Vertebrados/JADiaz/evolucion.htm La evolución de las especies: ¿por qué sobrevivió ''Pikaia''?] (Spanish)
* [http://facstaff.gpc.edu/~pgore/geology/geo102/burgess/burgess.htm Fossils of the Burgess Shale - Middle Cambrian]
* [http://onlinelibrary.wiley.com/doi/10.1111/j.1469-185X.2012.00220.x/abstract ''Pikaia gracilens'' Walcott, a stem-group chordate from the Middle Cambrian of British Columbia]

[[Category:Cephalochordata]]
[[Category:Cambrian chordates]]
[[Category:Burgess Shale animals]]
[[Category:Transitional fossils]]
[[Category:Fossil taxa described in 1911]]
[[Category:Taxa named by Charles Doolittle Walcott]]

Hadrocodium

2017-10-25T20:39:43Z

Wikkler: /* Family Tree */ Neither is Mammaliaformes.

{{Automatic taxobox
| fossil_range = [[Early Jurassic]], {{Fossil range|195}}
| image =
| image_caption =
| taxon = Hadrocodium
| authority = {{Harvnb|Luo|Crompton|Sun|2001}}
| subdivision_ranks = Species
| subdivision =
* '''''H. wui''''' {{Harvnb|Luo|Crompton|Sun|2001}}
}}

'''''Hadrocodium wui''''' (''hadro'' from Greek ἁδρός/''hadros'', "large, heavy, fullness";<ref name="Luo-etal-2001-ref1">{{Harvnb|Luo|Crompton|Sun|2001| loc=Note 1}}</ref> Latin: ''codium'', from Greek κώδεια/''kodeia'', "head [of a plant]" <ref>{{Harvnb|Liddell|Scott|1940}}</ref> (alluding to its enlarged cranial cavity);<ref>{{cite journal |last1=Luo |first1=Z.-X. |title=A New Mammaliaform from the Early Jurassic and Evolution of Mammalian Characteristics |journal=Science |volume=292 |issue=5521 |pages=1535–40 |year=2001 |pmid=11375489 |doi=10.1126/science.1058476 }}</ref> and ''wui'', the Latinized version of discoverer Xiao-Chun Wu's name<ref name="NG-2001"/>) is an extinct [[Mammaliaformes|mammaliaform]] that lived during the [[Sinemurian]] stage of the [[Early Jurassic]] approximately {{Mya|195}}<ref name="Luo-etal-2001-abst">{{harvnb|Luo|Crompton|Sun|2001| loc=Abstract}}</ref> in the [[Lower Lufeng Series|Lufeng basin]] in what is now the [[Yunnan]] province in south-western China<ref name="NG-2001">{{Harvnb|Parsell|2001}}</ref>
({{Coord|25.2|102.1|display=inline}}, paleocoordinates {{Coord|34.3|104.9|display=inline}}).<ref>{{Paleodb|collection|11805|Hei Koa Peng, Lufeng (CUP, IVPP) (Jurassic of China)}}. Retrieved April 2013.</ref>

The fossil of this mouse-like, paper-clip sized animal was discovered in 1985 but was then interpreted as a juvenile [[morganucodon]]tid.<ref name="Luo-etal-2001-ref1" /> ''Hadrocodium'' remained [[Undescribed taxon|undescribed]] until 2001; since then its large brain and advanced ear structure<ref name="CNN-2011">{{Harvnb|CNN|2001}}</ref> have greatly influenced the interpretation of the earliest stages of [[Evolution of mammals|mammalian evolution]], as these mammalian characters could previously be traced only to some {{Mya|150|million years ago}}.<ref name="Carnegie-2001">{{Harvnb|CMNH|2001}}</ref> ''Hadrocodium'' is known only from a skull, but the body is estimated to have been a mere {{Convert|3.2|cm|abbr=on}} in length and about {{Convert|2|g|abbr=on}} in mass, making it one of the smallest mammals ever.

==Features==
''Hadrocodium'' might have been the [[Evolution of mammalian auditory ossicles|first animal]] to have a nearly fully mammalian middle ear. It is the earliest known example of several features possessed only by mammals,<ref>[http://palaeos.com/vertebrates/mammaliformes/mammaliformes3.html Symmetrodonta - Palaeos]</ref> including the [[Middle ear|middle-ear]] structure characteristic of modern mammals and a relatively large [[brain]] cavity.<ref name="CNN-2011" /> These features had been considered limited to the [[crown group]] mammals, who emerged in the [[Middle Jurassic]]; the discovery of ''Hadrocodium'' suggests that these attributes appeared earlier (45 million years earlier) than previously thought.

Whether ''Hadrocodium'' was [[warm-blooded]] or [[Poikilotherm|cold-blooded]] has not been settled, although its apparent [[Nocturnal animal|nocturnal]] features would seem to place it in the endotherm group.

==Family Tree==
{{main article|Evolution of mammals}}

{{clade| style=font-size:90%;line-height:100%
|label1=[[Cynodontia]]
|1={{clade
|1={{clade
|1=''[[Dvinia]]''
|2=[[Procynosuchidae]]
}}
|label2=[[Epicynodontia]]
|2={{clade
|1=''[[Thrinaxodon]]''
|label2=[[Eucynodontia]]
|2={{clade
|1={{clade
|1=''[[Cynognathus]]''
|2={{clade
|1=[[Tritylodontidae]]
|2=[[Traversodontidae]]}} }}
|label2=[[Probainognathia]]
|2={{clade
|1={{clade
|1=[[Tritheledontidae]]
|2=[[Chiniquodontidae]]}}
|2={{clade
|1=''[[Prozostrodon]]''
|label2=[[Mammaliaformes]]
|2={{clade
|1=[[Morganucodon]]tidae
|2={{clade
|1=[[Docodonta]]
|2={{clade
|1='''''Hadrocodium'''''
|2={{clade
|1=[[Kuehneotheriidae]]
|2=[[crown group]] [[Mammals]]
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}

;Phylogeny <ref>{{cite journal|last1=Close|first1=Roger A.|last2=Friedman|first2=Matt|last3=Lloyd|first3=Graeme T.|last4=Benson|first4=Roger BJ|year=2015|title=Evidence for a mid-Jurassic adaptive radiation in mammals.|journal=Current Biology|volume=25|issue=16|pages=2137–2142|doi=10.1016/j.cub.2015.06.047|pmid=26190074}}</ref>
{{clade |style=font-size:95%;line-height:80%;
|label1=[[Mammaliaformes]] 
|1={{clade
|1= ''[[Adelobasileus]]''
|2={{clade
|1= ''[[Sinoconodon]]''
|2={{clade
|1={{clade
|1= ''[[Morganucodon]]''
|2= ''[[Megazostrodon]]''
}}
|2={{clade
|1= [[Haramiyida]]
|2={{clade
|1={{clade
|1= ''[[Haldanodon]]''
|2= ''[[Castorocauda]]''
}}
|2={{clade
|1= '''''Hadrocodium'''''
|2= [[Mammalia]]
}} }} }} }} }} }} }}

==See also==
*[[Evolution of mammalian auditory ossicles]]

==References==
{{Reflist|30em}}

==Bibliography==
{{Refbegin|30em}}
* {{Cite web
| title = Hadrocodium wui
| publisher = Carnegie Museum of Natural History | year = 2001
| url = http://www.carnegiemnh.org/vp/hadrocodium.html | accessdate = April 2013
| ref = {{Harvid|CMNH|2001}}}}
* {{Cite web
| title = Tiny creature may be ancestor of all mammals
| publisher = CNN | date = May 24, 2001
| url = http://archives.cnn.com/2001/TECH/science/05/24/tiny.ancestor/index.html | accessdate = April 2013
| ref = {{Harvid|CNN|2001}}}}
* {{Cite web
| last1 = Liddell | first1 = Henry George | authorlink1 = Henry Liddell
| last2 = Scott | first2 = Robert | authorlink2 = Robert Scott (philologist)
| title = A Greek–English Lexicon: κώδεια
| year = 1940 | publisher = Perseus Digital Library
| url = http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dkw%2Fdeia | accessdate = April 2013
| ref = harv}}
* {{Cite journal
| last1 = Luo | first1 = Zhe-Xi
| last2 = Crompton | first2 = Alfred W.
| last3 = Sun | first3 = Ai-Lin
| title = A New Mammaliaform from the Early Jurassic and Evolution of Mammalian Characteristics
| year = 2001 | journal = Science | volume = 292 | issue = 5521 | pages = 1535–1540
| url = http://www.bi.ku.dk/dna/course/papers/J1.luo.pdf
| doi = 10.1126/science.1058476 | pmid = 11375489 | bibcode = 2001Sci...292.1535L | ref = harv}}
* {{Cite news
| last = Parsell | first = D.L.
| title = Tiny Fossil From Early Jurassic Fills New Niche in Mammal Evolution
| date = May 24, 2001 | publisher = National Geographic News
| url = http://news.nationalgeographic.com/news/2001/05/0524_paperclipmammal.html | accessdate = April 2013
| ref = harv}}
* {{Cite journal
| last1 = Rowe | first1 = Timothy
| last2 = Macrini | first2 = Thomas E
| last3 = Luo | first3 = Zhe-Xi
| title = Fossil Evidence on Origin of the Mammalian Brain
|date=May 2011 | journal = Science | volume = 332 | issue = 955
| doi = 10.1126/science.1203117 | bibcode = 2011Sci...332..955R | pmid=21596988 | pages=955–7}} [http://www.sciencemag.org/content/suppl/2011/05/18/332.6032.955.DC1/Rowe-SOM.pdf (Supporting online material)]
{{Refend}}

==External links==
{{Wikispecies|Hadrocodium}}
* {{Cite web
| last1 = Rowe | first1 = Timothy
| last2 = Macrini | first2 = Thomas E
| last3 = Luo | first3 = Zhe-Xi
| title = Hadrocodium wui (On-line)
| publisher = Digital Morphology at the University of Texas | year = 2006
| url = http://www.digimorph.org/specimens/Hadrocodium_wui/
| accessdate = April 2013
}} — 3D models from CT scans of the original fossil
* [http://www.palaeocritti.com/by-group/mammaliaformes/hadrocodium Palaeocritti - a guide to prehistoric animals]

{{Mammaliaformes|B.}}

[[Category:Mammaliaformes]]
[[Category:Jurassic mammals]]
[[Category:Jurassic mammals of Asia]]

{{paleo-mammal-stub}}

Hadrocodium

2017-10-25T20:39:21Z

Wikkler: /* Family Tree */ Mammalia's not a genus.

{{Automatic taxobox
| fossil_range = [[Early Jurassic]], {{Fossil range|195}}
| image =
| image_caption =
| taxon = Hadrocodium
| authority = {{Harvnb|Luo|Crompton|Sun|2001}}
| subdivision_ranks = Species
| subdivision =
* '''''H. wui''''' {{Harvnb|Luo|Crompton|Sun|2001}}
}}

'''''Hadrocodium wui''''' (''hadro'' from Greek ἁδρός/''hadros'', "large, heavy, fullness";<ref name="Luo-etal-2001-ref1">{{Harvnb|Luo|Crompton|Sun|2001| loc=Note 1}}</ref> Latin: ''codium'', from Greek κώδεια/''kodeia'', "head [of a plant]" <ref>{{Harvnb|Liddell|Scott|1940}}</ref> (alluding to its enlarged cranial cavity);<ref>{{cite journal |last1=Luo |first1=Z.-X. |title=A New Mammaliaform from the Early Jurassic and Evolution of Mammalian Characteristics |journal=Science |volume=292 |issue=5521 |pages=1535–40 |year=2001 |pmid=11375489 |doi=10.1126/science.1058476 }}</ref> and ''wui'', the Latinized version of discoverer Xiao-Chun Wu's name<ref name="NG-2001"/>) is an extinct [[Mammaliaformes|mammaliaform]] that lived during the [[Sinemurian]] stage of the [[Early Jurassic]] approximately {{Mya|195}}<ref name="Luo-etal-2001-abst">{{harvnb|Luo|Crompton|Sun|2001| loc=Abstract}}</ref> in the [[Lower Lufeng Series|Lufeng basin]] in what is now the [[Yunnan]] province in south-western China<ref name="NG-2001">{{Harvnb|Parsell|2001}}</ref>
({{Coord|25.2|102.1|display=inline}}, paleocoordinates {{Coord|34.3|104.9|display=inline}}).<ref>{{Paleodb|collection|11805|Hei Koa Peng, Lufeng (CUP, IVPP) (Jurassic of China)}}. Retrieved April 2013.</ref>

The fossil of this mouse-like, paper-clip sized animal was discovered in 1985 but was then interpreted as a juvenile [[morganucodon]]tid.<ref name="Luo-etal-2001-ref1" /> ''Hadrocodium'' remained [[Undescribed taxon|undescribed]] until 2001; since then its large brain and advanced ear structure<ref name="CNN-2011">{{Harvnb|CNN|2001}}</ref> have greatly influenced the interpretation of the earliest stages of [[Evolution of mammals|mammalian evolution]], as these mammalian characters could previously be traced only to some {{Mya|150|million years ago}}.<ref name="Carnegie-2001">{{Harvnb|CMNH|2001}}</ref> ''Hadrocodium'' is known only from a skull, but the body is estimated to have been a mere {{Convert|3.2|cm|abbr=on}} in length and about {{Convert|2|g|abbr=on}} in mass, making it one of the smallest mammals ever.

==Features==
''Hadrocodium'' might have been the [[Evolution of mammalian auditory ossicles|first animal]] to have a nearly fully mammalian middle ear. It is the earliest known example of several features possessed only by mammals,<ref>[http://palaeos.com/vertebrates/mammaliformes/mammaliformes3.html Symmetrodonta - Palaeos]</ref> including the [[Middle ear|middle-ear]] structure characteristic of modern mammals and a relatively large [[brain]] cavity.<ref name="CNN-2011" /> These features had been considered limited to the [[crown group]] mammals, who emerged in the [[Middle Jurassic]]; the discovery of ''Hadrocodium'' suggests that these attributes appeared earlier (45 million years earlier) than previously thought.

Whether ''Hadrocodium'' was [[warm-blooded]] or [[Poikilotherm|cold-blooded]] has not been settled, although its apparent [[Nocturnal animal|nocturnal]] features would seem to place it in the endotherm group.

==Family Tree==
{{main article|Evolution of mammals}}

{{clade| style=font-size:90%;line-height:100%
|label1=[[Cynodontia]]
|1={{clade
|1={{clade
|1=''[[Dvinia]]''
|2=[[Procynosuchidae]]
}}
|label2=[[Epicynodontia]]
|2={{clade
|1=''[[Thrinaxodon]]''
|label2=[[Eucynodontia]]
|2={{clade
|1={{clade
|1=''[[Cynognathus]]''
|2={{clade
|1=[[Tritylodontidae]]
|2=[[Traversodontidae]]}} }}
|label2=[[Probainognathia]]
|2={{clade
|1={{clade
|1=[[Tritheledontidae]]
|2=[[Chiniquodontidae]]}}
|2={{clade
|1=''[[Prozostrodon]]''
|label2=[[Mammaliaformes]]
|2={{clade
|1=[[Morganucodon]]tidae
|2={{clade
|1=[[Docodonta]]
|2={{clade
|1='''''Hadrocodium'''''
|2={{clade
|1=[[Kuehneotheriidae]]
|2=[[crown group]] [[Mammals]]
}}
}}
}}
}}
}}
}}
}}
}}
}}
}}

;Phylogeny <ref>{{cite journal|last1=Close|first1=Roger A.|last2=Friedman|first2=Matt|last3=Lloyd|first3=Graeme T.|last4=Benson|first4=Roger BJ|year=2015|title=Evidence for a mid-Jurassic adaptive radiation in mammals.|journal=Current Biology|volume=25|issue=16|pages=2137–2142|doi=10.1016/j.cub.2015.06.047|pmid=26190074}}</ref>
{{clade |style=font-size:95%;line-height:80%;
|label1=''[[Mammaliaformes]]'' 
|1={{clade
|1= ''[[Adelobasileus]]''
|2={{clade
|1= ''[[Sinoconodon]]''
|2={{clade
|1={{clade
|1= ''[[Morganucodon]]''
|2= ''[[Megazostrodon]]''
}}
|2={{clade
|1= [[Haramiyida]]
|2={{clade
|1={{clade
|1= ''[[Haldanodon]]''
|2= ''[[Castorocauda]]''
}}
|2={{clade
|1= '''''Hadrocodium'''''
|2= [[Mammalia]]
}} }} }} }} }} }} }}

==See also==
*[[Evolution of mammalian auditory ossicles]]

==References==
{{Reflist|30em}}

==Bibliography==
{{Refbegin|30em}}
* {{Cite web
| title = Hadrocodium wui
| publisher = Carnegie Museum of Natural History | year = 2001
| url = http://www.carnegiemnh.org/vp/hadrocodium.html | accessdate = April 2013
| ref = {{Harvid|CMNH|2001}}}}
* {{Cite web
| title = Tiny creature may be ancestor of all mammals
| publisher = CNN | date = May 24, 2001
| url = http://archives.cnn.com/2001/TECH/science/05/24/tiny.ancestor/index.html | accessdate = April 2013
| ref = {{Harvid|CNN|2001}}}}
* {{Cite web
| last1 = Liddell | first1 = Henry George | authorlink1 = Henry Liddell
| last2 = Scott | first2 = Robert | authorlink2 = Robert Scott (philologist)
| title = A Greek–English Lexicon: κώδεια
| year = 1940 | publisher = Perseus Digital Library
| url = http://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.04.0057%3Aentry%3Dkw%2Fdeia | accessdate = April 2013
| ref = harv}}
* {{Cite journal
| last1 = Luo | first1 = Zhe-Xi
| last2 = Crompton | first2 = Alfred W.
| last3 = Sun | first3 = Ai-Lin
| title = A New Mammaliaform from the Early Jurassic and Evolution of Mammalian Characteristics
| year = 2001 | journal = Science | volume = 292 | issue = 5521 | pages = 1535–1540
| url = http://www.bi.ku.dk/dna/course/papers/J1.luo.pdf
| doi = 10.1126/science.1058476 | pmid = 11375489 | bibcode = 2001Sci...292.1535L | ref = harv}}
* {{Cite news
| last = Parsell | first = D.L.
| title = Tiny Fossil From Early Jurassic Fills New Niche in Mammal Evolution
| date = May 24, 2001 | publisher = National Geographic News
| url = http://news.nationalgeographic.com/news/2001/05/0524_paperclipmammal.html | accessdate = April 2013
| ref = harv}}
* {{Cite journal
| last1 = Rowe | first1 = Timothy
| last2 = Macrini | first2 = Thomas E
| last3 = Luo | first3 = Zhe-Xi
| title = Fossil Evidence on Origin of the Mammalian Brain
|date=May 2011 | journal = Science | volume = 332 | issue = 955
| doi = 10.1126/science.1203117 | bibcode = 2011Sci...332..955R | pmid=21596988 | pages=955–7}} [http://www.sciencemag.org/content/suppl/2011/05/18/332.6032.955.DC1/Rowe-SOM.pdf (Supporting online material)]
{{Refend}}

==External links==
{{Wikispecies|Hadrocodium}}
* {{Cite web
| last1 = Rowe | first1 = Timothy
| last2 = Macrini | first2 = Thomas E
| last3 = Luo | first3 = Zhe-Xi
| title = Hadrocodium wui (On-line)
| publisher = Digital Morphology at the University of Texas | year = 2006
| url = http://www.digimorph.org/specimens/Hadrocodium_wui/
| accessdate = April 2013
}} — 3D models from CT scans of the original fossil
* [http://www.palaeocritti.com/by-group/mammaliaformes/hadrocodium Palaeocritti - a guide to prehistoric animals]

{{Mammaliaformes|B.}}

[[Category:Mammaliaformes]]
[[Category:Jurassic mammals]]
[[Category:Jurassic mammals of Asia]]

{{paleo-mammal-stub}}

Simoliophiidae

2017-10-18T19:38:21Z

Wikkler: No... just... no.

{{automatic taxobox
| taxon = Simoliophiidae
| fossil_range = Mesozoic
}}

Simoliophiidae is an extinct [[family (biology)|family]] of limbed [[Tethys Ocean|Tethyan]] [[Marine reptile|marine snakes]] of the order [[Squamata]].<ref>{{cite journal|last1=Hsiang|first1=Allison Y|last2=Field|first2=Daniel J|last3=Webster|first3=Timothy H|last4=Behlke|first4=Adam DB|last5=Davis|first5=Matthew B|last6=Racicot|first6=Rachel A|last7=Gauthier|first7=Jacques A|title=The origin of snakes: revealing the ecology, behavior, and evolutionary history of early snakes using genomics, phenomics, and the fossil record|journal=BMC Evolutionary Biology|date=20 May 2015|volume=15|issue=1|doi=10.1186/s12862-015-0358-5}}</ref> The simoliophiidae occurred in the [[Cretaceous]] period.

== References ==
{{reflist}}

{{paleo-reptile-stub}}
{{snake-stub}}

[[Category:Snakes]]
[[Category:Extinct reptiles]]
[[Category:Marine reptiles]]

Template:Ornithischia

2017-10-16T21:48:21Z

Wikkler:

{{Navbox with collapsible groups
|name = Ornithodira
|state = {{{state|expanded}}}
|selected = {{{1|}}}
|title = [[Avemetatarsalia]]ns ([[Dinosauromorpha]] and relatives)
|bodyclass = hlist

|above =
* Kingdom: [[Animal]]ia
* Phylum: [[Chordate|Chordata]]
* Class: [[Reptile|Sauropsida]]
* ''Clade'': [[Archosauria]]
* ''Clade'': [[Avemetatarsalia]]
** [[Dinosauromorpha]]
** [[Pterosauromorpha]]

|group1 = [[Avemetatarsalia]]
|abbr1 = A.
|image1 = [[File:Scleromochlus_BW.jpg|140px|''Scleromochlus taylori'']]
|list1 =
{{Navbox|subgroup
|group1 = [[Avemetatarsalia]]
|list1 =
{{Navbox|subgroup
|list1 =
* {{extinct}}''[[Scleromochlus]]''
|group2 = {{extinct}}[[Aphanosauria]]
|list2 =
* ''[[Dongusuchus]]''
* ''[[Spondylosoma]]''
* ''[[Teleocrater]]''
* ''[[Yarasuchus]]''
|group3 = [[Ornithodira]]
|list3 =
* {{uu|'''[[Dinosauromorpha]]'''}}
** see below
* {{extinct}}{{uu|'''[[Pterosauromorpha]]'''}}
** see below
}}
}}

|group2 = [[Dinosauromorpha]]
|abbr2 = D.
|image2 = [[File:Silesaurus1.jpg|140px|''Silesaurus opolensis'']]
|list2 =
{{Navbox|subgroup
|group1 = {{extinct}}[[Lagerpetidae]]
|list1 =
* ''[[Dromomeron]]''
* ''[[Lagerpeton]]''
* ''[[Ixalerpeton]]''
|group2 = [[Dinosauriformes]]
|list2 =
{{Navbox|subgroup
|list1 =
* ''[[Marasuchus]]''
* ''[[Lagosuchus]]''
* ''[[Saltopus]]''
* ''[[Nyasasaurus]]''?
|group2 = {{extinct}}[[Silesauridae]]
|list2 =
* ''[[Agnosphitys]]''
* ''[[Asilisaurus]]''
* ''[[Diodorus (genus)|Diodorus]]''
* ''[[Eucoelophysis]]''
* ''[[Ignotosaurus]]''
* ''[[Lewisuchus]]''
* ''[[Lutungutali]]''
* ''[[Pisanosaurus]]''
* ''[[Pseudolagosuchus]]''
* ''[[Silesaurus]]''
* ''[[Sacisaurus]]''
* ''[[Technosaurus]]''?
|group3 = {{uu|[[Dinosauria]]}} ([[Dinosaur]]s)
|list3 =
{{Navbox|subgroup
|list1 =
* {{extinct}}{{uu|'''[[Ornithischia]]'''}}
** see below
|group2 = {{uu|[[Saurischia]]}}
|list2 = {{Navbox|subgroup
| group1 = {{extinct}}Basal
| list1 =
* ''[[Alwalkeria]]''
* ''[[Teyuwasu]]''?
| group2 = [[Eusaurischia]]
| list2 =
* {{extinct}}''[[Eoraptor]]''
* {{extinct}}{{uu|'''[[Sauropodomorpha]]'''}}
** see below
* {{uu|'''[[Theropoda]]'''}}
** see below
}}
}}
}}
}}
|group3 = {{extinct}}[[Pterosauromorpha]]
|abbr3 = P.
|image3 = [[File:Eudimorphodon_BW.jpg|140px|''Eudimorphodon ranzii'']]
|list3 =
{{Navbox|subgroup
|group1 = [[Scleromochlidae]]
|list1 =
* ''[[Scleromochlus]]''
|group2 = [[Pterosauria]]
|list2 =
{{Navbox|subgroup
|group1 = [[Eopterosauria]]
|list1 =
{{Navbox|subgroup
|group1 = [[Preondactylia]]
|list1 =
* ''[[Austriadactylus]]''
* ''[[Preondactylus]]''
|group2 = [[Eudimorphodontoidea]]
|list2 =
{{Navbox|subgroup
|list1 =
* ''[[Peteinosaurus]]''
|group2 = [[Eudimorphodontidae]]
|list2 =
* ''[[Eudimorphodon]]''
}}
}}
|group2 = [[Macronychoptera]]
|list2 =
{{Navbox|subgroup
|group1 = [[Dimorphodontia]]
|list1 =
* ''[[Dimorphodon]]''
* ''[[Parapsicephalus]]''
|group2 = [[Novialoidea]]
|list2 =
{{Navbox|subgroup
|list1 =
* ''[[Campylognathoides]]''
|group2 = [[Breviquartossa]]
|list2 =
* '''[[Rhamphorhynchidae]]'''
* '''[[Pterodactylomorpha]]'''
**''[[Sordes]]''
**''[[Allkaruen]]''
**'''[[Monofenestrata]]'''
}}
}}
}}
}}

|group4 = {{extinct}}[[Ornithischia]]
|abbr4 = O.
|image4 = [[File:Eocursor_BW.jpg|140px|''Eocursor parvus'']]
|list4 =
{{Navbox|subgroup
|group1 = Basal [[ornithischia]]ns
|list1 =
* ''[[Eocursor]]''
* ''[[Fabrosaurus]]''
|group2 = [[Heterodontosauridae]]
|list2 =
{{Navbox|subgroup
|list1 =
* ''[[Echinodon]]''
* ''[[Fruitadens]]''
* ''[[Tianyulong]]''
|group2 = [[Heterodontosaurinae]]
|list2 =
* ''[[Lycorhinus]]''
* ''[[Pegomastax]]''
* ''[[Manidens]]''
* ''[[Abrictosaurus]]''
* ''[[Heterodontosaurus]]''
}}
|group3 = [[Genasauria]]
|list3 = {{Navbox|subgroup
|group1 = {{uu|[[Thyreophora]]}}
|list1 =
{{Navbox|subgroup
|list1 =
* ''[[Scutellosaurus]]''
* ''[[Emausaurus]]''
* ''[[Tatisaurus]]''
|group2 = [[Scelidosauridae]]
|list2 =
* ''[[Scelidosaurus]]''
|group3 = [[Eurypoda]]
|list3 =
* '''[[Ankylosauria]]'''
* '''[[Stegosauria]]'''
}}
|group2 = [[Neornithischia]]
|list2 =
{{Navbox|subgroup
|list1 =
* ''[[Stormbergia]]''
* ''[[Agilisaurus]]''
* ''[[Hexinlusaurus]]''
* ''[[Othnielosaurus]]''
* ''[[Kulindadromeus]]''
|group2 = [[Cerapoda]]
|list2 =
* {{uu|'''[[Ornithopoda]]'''}}
** includes [[Iguanodontia]]
* {{uu|'''[[Marginocephalia]]'''}}
** See [[Template:Marginocephalia|Marginocephalia]]
}}}}
}}

|group5 = {{extinct}}[[Sauropodomorpha]]
|abbr5 = S.
|image5 = [[File:Sellosaurus.jpg|140px|''Plateosaurus gracilis'']]
|list5 =
{{Navbox|subgroup
|group1 = [[Sauropodomorpha]]
|list1 =
{{Navbox|subgroup
|list1 =
* ''[[Pantydraco]]''
* ''[[Thecodontosaurus]]''
* ''[[Nambalia]]''
* ''[[Efraasia]]''
* ''[[Plateosauravus]]''
* ''[[Ruehleia]]''
* ''[[Buriolestes]]''
|group2 = [[Guaibasauridae]]
|list2 =
{{Navbox|subgroup
|list1 =
* ''[[Guaibasaurus]]''
|group2 = [[Saturnaliinae]]
|list2 =
* ''[[Saturnalia (genus)|Saturnalia]]''
* ''[[Chromogisaurus]]''
}}
|group3 = [[Plateosauria]]
|list3 =
{{Navbox|subgroup
|list1 =
* ''[[Sarahsaurus]]''
|group2 = [[Plateosauridae]]
|list2 =
* ''[[Euskelosaurus]]''
* ''[[Jaklapallisaurus]]''
* ''[[Plateosaurus]]''
* ''[[Unaysaurus]]''
|group3 = [[Riojasauridae]]
|list3 =
* ''[[Eucnemesaurus]]''
* ''[[Riojasaurus]]''
|group4 = [[Massospondylidae]]
|list4 =
* ''[[Adeopapposaurus]]''
* ''[[Coloradisaurus]]''
* ''[[Glacialisaurus]]''
* ''[[Leyesaurus]]''
* ''[[Lufengosaurus]]''
* ''[[Massospondylus]]''
* ''[[Pradhania]]''
|group5 = [[Anchisauria]]
|list5 =
* ''[[Anchisaurus]]''
* ''[[Aardonyx]]''
* ''[[Melanorosaurus]]''
* ''[[Sefapanosaurus]]''
* ''[[Leonerasaurus]]''
* ''[[Lessemsaurus]]''
* ''[[Antetonitrus]]''
* {{uu|'''[[Sauropoda]]'''}}
}}
}}
}}
|group6 = [[Theropoda]]
|abbr6 = T.
|image6 = [[File:Coelophysis_Jeff_Martz_(flipped).jpg|140px|''Coelophysis bauri'']]
|list6 =
{{Navbox|subgroup
|group1 = [[Theropoda]]
|list1 =
{{Navbox|subgroup
|group1 = {{extinct}}Basal
|list1 =
* ''[[Eodromaeus]]''
* ''[[Daemonosaurus]]''
|group2 = {{extinct}}[[Herrerasauridae]]
|list2 =
* ''[[Caseosaurus]]''
* ''[[Chindesaurus]]''
* ''[[Herrerasaurus]]''
* ''[[Sanjuansaurus]]''
* ''[[Staurikosaurus]]''
|group3 = [[Neotheropoda]]
|list3 =
{{Navbox|subgroup
|list1 =
* ''[[Liliensternus]]''
* ''[[Tachiraptor]]''
* ''[[Zupaysaurus]]''
|group2 = {{extinct}}[[Coelophysoidea]]
|list2 =
{{Navbox|subgroup
|list1 =
*''[[Dracoraptor]]''
*''[[Dolichosuchus]]''
*''[[Gojirasaurus]]''?
*''[[Lophostropheus]]''
*''[[Podokesaurus]]''
*''[[Sarcosaurus]]''
*''[[Tawa (dinosaur)|Tawa]]''
|group2 = {{extinct}}[[Coelophysidae]]
|list2 =
*''[[Coelophysis]]''
*''[[Camposaurus]]''
*''[[Panguraptor]]''
*''[[Procompsognathus]]''
*''[[Pterospondylus]]''
*''[[Segisaurus]]''
*''[[Lepidus (dinosaur)|Lepidus]]''
|group3 = {{extinct}}[[Dilophosauridae]]
|list3 =
* ''[[Cryolophosaurus]]''?
* ''[[Dilophosaurus]]''
* ''[[Dracovenator]]''
* ''[[Shuangbaisaurus]]''?
* ''[[Sinosaurus]]''?
|group4 = [[Averostra]]
|list4 =
* {{extinct}}{{uu|'''[[Ceratosauria]]'''}}
** see [[Template:Ceratosauria|Ceratosauria]]
* {{uu|'''[[Tetanurae]]'''}}
** includes [[bird]]s
}}}}
}}
}}
| below =
* Major clades {{uu|underlined}}
* See also categories:
** [[:Category:Archosaurs|Archosaurs]]
** [[:Category:Dinosaurs|Dinosaurs]]
** [[:Category:Pterosaurs|Pterosaurs]]
** [[:Category:Ornithodirans|Ornithodirans]]
* [[Cretaceous–Paleogene extinction event]]
}}<noinclude>
{{collapsible option}}
[[Category:Archosaur navigational boxes]]
</noinclude>

Sunda flying lemur

2017-10-15T18:46:45Z

Wikkler: A body section of the article was placed at the beginning, somehow.

{{Taxobox
| name = Sunda flying lemur<ref name=staff>{{cite journal |author1=Stafford B.J. |author2=Szalay F.S. | year = 2000 | title = Craniodental functional morphology and taxonomy of Dermopterans | journal = Journal of Mammalogy | volume = 81 | issue = | pages = 360–385 | doi = 10.1093/jmammal/81.2.360 }}</ref>
| status = LC
| status_system = iucn3.1
| status_ref = <ref name=IUCN>{{cite journal | authors = Boeadi; Steinmetz, R. | title = ''Galeopterus variegatus'' | journal = [[IUCN Red List of Threatened Species]] | volume= 2008 | page = e.T41502A10479343 | publisher = [[IUCN]] | year = 2008 | url = http://www.iucnredlist.org/details/41502/0 | doi = 10.2305/IUCN.UK.2008.RLTS.T41502A10479343.en | accessdate = 24 November 2016}}</ref>
| image = Kaguaani 02.jpg
| image_caption = ''Galeopterus variegatus''
| regnum = [[Animal]]ia
| phylum = [[Chordata]]
| classis = [[Mammal]]ia
| ordo = [[Dermoptera]]
| familia = [[Cynocephalidae]]
| genus = '''''Galeopterus'''''
| genus_authority = Thomas, 1908
| species = '''''G. variegatus'''''
| binomial = ''Galeopterus variegatus''
| binomial_authority = ([[Jean-Baptiste Audebert|Audebert]], 1799)
| range_map = Sunda Flying Lemur area.png
| range_map_caption = Sunda flying lemur range
| type_species = ''Galeopitiecus temminckii''
| type_species_authority = [[George Robert Waterhouse|Waterhouse]], 1838
}}
The '''Sunda flying lemur''' (''Galeopterus variegatus''), also known as the '''Malayan flying lemur, Sunda Colugo''' or '''Malayan colugo''', is a species of [[colugo]]. Until recently, it was thought to be one of only two species of flying lemur, the other being the [[Philippine flying lemur]] which is found only in the [[Philippines]]. The Sunda flying lemur is found throughout Southeast Asia in [[Indonesia]], [[Thailand]], [[Malaysia]], and [[Singapore]].<ref>[http://zipcodezoo.com/Animals/C/Cynocephalus_variegatus.asp Malayan flying lemur]</ref>

The Sunda flying lemur is not a [[lemur]] and does not fly. Instead, it glides as it leaps among trees. It is strictly [[arboreal]], is active at night, and feeds on soft plant parts such as young leaves, shoots, flowers, and fruits. After a 60-day gestation period, a single offspring is carried on the mother's abdomen held by a large skin membrane.<ref>{{Animal Burnie}}</ref> It is a forest-dependent species.

The head-body length of Sunda flying lemur is about {{convert|33|to|42|cm|abbr=on}}. Its tail length measures {{convert|18|to|27|cm|in|abbr=on}},<ref>{{Cite book|title = A Naturalist's Guide to the Mammals of Southeast Asia|last = Shepherd|first = Chris R.|publisher = John Beaufoy Publishing|year = 2012|isbn = 978-1-906780-71-5|location = Wiltshire, UK|pages = 17|last2 = Shepherd|first2 = Loretta Ann}}</ref> and its weight is {{convert|0.9|to|1.3|kg|lb|abbr=on}}.

The Sunda flying lemur is protected by national legislation. In addition to deforestation and loss of habitat, local subsistence hunting poses a serious threat to this animal. Competition with the [[plantain squirrel]] (''Callosciurus notatus'') represents another challenge for this species. More information is needed on population declines, but at present, the rate of the decline is not believed to merit listing in any category lower than [[Least Concern]].<ref name="IUCN" />

== Classification and evolution ==
[[image:Galeopterus varigatus Skull.jpg|thumb|Skull]]
The Sunda flying lemurs' two forms are not morphologically distinct from one another; the large form occurs on the mainland of the [[Sunda Shelf]] area and the mainland of Southeast Asia, while the dwarf form occurs in central [[Laos]] and some other adjacent islands.<ref name="staff" /> The Laos specimen is smaller (about 20%) than the other known mainland population.<ref name=rugg>{{cite journal | author = Ruggeri N. | author2 = Etterson M. | year = 1998 | title = The first record of colugo (Cynocephalus variegatus) from the Lao P.D.R. | url = | journal = Mammalia | volume = 62 | issue = | pages = 450–451 }}</ref> Despite the large and dwarf forms, four subspecies are known: ''G. v. variegatus'' ([[Java]]), ''G. v. temminckii'' ([[Sumatra]]), ''G. v. borneanus'' ([[Borneo]]), and ''G. v. peninsulae'' (Peninsular [[Malaysia]] and mainland of Southeast Asia)<ref name="staff" /> incorporating on the genetic species concept due to geographic isolation and genetic divergence. Recent molecular and morphological data provide the evidence that the mainland, Javan, and Bornean Sunda flying lemur subspecies may be recognised as three separate species in the genus ''Galeopterus''.<ref name=jan>{{cite journal | author = Janecka J.E. | author2 = Helgen K.M. | author3 = Lim N.T.L. | author4 = Baba M. | author5 = Izawa M. | author6 = Boeadi | author7 = Murphy W.J. | year = 2008 | title = Evidence for multiple species of Sunda Colugo | url = | journal = Current Biology | volume = 18 | issue = | page = 21 | doi=10.1016/j.cub.2008.09.005 | pmid=19000793}}</ref>

== Habits and lifestyle ==

Sunda flying lemurs live either solitary or in small groups that are loosely connected. They can be territorial as regards foraging and sleeping areas. They are mainly nocturnal. They are strictly arboreal and in the daytime they sleep high within dense foliage in the treetops or in holes in trees. With all four of their feet, they cling on to the trunk of a tree or the underside of branches. Climbing involves stretching out their two front legs and then bringing up their two back legs, which results in an awkward hopping. They can glide more than 100 m with minimal loss in elevation. When threatened they either climb higher up or remain motionless. These animals are quite helpless if on the forest floor.

== Behaviour and ecology ==

The Sunda flying lemur is a skillful climber, but is helpless when on the ground.<ref>Vaughan, T.A. (1986). Mammalogy. 3rd edition. Saunders College Publishing, Philadelphia.</ref> Its gliding membrane connects from the neck, extending along the limbs to the tips of the fingers, [[toe]]s and nails.<ref name=feld>Feldhamer, G.A., Drickamer, L.C., Vessey, S.H. and Merritt, J.F. (2003). Mammalogy: adaptation, diversity and ecology. 2nd edition. McGraw-Hill Companies Inc., New York.</ref> This kite-shaped skin is known as a [[patagium]], which is expanded for gliding. It can glide over a distance of 100 m with a loss of less than 10 m in elevation.<ref name="feld" /> It can maneuver and navigate while gliding, but strong rain and wind can affect its ability to glide. Gliding usually occurs in open areas or high in the canopy, especially in dense [[tropical rainforest]]. The Sunda flying lemur needs a certain distance to glide and to land to avoid injury.<ref name=byrnes08>{{cite journal |author1=Byrnes G. |author2=Norman T.-L. Lim |author3=Andrew J. Spence | year = 2008 | title = Take-off and landing kinetics of a free-ranging gliding mammal, the Malayan Colugo (Galeopterus variegatus) | url = http://rspb.royalsocietypublishing.org/content/275/1638/1007.short | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 275 | issue = 1638| pages = 1007–1013 | doi=10.1098/rspb.2007.1684 | pmid=18252673 | pmc=2600906}}</ref> The highest landing forces are experienced after short glides; longer glides lead to softer landings, due to the colugo's ability to brake its glide aerodynamically. The ability to glide increases a colugo's access to scattered food resources in the rainforest, without increasing exposure to terrestrial or arboreal predators.<ref>{{cite journal | author = Byrnes G., Libby Thomas, Norman , Lim T.-L., Andrew J. Spence | year = 2011 | title = Gliding saves time but not energy in Malayan Colugos | url = | journal = Journal of Experimental Biology | volume = 214 | issue = | pages = 2690–2696 | doi=10.1242/jeb.052993 | pmid=21795564}}</ref>

In general, the diet of the Sunda flying lemur consists mainly of leaves. It usually consumes leaves with less [[potassium]] and [[nitrogen]]-containing compounds, but with higher [[tannin]].<ref>{{cite journal |author1=Agoramoorthy G. |author2=Sha C.M. |author3=Hsu M.J. | year = 2006 | title = Population, diet and conservation of Malayan flying lemurs in altered and fragmented habitats in Singapore | url = | journal = Biodiversity and Conservation | volume = 15 | issue = | pages = 2177–2185 | doi=10.1007/s10531-004-6900-1}}</ref> It also feeds on buds,<ref name=yasu>Yasuma, S. and Andau, M. (2000). Mammals of Sabah Part-2, habitat and ecology. Tian Sing Printing Co. Sdn.Bhd, Kota Kinabalu.</ref> shoots,<ref name="fran"/> coconut flowers, durian flowers,<ref name=ket>{{cite journal |author1=Ketol B. |author2=Abdullah M.T. |author3=Tedong S. | year = 2006 | title = Short notes: Distribution records of the rare flying lemur in Kota Samarahan and Kuching Area, Sarawak | url = | journal = Sarawak Museum Journal | volume = 83 | issue = | pages = 237–241 }}</ref> fruits, and sap<ref name=lim>Lim, B.L. (1967). Observations on the food habits and ecological habitat of the Malaysian flying lemur. International Zoo Yearbook. Scotland, Aberdeen University Press. 7: 196-197.</ref> from selected tree species. It also feeds on insects in [[Sarawak]], Malaysian Borneo.<ref>Davis D D. (1958). Mammals of the Kelabit Plateau Northern Sarawak. Chicago: Chicago Natural History Museum.</ref> The selected food sources depend on the localities, habitat, vegetation types, and availability.<ref>Sih, A. (1993). Effects of ecological interactions on forager diets: Competition, predation risk, parasitism and prey behaviour. In diet selection: An interdisciplinary approach to foraging behaviour (Hughes, R.N. eds). Blackwell scientific publications. Cambridge University Press, England.</ref>

The Sunda flying lemur mainly forages in tree canopies. It may forage on several different tree species in a single night,<ref>Wischusen, E.W. (1990). The foraging ecology and natural history of the Philippine flying lemur (Cynocephalus volans). PhD Thesis. Cornell University, Ithaca, New York.</ref><ref>{{cite journal |author1=Wischusen E.W. |author2=Richmond M.E. | year = 1998 | title = Foraging ecology of the Philippine Flying Lemur (Cynocephalus volans) | url = | journal = Journal of Mammalogy | volume = 79 | issue = 4| pages = 1288–1295 | doi=10.2307/1383020}}</ref> or on a single species. It can also be seen licking tree bark of selected tree species to obtain water, nutrients, salts, and minerals.<ref>Lim, N.T-L. (2007). Colugo: The flying lemur of South-East Asia. Draco Publishing and Distribution Pte Ltd., Singapore.</ref> Though the Sunda flying lemur has been reported to also occur in gardens and plantations, the species resides in forests primarily.<ref name=lim2>{{cite journal | author = Lim N.T-L. | author2 = Giam X. | author3 = Byrnes G. | author4 = Clements G.R. | year = 2013 | title = Occurrence of the Sunda colugo (Galeopterus variegatus) in the tropical forests of Singapore: A Bayesian approach | url = | journal = Mammalian Biology | volume = 78 | issue = | pages = 63–67 | doi=10.1016/j.mambio.2012.06.008}}</ref>

== Distributions and habitats ==

The Sunda flying lemur is widely distributed throughout Southeast Asia, ranging from the Sunda Shelf mainland to other islands – Northern Laos,<ref name="rugg" /> [[Cambodia]], [[Vietnam]], [[Thailand]], Malaysia (Peninsular, [[Sabah]] and [[Sarawak]]), [[Singapore]], [[Brunei]], [[Indonesia]] ([[Kalimantan]], [[Sumatera]], [[Bali]], Java),<ref name="staff" /><ref>Corbet, G.B. and Hill, J.E. (1992). The mammals of the Indomalayan region: A systematic review. Natural History Museum Publications. Oxford University Press, Oxford</ref> and many adjacent islands.<ref name=fran>Francis, C.M. (2008). A field guide to the mammals of south-east Asia. New Holland Publishers (UK) Ltd., London.</ref> Conversely, the [[Philippine flying lemur]] (''C. volans'') is confined to the southern parts of the [[Philippines]] only.<ref name="staff" />

The Sunda flying lemur is adapted to many different vegetation types, including gardens, primary and secondary forest,<ref>Lekagul, B. and McNeely, J.A. (1977). Mammals of Thailand. Kurusapha Ladprao Press, Bangrak (Bangkok).</ref> [[rubber]] and [[coconut]] plantations,<ref>Hill, J.E. (1993). Flying lemurs (in encyclopedia of animals). Weldon Owen Pty Limited, Singapore.</ref> fruit orchards (dusun),<ref name="ket" /> [[mangrove swamp]]s,<ref name="yasu" /> lowlands and upland forests,<ref name="feld" /><ref>Payne, J., Francis, C.M. and Phillipps, K. (1985). A field guide to the mammals of Borneo. Sabah Society, Kota Kinabalu.</ref> tree plantations,<ref name="fran" /> lowland [[dipterocarp]] forests, and mountainous areas.<ref name="lim" /> However, not all of the mentioned habitats can sustain large colugo populations.<ref name="lim2" />
{{commons|Galeopterus variegatus}}
{{wikispecies|Galeopterus variegatus}}

==References==

{{Reflist|30em}}

== External links ==
* {{UCSC genomes|galVar1}}

==Further reading==
{{refbegin}}
*Anon, (2008). Flying lemurs mating, Bako National Park, Sarawak, Malaysia. https://www.youtube.com/watch?v=wavNc4nuVyk. Accessed date October 7, 2008.
*{{cite journal | author = Byrnes G., Libby Thomas, Norman , Lim T.-L., Andrew J. Spence | year = 2011 | title = Gliding saves time but not energy in Malayan Colugos | url = | journal = Journal of Experimental Biology | volume = 214 | issue = | pages = 2690–2696 | doi=10.1242/jeb.052993 | pmid=21795564}}
*{{cite journal | author = Chapman H.C. | year = 1902 | title = Observations upon Galeopithecus volans | url = | journal = Proceedings of the Academy of the Academy of Natural Science of Philadelphia | volume = 54 | issue = | pages = 241–254 }}
*{{cite journal |author1=Chasen F.N. |author2=Kloss C.B. | year = 1929 | title = Notes on flying lemurs (Galeopterus) | url = | journal = Bulletin of the Raffles Museum | volume = 2 | issue = | pages = 12–22 }}
*{{cite journal |author1=Dzulhelmi M.N. |author2=Abdullah M.T. | year = 2009a | title = An ethogram construction for the Malayan flying lemur (Galeopterus variegatus) in Bako National Park, Sarawak, Malaysia | url = | journal = Journal of Tropical Biology and Conservation | volume = 5 | issue = 1| pages = 31–42 }}
*{{cite journal |author1=Dzulhelmi M.N. |author2=Abdullah M.T. | year = 2009b | title = The foraging ecology of the Sunda Colugo (Galeopterus variegatus) in Bako National Park, Sarawak, Malaysia | url = | journal = Malayan Nature Journal | volume = 61 | issue = 4| pages = 285–294 }}
*{{cite journal |author1=Dzulhelmi M.N. |author2=Abdullah M.T. | year = 2010 | title = Distribution of the Sunda Colugo (Galeopterus variegatus) in Malaysia (Peninsular, Sabah and Sarawak) | url = | journal = Journal of Tropical Life Sciences Research | volume = 21 | issue = 2| pages = 69–83 }}
*Dzulhelmi, M.N., Marzuki, H. and Abdullah, M.T. (2010). Observation on the roosting selection of the Sunda Colugo (Galeopterus variegatus) in Bako National Park, Sarawak, Malaysia. Proceedings of Conference on Natural Resources in the Tropics3: Harnessing Tropical Natural Resources Through Innovations and Technologies. pp. 433–439.
*Dzulhelmi, M.N. (2011). Behavioural Ecology of the Sunda Colugo ''Galeopterus'' ''variegatus'' (Mammalia: Dermoptera) in Bako National Park, Sarawak, Malaysia . MSc. Dissertations. Universiti Malaysia Sarawak, Kota Samarahan.
*Dzulhelmi, N. (2013). Natural History of the Colugo. UKM Press: Bangi.
*Ellerman, J.R. and Morrison-Scott, T.C.S. (1955). Supplement to Chasen, F.N. (1940): A handlist of Malaysian mammals. British Museum. Tonbridge Printers Ltd., London.
*{{cite journal |author1=Henry C. |author2=Chapman M.D. | year = 1902 | title = Observation upon Galeopithecus volans | url = | journal = Proceedings of the Academy of Natural Sciences of Philadelphia | volume = 54 | issue = 1| pages = 241–254 }}
*{{cite journal |author1=Janecka J.E. |author2=Miller W. |author3=Pringle T.H. |author4=Wiens F. |author5=Zitzmann A. |author6=Helgen K.M. |author7=Springer M.S. |author8=Murphy W.J. | year = 2007 | title = Molecular and genomic data identify the closest living relative of primates | url = | journal = Science | volume = 318 | issue = 5851| pages = 792–794 | doi=10.1126/science.1147555 | pmid=17975064}}
*{{cite journal |author1=Karim C. |author2=Tuen A.A. |author3=Abdullah M.T. | year = 2004 | title = Mammals. Sarawak Bau Limestone Biodiversity | url = | journal = Sarawak Museum Journal | volume = 6 | issue = 80| pages = 221–234 }}
*Khan, M.M. (1992). Mamalia semenanjung Malaysia. Department of Wildlife and National Parks, Kuala Lumpur.
*Kool, K.M. and Nawi, Y. (1995). Catalogue of skin in Sarawak museum, Kuching, Sarawak. Universiti Malaysia Sarawak, Kota Samarahan.
*{{cite journal | author = Lewis R.E. | year = 1986 | title = A rain-forest raptor in danger | url = | journal = Oryx | volume = 20 | issue = | pages = 170–175 | doi=10.1017/s0030605300020032}}
*Lim, N.T. (2004). Autecology and a preliminary population census of the Malayan flying lemur Cynocephalus variegatus in Singapore. BSc. Final Year Project. National University of Singapor, Singapore.
*{{cite journal | author = Martin R.D. | year = 2008 | title = Colugos: obscure mammals glide into the evolutionary limelight | url = | journal = Journal of Biology | volume = 7 | issue = | page = 13 | doi=10.1186/jbiol74}}
*Maryanto, I., Anang, S.A., and Agus, P.K. (2008). Mamalia dilindungi perundang-undangan Indonesia. LIPI Press, Jakarta.
*Medway, L. (1978). The wild mammals of Malaya (Peninsular Malaysia) and Singapore. Oxford University Press, Kuala Lumpur.
*{{cite journal |author1=Mohd-Azlan J. |author2=Fauzi M.F. | year = 2006 | title = Ethnozoological survey in selected areas in Sarawak | url = | journal = Sarawak Museum Journal | volume = 83 | issue = | pages = 185–200 }}
*{{cite journal | author = Miller G.S., Jr | year = 1906 | title = The nomenclature of flying lemur | url = | journal = Proceedings of the Biological Society of Washington | volume = 19 | issue = | pages = 41–48 }}
*{{cite journal |author1=Nie W. |author2=Fu B. |author3=O'Brian P.C.M. |author4=Wang J. |author5=Su W. |author6=Tanomtong A. |author7=Volobouev V. |author8=Ferguson-Smith M.A. |author9=Yang F. | year = 2008 | title = Flying lemurs – the "flying tree-shrews"? molecular cytogenetic evidence for a Scandentia-Dermoptera sister clade | url = | journal = BMC Biology | volume = 6 | issue = | page = 18 | doi=10.1186/1741-7007-6-18}}
*{{cite journal | author = Novacek M.J. | year = 1992 | title = Fossils, topologies, missing data, and higher level phylogeny of eutherian mammals | url = | journal = Systematics Biology | volume = 41 | issue = 1| pages = 58–73 | doi=10.1093/sysbio/41.1.58}}
*{{cite journal | author = Novacek M.J. | year = 1993 | title = Mammalian phylogeny: morphology and molecules | url = | journal = Trends in Ecological and Evolution | volume = 8 | issue = 9| pages = 339–340 | doi=10.1016/0169-5347(93)90245-k}}
*Nowak, R.M. (1999). Mammals of the world. 6th edition. Johns Hopkins University Press, Baltimore.
*Parr, J.W.K., Komolphalin, K. and Wongkalasin, M. (2003). A guide to the large mammals of Thailand. Sarakadee Press, Bangkok.
*Penry, D.L. (1993). Digestive constraints on diet selection. In diet selection: An interdisciplinary approach to foraging behaviour (Hughes, R.N. eds). Blackwell Scientific Publications. Cambridge University Press, Cambridge.
*{{cite journal | author = Pettigrew J.D. | year = 1995 | title = Flying primates: crashed or crashed through?. | url = | journal = Symp. Zoological Society of London | volume = 67 | issue = | pages = 3–26 }}
*{{cite journal |author1=Schmitz J. |author2=Ohme M. |author3=Suryobroto B. |author4=Zichler H. | year = 2002 | title = The colugo (Cynocephalus variegatus), Dermoptera: the Primates' gliding sister? | url = | journal = Molecular Phylogenetics and Evolution | volume = 19 | issue = | pages = 2308–2312 | doi=10.1093/oxfordjournals.molbev.a004054}}
*{{cite journal |author1=Schmitz J. |author2=Ohme M. |author3=Zichler H. | year = 2003 | title = A novel family of tRNA-derived SINEs in the colugo and two new retrotransposable markers separating dermopterans from primates | url = | journal = Molecular Phylogenetics and Evolution | volume = 28 | issue = | pages = 341–349 | doi=10.1016/s1055-7903(03)00060-5}}
*Stephen, D.W. and Krebs, J.R. (1986). Foraging theory. Princeton University Press, England.
*{{cite journal | author = Wharton C.H. | year = 1950 | title = Notes on the life history of the flying lemur | url = | journal = Journal of Mammalogy | volume = 31 | issue = | pages = 269–273 | doi=10.2307/1375292}}
*{{cite journal |author1=Wischusen E.W. |author2=Richmond M.E. | year = 1989 | title = Techniques for capturing and marking the Philippine flying lemurs (Cynocephalus volans) | url = | journal = Malayan Nature Journal | volume = 43 | issue = | pages = 100–105 }}
{{refend}}

{{Colugo nav}}

[[Category:Mammals of Brunei]]
[[Category:Mammals of Borneo]]
[[Category:Mammals of Indonesia]]
[[Category:Mammals of Thailand]]
[[Category:Mammals of Malaysia]]
[[Category:Mammals of Singapore]]
[[Category:Mammals described in 1799]]
[[Category:Colugos]]

Opisthocomidae

2017-10-07T11:35:10Z

Wikkler: /* Taxonomy */ Opisthocomidae's not extinct. Hoatzins are extant.

{{automatic taxobox
| name = Opisthocomids
| fossil_range = [[Late Eocene]] - Recent, {{fossilrange|Late Eocene|0|earliest=54}}
| image = Hoatzins in Brazil.jpg
| image_caption = [[Hoatzin]] (''Opisthocomus hoazin'')
| display_parents = 2
| parent_authority = L'Herminier, 1837
| taxon = Opisthocomidae
| authority = [[William John Swainson|Swainson]], 1837
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
* ?†''[[Foro (bird)|Foro]]''
* ?†''[[Hoatzi]]''
* ?†''[[Onychopteryx]]''
* †''[[Hoazinoides]]''
* †''[[Namibiavis]]''
* †''[[Protoazin]]''
* †''[[Hoazinavis]]''
* ''[[Opisthocomus]]''
| synonyms =
* Foratidae Olson 1992
* Hoazinoididae Rasmussen 1997
* Onychopterygidae Cracraft 1971
}}
'''Opisthocomidae''' is a group of [[bird]]s, the only named family within the order '''Opisthocomiformes'''. The only living representative is the [[hoatzin]] (''Opisthocomus hoazin'') which lives in the [[Amazon basin|Amazon]] and the [[Orinoco delta]] in [[South America]]. Several fossil species have been identified, including one from [[Africa]]<ref>{{cite journal | author = Mayr G., Alvarenga H., Mourer-Chauviré C. | year = 2011 | title = Out of Africa: Fossils shed light on the origin of the hoatzin, an iconic Neotropic bird | url = | journal = Naturwissenschaften | volume = 98| issue = | pages = 961–6| doi = 10.1007/s00114-011-0849-1 | pmid = 21964974 }}</ref> and one from [[Europe]].<ref name=Protoazins>{{Cite journal|author=[[Gerald Mayr]] and Vanesa L. De Pietri |year=2014 |title=Earliest and first Northern Hemispheric hoatzin fossils substantiate Old World origin of a "Neotropic endemic" |journal=Naturwissenschaften |volume= 101|issue= |pages= 143–148|doi=10.1007/s00114-014-1144-8 |pmid=24441712}}</ref>

==Phylogeny==
Based on the work of Mayr & De Pietri 2014.<ref name=Protoazins />

{{clade|style=font-size:100%;line-height:80%
|label1=
|1={{Clade
|1=†''[[Namibiavis]]''
|2={{Clade
|1=†''[[Protoazin]]''
|2={{Clade
|1=†''[[Hoazinavis]]''
|2=''[[Opisthocomus]]''
}}
}}
}}
}}

==Taxonomy==
* Family '''Opisthocomidae''' Swainson 1837<ref name="mikko">[[Mikko's Phylogeny Archive]] [http://www.helsinki.fi/~mhaaramo/] {{cite web|last=Haaramo|first=Mikko|year=2007|title=''Primitive Land-bird Assemblage'' |url=http://www.helsinki.fi/~mhaaramo/metazoa/deuterostoma/chordata/archosauria/aves/primitive_landbirds_1.html |accessdate= 30 December 2015}}</ref>
** Genus ?†''[[Foro (bird)|Foro]]'' Olson 1992 (mid-Eocene, USA) - cuculiform?
*** Species †''[[Foro panarium]]'' Olson 1992
** Genus ?†''[[Hoatzi]]'' Thomas 1996 (Eocene, Argentina)
*** Species †''[[Hoatzi panarium]]'' Thomas 1996
** Genus ?†''[[Onychopteryx]]'' Cracraft 1971 (Early Eocene of Argentina) – falconid? A ''[[nomen dubium]]''
*** Species †''[[Onychopteryx simpsoni]]'' Cracraft 1971
** Genus †''[[Protoazin]]'' Mayr & De Pietri 2014 (late Eocene of France)
*** Species †''[[Protoazin parisiensis]]'' Mayr & De Pietri 2014
** Genus †''[[Namibiavis]]'' Mourer-Chauviré 2003 (Middle Miocene of Namibia)
*** Species †''[[Namibiavis senutae]]'' Mourer-Chauviré 2003
** Genus †''[[Hoazinavis]]'' Alvarenga, Mayr & Mourer-Chauviré 2011 (Late Oligocene and Early Miocene of Brazil)
*** Species †''[[Hoazinavis lacustris]]'' Alvarenga, Mayr & Mourer-Chauviré 2011
** Genus †''[[Hoazinoides]]'' Miller 1953 (middle Miocene of Colombia)
*** Species †''[[Hoazinoides magdalenae]]'' Miller 1953
** Genus ''[[Opisthocomus]]'' Illiger 1811
*** Species ''[[Opisthocomus hoazin]]'' (Müller 1776) Illiger 1811 [''Phasianus hoazin'' Müller 1776]

==References==
{{reflist}}

{{Birds}}

[[Category:Higher-level bird taxa restricted to the Neotropics]]
[[Category:Monotypic bird families]]
[[Category:Opisthocomiformes]]
[[Category:Herbivorous vertebrates]]
[[Category:Extant Eocene first appearances]]

{{bird-stub}}

Azhdarchoidea

2017-10-04T17:29:04Z

Wikkler: Extinct markers are unnecessary for every member of an extinct clade.

{{Automatic taxobox
| name = Azhdarchoids
| fossil_range = [[Late Jurassic]] - [[Late Cretaceous]], {{fossilrange|150|66}}
| image = Quetzalcoatlus 1.JPG
| image_caption = Reconstructed skeleton of ''[[Quetzalcoatlus northropi]]''
| authority = [[L.A Nesov|Nesov]], 1984
| subdivision_ranks = Subgroups
| subdivision =
* ''[[Ornithostoma]]''
* ''[[Vectidraco]]''
* ''[[Montanazhdarcho]]''
* [[Tapejaromorpha]]
* '''Neoazhdarchia'''
** ''[[Microtuban]]''
** '''Dsungaripteromorpha'''
*** [[Dsungaripteridae]]
*** [[Thalassodromidae]]
** '''Neopterodactyloidea'''
*** [[Azhdarchidae]]
*** [[Chaoyangopteridae]]
| synonyms =
Tapejaroidea Unwin, 2003
}}
'''Azhdarchoidea''' is a group of [[pterosaurs]] within the suborder [[Pterodactyloidea]].

==Classification==
Azhdarchoidea was given a phylogenetic definition by David Unwin in 2003. Unwin defined the group as the most recent common ancestor of ''Quetzalcoatlus'' and ''Tapejara'', and all its descendants.<ref name="unwin2003">Unwin, D. M., (2003). "On the phylogeny and evolutionary history of pterosaurs." Pp. 139-190. in Buffetaut, E. & Mazin, J.-M., (eds.) (2003). ''Evolution and Palaeobiology of Pterosaurs''. Geological Society of London, Special Publications 217, London, 1-347.</ref>

There have been several competing views of azhdarchoid relationships. The first, presented by Felipe Pinheiro and colleagues in 2011, considered the tapejarids to be a monophyletic clade including the thalassodromids and chaoyangopterids.<ref name=pinheiroetal2011>Pinheiro, F.L., Fortier, D.C., Schultz, C.L., De Andrade, J.A.F.G. and Bantim, R.A.M. (in press). "New information on ''Tupandactylus imperator'', with comments on the relationships of Tapejaridae (Pterosauria)." ''Acta Palaeontologica Polonica'', in press, available online 03 Jan 2011. {{doi|10.4202/app.2010.0057}}</ref> The second, found by Naish & Martill 2006 as well as Lu ''et al.'' 2008, considered the traditional "tapejarids" to be a paraphyletic grade of primitive azhdarchoids. with true tapejarids most basal, and the thalassodromids and chaoyangopterids being successively more closely related to azhdarchids.<ref name=luetal2008>{{cite journal | last1 = Lü | first1 = J. | last2 = Unwin | first2 = D.M. | last3 = Xu | first3 = L. | last4 = Zhang | first4 = X. | year = 2008 | title = A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution | url = | journal = Naturwissenschaften | volume = 95 | issue = 9| pages = 891–897 | doi = 10.1007/s00114-008-0397-5 | pmid = 18509616 }}</ref> More recent and larger studies

All azhdarchoids which are part of the clade formed by ''Quetzalcoatlus'' and ''Tupuxuara'' are included in the group '''Neoazhdarchia''' ("new azhdarchids") as defined by Unwin in 2003. In 2003 Unwin also defined the clade '''Tapejaroidea''', which he defined as the most recent common ancestor and all descendants of ''Tapejara'', ''Quetzalcoatlus'', and ''[[Dsungaripterus]]''.<ref name="unwin2003"/>

There are competing theories of azhdarchoid phylogeny. Below is a [[cladogram]] showing the results of a [[phylogenetic]] analysis presented by Andres, Clark & Xu, 2014. This study found the a grouping of tapejarids at the base of the clade, with thalassodromids more closely related to azhdarchids and chaoyangopterids, as well as [[Dsungaripteridae|dsungaripterids]].<ref name=kryptodrakon>{{Cite journal | doi = 10.1016/j.cub.2014.03.030| title = The Earliest Pterodactyloid and the Origin of the Group| journal = Current Biology| year = 2014| last1 = Andres | first1 = B. | last2 = Clark | first2 = J. | last3 = Xu | first3 = X. | pmid=24768054 | volume=24 | pages=1011–6}}</ref>

{{clade| style=font-size:100%;line-height:100%
|label1= '''Azhdarchoidea''' 
|1={{clade
|1=[[Tapejaromorpha]][[File:Bakonydraco1DB.jpg|80 px]]
|label2= '''Neoazhdarchia''' 
|2={{clade
|2={{clade
|label1='''Neopterodactyloidea'''
|1={{clade
|1=[[Chaoyangopteridae]]
|2=[[Azhdarchidae]][[File:Quetzalcoatlus07.jpg|80 px]] }} }}
|1={{clade
|label1='''Dsungaripteromorpha'''
|1={{clade
|1=[[Thalassodromidae]]
|2=[[Dsungaripteridae]] }}
}} }} }} }}

The result of another analysis, by Vidovic and Martill, is shown below. They found tapejarids (including chaoyangopterines) to be the closest relatives of azhdarchids, followed by thalassodromids (represented by ''[[Tupuxuara]]'') and then by dsungaripterids, making Azhdarchoidea itself a small subgroup of dsungaripteroids.<ref name=aerodactylus>{{Cite journal | doi = 10.1371/journal.pone.0110646| title = ''Pterodactylus scolopaciceps'' Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: The Problem of Cryptic Pterosaur Taxa in Early Ontogeny| journal = PLoS ONE| volume = 9| issue = 10| pages = e110646| year = 2014| last1 = Vidovic | first1 = S. U. | last2 = Martill | first2 = D. M. }}</ref>

{{clade| style=font-size:100%;line-height:100%
|label1=[[Dsungaripteroidea]]
|1={{clade
|1=''[[Germanodactylus cristatus]]''[[File:Germ rhamph DB.jpg|80 px]]
|label2='''Tapejaroidea'''
|2={{clade
|1=''[[Dsungaripterus weii]]''
|label2='''Neoazhdarchia'''
|2={{clade
|1=''[[Tupuxuara]]''[[File:Tupux longDB2.jpg|80 px]]
|label2='''Azhdarchoidea'''
|2={{clade
|2=[[Azhdarchidae]][[File:Quetzalcoatlus07.jpg|80 px]]
|1={{clade
|label1=[[Tapejaromorpha]]
|1={{clade
|1=''[[Shenzhoupterus]]''
|2=[[Tapejaridae]][[File:Bakonydraco1DB.jpg|80 px]]
}} }} }} }} }} }} }}

==References==
{{reflist}}

[[Category:Azhdarchoids]]
[[Category:Tithonian first appearances]]
[[Category:Maastrichtian extinctions]]

{{pterosaur-stub}}

Notocolossus

2017-10-02T21:04:26Z

Wikkler:

{{Distinguish2|another recent titanosaur discovery in Argentina, [[Patagotitan]]}}
{{Italic title}}{{speciesbox
| name = ''Notocolossus''
| fossil_range = [[Late Cretaceous]], {{fossilrange|86}}
| image=Notocolossus.jpg
| image_caption = Skeletal restoration
| genus = Notocolossus
| parent_authority = González Riga et al., [[2016 in paleontology|2016]]
| authority = González Riga et al., 2016
| species = gonzalezparejasi
}}
'''''Notocolossus''''' is a [[genus]] of herbivorous [[lithostrotia]]n [[titanosaur]] [[sauropod]] [[dinosaur]] from late [[Cretaceous]] strata of [[Mendoza Province]], [[Argentina]].

==Discovery and naming==
[[File:Notocolossus limbs.jpg|thumb|upright|left|Limb bones]]
A fossil of a large sauropod was discovered by the Argentine paleontologist Dr. [[Bernardo Javier González Riga]] in [[Mendoza province]] <ref name="González RigaLamanna2016"/>

In 2016, the [[type species]] ''Notocolossus gonzalezparejasi'' was named and described by Bernardo J. González Riga, [[Matt Lamanna|Matthew Carl Lamanna]], [[Leonardo Daniel Ortiz David]],
[[Jorge Orlando Calvo]] and Juan P. Coria. The generic name combines the Greek words νότος, ''notos'', "south wind", and κολοσσός, ''kolossos'', "giant statue", in reference to the provenance from the [[Southern Hemisphere]] and the gigantic size of the animal. The [[specific name (zoology)|specific name]] honours [[Jorge González Parejas]], for having studied the dinosaur fossils of Mendoza province for two decades.<ref name="González RigaLamanna2016"/>

[[File:Notocolossus NT small.jpg|thumb|left|Reconstruction of ''Notocolossus gonzalesparejasi'']]

The [[holotype]], '''UNCUYO-LD 301''', was found in a layer of the [[Plottier Formation]] dating from the [[Coniacian]]-[[Santonian]], about eighty-six million years old. It consists of a partial skeleton lacking the skull. It contains an anterior back vertebra, an anterior tail vertebra, a right humerus and the upper part of a left pubic bone. Although the bones have not been discovered in articulation, they were considered to represent a single individual because they were found in close association on a surface of eight by eight metres. A second partial skeleton lacking the skull was referred to ''Notocolossus'': specimen UNCUYO-LD 302 representing a smaller individual. It contains a series of five anterior tail vertebrae and a complete right foot connected to an astragalus. It was recovered on a distance of 403 metres to the holotype from a surface of five by five metres.<ref name="González RigaLamanna2016"/>

==Description==
[[File:Notocolossus vertebrae.jpg|thumb|Vertebral bones]]
The evidence suggests that ''Notocolossus'' was among the largest titanosaurs, and therefore one of the heaviest land animals, yet discovered. Although the incompleteness of the skeleton of the new sauropod has prevented scientists from making precise estimates of its size, its humerus, or upper arm bone, is 1.76 m (5 ft 9 in) in length, which is longer than that of any other titanosaur for which this bone is known, including other giants such as ''[[Dreadnoughtus]]'', ''[[Futalognkosaurus]]'', and ''[[Paralititan]]''. If, as is likely, the body proportions of ''Notocolossus'' were comparable to those of better preserved titanosaurs, the new dinosaur may have weighed in the range of {{convert|44.9|-|75.9|t}}, most likely {{convert|60.4|t}}.<ref name="González RigaLamanna2016"/>

In 2016, several distinguishing traits of ''Notocolossus'' were determined. Nine of these were [[autapomorphies]], unique derived qualities. At the anterior back vertebra, the depression between the front edge of the side process and the lower edge of the side process is divided by two accessory ridges, one of which runs vertically while the other crosses the first horizontally. On the front side of the neural spines of the front tail vertebrae, run vertical ridges fusing at their lower ends, above the level of the front joint processes, creating a V-shaped structure. The upper inner corner of the humerus is strongly expanded, extending far beyond the inner side of the shaft. The humerus has in general a strongly expanded upper edge, 2.9 wider than the shaft diameter. On the humerus shaft, below the scar for the ''[[musculus coracobrachialis]]'', runs a diagonal ridge, from the upper and outer side to the inner and lower side. The first metatarsal has an upper width exceeding its length. The third metatarsal is relatively short with 1.2 times the length of the first metatarsal. The upper toe phalanges are 50% wider at their tops than their respective metatarsals are long. The toe claws are reduced, rough and truncated.<ref name="González RigaLamanna2016"/>

The foot or [[Pes (anatomy)|pes]] of ''Notocolossus'', a skeleton part that is not often preserved in sauropods, shows a unique build. It contains a compact, homogeneous [[metatarsus]], thought to be adapted for bearing extraordinary weight. It also presents truncated [[ungual]]s, characteristics otherwise unknown in the Sauropoda. This parallels the vertical position of the metacarpals and the finger reduction in the hand of other titanosaurs.<ref name="González RigaLamanna2016">{{cite journal|last1=González Riga|first1=Bernardo J.|last2=Lamanna|first2=Matthew C.|last3=Ortiz David|first3=Leonardo D.|last4=Calvo|first4=Jorge O.|last5=Coria|first5=Juan P.|title=A gigantic new dinosaur from Argentina and the evolution of the sauropod hind foot|journal=Scientific Reports|volume=6|year=2016|pages=19165|issn=2045-2322|doi=10.1038/srep19165|pmid=26777391|pmc=4725985}}</ref>

==Phylogeny==
''Notocolossus'' is included in the [[clade]] of the [[Titanosauria]], a group of diverse sauropods, dinosaurs with four-legged [[herbivore]]s with long necks and bodies.<ref name="González RigaLamanna2016"/> In 2016, a [[cladistic]] analysis placed ''Notocolossus'', within the Titanosauria, in the [[Lithostrotia]], in a rather basal position as a [[sister species]] of ''Dreadnoughtus''.<ref name="González RigaLamanna2016"/>

==References==
{{Portal|Dinosaurs}}
{{Reflist}}

{{DEFAULTSORT:Notocolossus}}
[[Category:Lithostrotians]]
[[Category:Dinosaur genera]]
[[Category:Late Cretaceous dinosaurs of South America]]
[[Category:Fossils of Argentina]]
[[Category:Cretaceous Argentina]]
[[Category:Fossil taxa described in 2016]]
[[Category:Vertebrates described in 2016]]
[[Category:Reptiles described in the 21st century]]
[[Category:Taxa named by Matt Lamanna]]
[[Category:Taxa named by Rodolfo Coria]]

Neornithischia

2017-09-30T20:03:54Z

Wikkler: Hypsilophodon's been removed from the ornithopod page so we need somewhere to put it. Also, extinct markers are unnecessary for every member of an extinct clade.

{{Automatic taxobox
| name = Neornithischians
| fossil_range = [[Early Jurassic]]–[[Late Cretaceous]], {{fossilrange|199.6|66}}
| image = Museum of the Rockies Dinosaur Heads.JPG
| image_caption = Skulls of neornithischian dinosaurs
| taxon = Neornithischia
| authority = Cooper, 1985
| subdivision_ranks = Subgroups
| subdivision =
*''[[Lesothosaurus]]''?
*''[[Kulindadromeus]]''
*''[[Agilisaurus]]''
*''[[Alocodon]]''
*''[[Ferganocephale]]''
*''[[Gongbusaurus]]''
*''[[Hexinlusaurus]]''
*''[[Hypsilophodon]]''
*''[[Nanosaurus]]''
*''[[Othnielia]]''
*''[[Othnielosaurus]]''
*''[[Phyllodon]]''
*''[[Stormbergia]]''
*''[[Xiaosaurus]]''
*'''[[Parksosauridae]]'''?
*'''[[Cerapoda]]'''
}}
'''Neornithischia''' ("new ornithischians") is a [[clade]] of the [[dinosaur]] order [[Ornithischia]]. They are the sister group of the [[Thyreophora]] within the clade [[Genasauria]]. Neornithischians are united by having a thicker layer of asymmetrical enamel on the inside of their lower teeth. The teeth wore unevenly with chewing and developed sharp ridges that allowed neornithischians to break down tougher plant food than other dinosaurs. Neornithischians include a variety of basal forms historical known as "hypsilophodonts", including the [[Parksosauridae]]; in addition, there are derived forms classified in the groups [[Marginocephalia]] and [[Ornithopoda]]. The former includes clades [[Pachycephalosauria]] and [[Ceratopsia]], while the latter includes ''[[Hypsilophodon]]'' and the more derived [[Iguanodontia]].

==Classification==
Neornithischia was first named by Cooper in 1985 and defined as "all genasaurians more closely related to ''[[Parasaurolophus walkeri]]'' Parks, 1922, than to ''[[Ankylosaurus magniventris]]'' Brown, 1908 or ''[[Stegosaurus stenops]]'' Marsh, 1877a".<ref name=Butler08>{{Cite journal |author1=Richard J. Butler |author2=Paul Upchurch |author3=David B. Norman |year=2008 |title=The phylogeny of the ornithischian dinosaurs |journal=Journal of Systematic Palaeontology |volume=6 |issue=1 |pages=1–40 |doi=10.1017/S1477201907002271 }}</ref> The [[cladogram]] below follows a 2011 analysis by paleontologists Richard J. Butler, Jin Liyong, Chen Jun and [[Pascal Godefroit]].<ref name=ButlerEtAl>{{Cite journal |author1=Richard J. Butler |author2=Jin Liyong |author3=Chen Jun |author4=Pascal Godefroit | year = 2011 | title = The postcranial osteology and phylogenetic position of the small ornithischian dinosaur ''Changchunsaurus parvus'' from the Quantou Formation (Cretaceous: Aptian–Cenomanian) of Jilin Province, north-eastern China | journal = Palaeontology | volume = 54 | issue = 3 | pages = 667–683 | doi = 10.1111/j.1475-4983.2011.01046.x }}</ref>

{{clade| style=font-size:85%;line-height:80%
|label1='''Neornithischia'''
|1={{clade
|1=''[[Stormbergia]]''
|2={{clade
|1=''[[Agilisaurus]]''
|2={{clade
|1=''[[Hexinlusaurus]]''
|2={{clade
|1=''[[Othnielosaurus]]''
|label2='''[[Cerapoda]]'''
|2={{clade
|1='''[[Ornithopoda]]'''
|2='''[[Marginocephalia]]''' }} }} }} }} }} }}

A recent study by [[Matthew G. Baron]],<ref>Wade, Nicholas, ''[https://www.nytimes.com/2017/03/22/science/dinosaur-family-tree.html?em_pos=medium&emc=edit_sc_20170327&nl=science-times&nl_art=5&nlid=68634180&ref=headline&te=1 Shaking Up the Dinosaur Family Tree]'', Science, March 22, 2017 </ref> [[David B. Norman]], and Paul M. Barrett recovered the Early Jurassic taxon ''Lesothosaurus diagnosticus'' from Southern Africa as the most basal known member of Neornithischia – a position previously held by ''Stormbergia dangershoeki'' (a taxon considered by the authors to be an adult form of ''Lesothosaurus'' and therefore a junior subjective synonym). However, Baron et al. go on to state that this result is only poorly supported and that future studies will be needed in order to better resolve the base of the ornithischian tree. The cladogram below follows Baron et al. 2016.<ref>{{Cite journal |author1=Matthew G. Baron |author2=David B. Norman |author3=Paul M. Barrett |date=2016 |title=Postcranial anatomy of ''Lesothosaurus diagnosticus'' (Dinosauria: Ornithischia) from the Lower Jurassic of southern Africa: implications for basal ornithischian taxonomy and systematics |journal=Zoological Journal of the Linnean Society |doi=10.1111/zoj.12434}}</ref>

{{clade| style=font-size:85%;line-height:80%
|label1='''Neornithischia'''
|1={{clade
|1=''[[Lesothosaurus]]''
|2={{clade
|1=''[[Agilisaurus]]'' [[File:Agilisaurus2.jpg|50px]]
|2={{clade
|1=''[[Hexinlusaurus]]''
|2={{clade
|1=''[[Othnielosaurus]]''
|label2='''[[Cerapoda]]'''
|2={{clade
|1='''[[Ornithopoda]]''' [[File:Parasaurolophuspic steveoc.jpg|50px]]
|2='''[[Marginocephalia]]''' [[File:Triceratops BW.jpg|50px]] }} }} }} }} }} }}

==References==
{{Reflist}}
* {{cite journal |author=Butler, R.J. |year=2005 |title=The 'fabrosaurid' ornithischian dinosaurs of the Upper Elliot Formation (Lower Jurassic) of South Africa and Lesotho |journal=[[Zoological Journal of the Linnean Society]] |volume=145 |issue=2 |page=175-18 |doi=10.1111/j.1096-3642.2005.00182.x}}
* {{cite journal |author=[[Paul Sereno|Sereno, P.C.]] |year=1986 |title=Phylogeny of the bird-hipped dinosaurs (order Ornithischia) |journal=National Geographic Research |volume=2 |issue=2 |pages=234–56}}

==External links==
* [http://www.britannica.com/eb/article-225959 Encyclopædia Britannica]
* [http://www.palaeos.com/Vertebrates/Units/320Ornithischia/500.html Palæos]

{{Portal|Dinosaurs}}
{{taxonbar}}

[[Category:Ornithischians| ]]

{{ornithischian-stub}}

Beipiaognathus

2017-09-30T16:31:02Z

Wikkler:

{{speciesbox
| image =
| image_caption =
| fossil_range = [[Early Cretaceous]], {{fossilrange|125-121|max=125|min=121}}
| genus = Beipiaognathus
| parent_authority = Hu et al., 2016
| species = jii
| authority = Hu et al., 2016
}}


'''''Beipiaognathus''''' (meaning [[Beipiao]] jaw) is a genus of [[coelurosauria]]n [[theropod]] from the [[Early Cretaceous]] [[Yixian Formation]] of [[Liaoning]], [[China]].<ref name="huetal">{{cite journal | last1 = Hu | first1 = Y | last2 = Wang | first2 = X | last3 = Huang | first3 = J | date = 2016 | title = A new species of compsognathid from the Early Cretaceous Yixian Formation of Western Liaoning, China. | journal = Journal of Geology | volume = 40 | pages = 191–196 | url = http://en.cnki.com.cn/Article_en/CJFDTotal-JSDZ201602029.htm}}</ref>

The genus was initially assigned to the [[Compsognathidae]] based on the presence of two traits: fan-shaped dorsal neural spines and a robust I-1 [[phalanx bone|phalanx]] on the hand.<ref name="huetal"/> However, it also differs from other compsognathids in several ways: the teeth are [[serration|unserrated]] and conical; the ulna is proportionally longer; the II-1 phalanx on the hand is longer and more robust; and the tail is much shorter.<ref name="huetal"/>

However, Andrea Cau has informally noted a number of points in the fossil that are indicative of it having been artificially assembled, thus rendering the specimen a phylogenetically uninformative chimaera.<ref name="cau">Cau, A (2016). [http://www.theropoda.blogspot.com/2016/08/lo-status-paleontologico-di.html Lo status (pale)ontologico di ''Beipiaognathus''] (in Italian) ''Theropoda''.</ref>

==References==
{{reflist}}

{{Portal|Paleontology|Dinosaurs}}

[[Category:Early Cretaceous dinosaurs of Asia]]
[[Category:Coelurosaurs]]
[[Category:Fossil taxa described in 2016]]
[[Category:Paleontology in Liaoning]]

Orodrominae

2017-09-30T12:54:13Z

Wikkler: It's probably not the Paleobiology "Datagase".

{{automatic taxobox
| name = Orodromines
| fossil_range = [[Cretaceous]], {{fossil range|113|75|latest=70}}
| image = Orodromeus.jpg
| image_width = 250px
| image_caption = Illustration of the type species, ''[[Orodromeus|Orodromeus makelai]]''
| authority = [[C. M. Brown|Brown]] ''et al'', 2013
| type_species = ''[[Orodromeus|Orodromeus makelai]]'' Horner & Weishampel, [[1988 in paleontology|1988]]
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
* ''[[Albertadromeus]]''
* ''[[Orodromeus]]'' ([[Type species|type]])
* ''[[Koreanosaurus]]''
* ''[[Oryctodromeus]]''
* ''[[Zephyrosaurus]]''
* ''[[Changchunsaurus]]''<ref name=citedoi17>{{Cite journal | last1 = Madzia | first1 = Daniel | last2 = Boyd | first2 = Clint A. | last3 = Mazuch | first3 = Martin | title = A basal ornithopod dinosaur from the Cenomanian of the Czech Republic | doi = 10.1080/14772019.2017.1371258 | journal = Journal of Systematic Paleontology | year = 2017 | pmid = | pmc = }}</ref>
* ''[[Haya_(dinosaur)|Haya]]''<ref name = citedoi17/>
}}

'''Orodrominae''' is a [[subfamily]] of [[Thescelosauridae|Thescelosaurid]] [[dinosaurs]] from the [[Cretaceous]] of [[North America]].<ref name=PaleobiologyDatabase>{{cite web|title=Orodrominae|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=267487|work=Paleobiology Database|accessdate=20 June 2013}}</ref>

==Distribution==
Orodromines were a [[North America]]n based group with [[fossil]]s from Canada and United States only.<ref name = PaleobiologyDatabase/> ''[[Albertadromeus]]'', as its name suggests, is only from the upper (later) part of the [[Oldman Formation]] in the [[Belly River Group]] of [[Alberta]], Canada.<ref name=citedoi13>{{Cite journal | last1 = Brown | first1 = C. M. | last2 = Evans | first2 = D. C. | last3 = Ryan | first3 = M. J. | last4 = Russell | first4 = A. P. | title = New data on the diversity and abundance of small-bodied ornithopods (Dinosauria, Ornithischia) from the Belly River Group (Campanian) of Alberta | doi = 10.1080/02724634.2013.746229 | journal = Journal of Vertebrate Paleontology | volume = 33 | issue = 3 | pages = 495 | year = 2013 | pmid = | pmc = }}</ref><ref name=Albertadromeus>{{cite web|title=Albertadromeus|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=267486|work=Paleobiology Database|accessdate=20 June 2013}}</ref> ''[[Orodromeus]]'', the type [[genus]], was widespread through [[Montana]].<ref name = PaleobiologyDatabase /> Its [[holotype]] was found at the [[Egg Mountain]] in the [[Two Medicine Formation]].<ref name=Orodromeus>{{cite web|title=Orodromeus|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=64338|work=Paleobiology Database|accessdate=20 June 2013}}</ref> ''[[Oryctodromeus]]'' fossils were found in the [[Lima Peaks]] section of the [[Blackleaf Formation]], also from [[Montana]].<ref name = PaleobiologyDatabase /><ref name=Oryctodromeus>{{cite web|title=Oryctodromeus|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=103709|work=Palaobiology Database|accessdate=20 June 2013}}</ref> ''[[Zephyrosaurus]]'', the most widespread genus, lived in southern [[Montana]] and northern [[Wyoming]].<ref name = PaleobiologyDatabase /><ref name=Zephyrosaurus>{{cite web|title=Zephyrosaurus|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=38743|work=Paleobiology Database|accessdate=20 June 2013}}</ref> Its [[holotype]]s locality is the [[Wolf Creek Canyon]], which is a sandstone in the [[Cloverly Formation]].<ref name=ZephyrosaurusSpecies>{{cite web|title=Zephyrosaurus species|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=52889|work=Paleobiology Database|accessdate=20 June 2013}}</ref>

==Age==
Orodromines are widespread throughout time starting in the [[Aptian]] and ending in the [[Campanian]]. The earliest fossils are of ''Zephyrosaurus'' and are from the Aptian (113 Ma).<ref name = Zephyrosaurus/><ref name = ZephyrosaurusSpecies/> After a 13 million year gap in the fossil record, fossils of a less common ''Oryctodromeus'' date to about 95 Ma in the [[Cenomanian]].<ref name = Oryctodromeus/> The next chronological fossils are from 76.5 Ma and belong to ''Albertadromeus''.<ref name=citedoi13/><ref name = Albertadromeus/> The latest fossils in the fossil record belonging to '''Orodromines''' are from the type genus, ''Orodromeus'' and date to 75 Ma.<ref name = Orodromeus/>

==Paleoecology==
[[File:Oryctodromeus.jpg|thumb|right|An illustration of ''[[Oryctodromeus]]'' burrowing]]
All Orodromines lived the lifestyle of a ground dwelling herbivore. ''[[Oryctodromeus]]'' burrows have been discovered. ''[[Orodromeus]]'' and ''[[Zephyrosaurus]]'' also probably lived in burrows.<ref name = PaleobiologyDatabase/>

==Classification==
Orodrominae is sister taxa to [[Thescelosaurinae]]. Its parent taxon is [[Thescelosauridae]] (Brown ''et al'', 2013).<ref name = PaleobiologyDatabase/>

===Phylogeny===
Previously, all genera in Orodrominae (except ''Albertadromeus'', which was named along with Orodrominae) were classified in the now unnatural group [[Hypsilophodontidae]]. They are all now simply considered to be basal members of [[Euornithopoda]].<ref name=Horner1988>Horner, J. and Weishampel, D. (1988), "Acomparative embryological study of two ornithischian dinosaurs". ''Nature'' (London), '''332'''(No. 6161): 256-257 (1988)</ref> The cladogram below is based on a phylogenetic analysis by Brown ''et al.'', 2013.<ref name=citedoi13/>
{{clade| style=font-size:100%; line-height:100%
|label1='''[[Thescelosauridae]]'''
|1={{clade
|label1='''Orodrominae'''
|1={{clade
|1={{clade
|1='''TMP 2008.045.0002'''
|2=''[[Oryctodromeus]]'' }}
|2={{clade
|1=''[[Albertadromeus]]''
|2=''[[Orodromeus]]''
|3=''[[Zephyrosaurus]]'' }} }}
|label2='''[[Thescelosaurinae]]'''
|2={{clade
|1=''[[Parksosaurus]]''
|2={{clade
|1=''[[Changchunsaurus]]''
|2=''[[Jeholosaurus]]''
|3=''[[Haya (dinosaur)|Haya]]''
|4=''[[Thescelosaurus]]'' }} }} }} }}

==References==
{{reflist}}

[[Category:Cretaceous dinosaurs]]

Orodrominae

2017-09-30T12:50:07Z

Wikkler: New phylogeny based on the Boyd analysis of 2015 recovers these two as orodromines.

{{automatic taxobox
| name = Orodromines
| fossil_range = [[Cretaceous]], {{fossil range|113|75|latest=70}}
| image = Orodromeus.jpg
| image_width = 250px
| image_caption = Illustration of the type species, ''[[Orodromeus|Orodromeus makelai]]''
| authority = [[C. M. Brown|Brown]] ''et al'', 2013
| type_species = ''[[Orodromeus|Orodromeus makelai]]'' Horner & Weishampel, [[1988 in paleontology|1988]]
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
* ''[[Albertadromeus]]''
* ''[[Orodromeus]]'' ([[Type species|type]])
* ''[[Koreanosaurus]]''
* ''[[Oryctodromeus]]''
* ''[[Zephyrosaurus]]''
* ''[[Changchunsaurus]]''<ref name=citedoi17>{{Cite journal | last1 = Madzia | first1 = Daniel | last2 = Boyd | first2 = Clint A. | last3 = Mazuch | first3 = Martin | title = A basal ornithopod dinosaur from the Cenomanian of the Czech Republic | doi = 10.1080/14772019.2017.1371258 | journal = Journal of Systematic Paleontology | year = 2017 | pmid = | pmc = }}</ref>
* ''[[Haya_(dinosaur)|Haya]]''<ref name = citedoi17/>
}}

'''Orodrominae''' is a [[subfamily]] of [[Thescelosauridae|Thescelosaurid]] [[dinosaurs]] from the [[Cretaceous]] of [[North America]].<ref name=PaleobiologyDatabase>{{cite web|title=Orodrominae|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=267487|work=Paleobiology Datagase|accessdate=20 June 2013}}</ref>

==Distribution==
Orodromines were a [[North America]]n based group with [[fossil]]s from Canada and United States only.<ref name = PaleobiologyDatabase/> ''[[Albertadromeus]]'', as its name suggests, is only from the upper (later) part of the [[Oldman Formation]] in the [[Belly River Group]] of [[Alberta]], Canada.<ref name=citedoi13>{{Cite journal | last1 = Brown | first1 = C. M. | last2 = Evans | first2 = D. C. | last3 = Ryan | first3 = M. J. | last4 = Russell | first4 = A. P. | title = New data on the diversity and abundance of small-bodied ornithopods (Dinosauria, Ornithischia) from the Belly River Group (Campanian) of Alberta | doi = 10.1080/02724634.2013.746229 | journal = Journal of Vertebrate Paleontology | volume = 33 | issue = 3 | pages = 495 | year = 2013 | pmid = | pmc = }}</ref><ref name=Albertadromeus>{{cite web|title=Albertadromeus|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=267486|work=Paleobiology Database|accessdate=20 June 2013}}</ref> ''[[Orodromeus]]'', the type [[genus]], was widespread through [[Montana]].<ref name = PaleobiologyDatabase /> Its [[holotype]] was found at the [[Egg Mountain]] in the [[Two Medicine Formation]].<ref name=Orodromeus>{{cite web|title=Orodromeus|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=64338|work=Paleobiology Database|accessdate=20 June 2013}}</ref> ''[[Oryctodromeus]]'' fossils were found in the [[Lima Peaks]] section of the [[Blackleaf Formation]], also from [[Montana]].<ref name = PaleobiologyDatabase /><ref name=Oryctodromeus>{{cite web|title=Oryctodromeus|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=103709|work=Palaobiology Database|accessdate=20 June 2013}}</ref> ''[[Zephyrosaurus]]'', the most widespread genus, lived in southern [[Montana]] and northern [[Wyoming]].<ref name = PaleobiologyDatabase /><ref name=Zephyrosaurus>{{cite web|title=Zephyrosaurus|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=38743|work=Paleobiology Database|accessdate=20 June 2013}}</ref> Its [[holotype]]s locality is the [[Wolf Creek Canyon]], which is a sandstone in the [[Cloverly Formation]].<ref name=ZephyrosaurusSpecies>{{cite web|title=Zephyrosaurus species|url=http://paleodb.org/?a=basicTaxonInfo&taxon_no=52889|work=Paleobiology Database|accessdate=20 June 2013}}</ref>

==Age==
Orodromines are widespread throughout time starting in the [[Aptian]] and ending in the [[Campanian]]. The earliest fossils are of ''Zephyrosaurus'' and are from the Aptian (113 Ma).<ref name = Zephyrosaurus/><ref name = ZephyrosaurusSpecies/> After a 13 million year gap in the fossil record, fossils of a less common ''Oryctodromeus'' date to about 95 Ma in the [[Cenomanian]].<ref name = Oryctodromeus/> The next chronological fossils are from 76.5 Ma and belong to ''Albertadromeus''.<ref name=citedoi13/><ref name = Albertadromeus/> The latest fossils in the fossil record belonging to '''Orodromines''' are from the type genus, ''Orodromeus'' and date to 75 Ma.<ref name = Orodromeus/>

==Paleoecology==
[[File:Oryctodromeus.jpg|thumb|right|An illustration of ''[[Oryctodromeus]]'' burrowing]]
All Orodromines lived the lifestyle of a ground dwelling herbivore. ''[[Oryctodromeus]]'' burrows have been discovered. ''[[Orodromeus]]'' and ''[[Zephyrosaurus]]'' also probably lived in burrows.<ref name = PaleobiologyDatabase/>

==Classification==
Orodrominae is sister taxa to [[Thescelosaurinae]]. Its parent taxon is [[Thescelosauridae]] (Brown ''et al'', 2013).<ref name = PaleobiologyDatabase/>

===Phylogeny===
Previously, all genera in Orodrominae (except ''Albertadromeus'', which was named along with Orodrominae) were classified in the now unnatural group [[Hypsilophodontidae]]. They are all now simply considered to be basal members of [[Euornithopoda]].<ref name=Horner1988>Horner, J. and Weishampel, D. (1988), "Acomparative embryological study of two ornithischian dinosaurs". ''Nature'' (London), '''332'''(No. 6161): 256-257 (1988)</ref> The cladogram below is based on a phylogenetic analysis by Brown ''et al.'', 2013.<ref name=citedoi13/>
{{clade| style=font-size:100%; line-height:100%
|label1='''[[Thescelosauridae]]'''
|1={{clade
|label1='''Orodrominae'''
|1={{clade
|1={{clade
|1='''TMP 2008.045.0002'''
|2=''[[Oryctodromeus]]'' }}
|2={{clade
|1=''[[Albertadromeus]]''
|2=''[[Orodromeus]]''
|3=''[[Zephyrosaurus]]'' }} }}
|label2='''[[Thescelosaurinae]]'''
|2={{clade
|1=''[[Parksosaurus]]''
|2={{clade
|1=''[[Changchunsaurus]]''
|2=''[[Jeholosaurus]]''
|3=''[[Haya (dinosaur)|Haya]]''
|4=''[[Thescelosaurus]]'' }} }} }} }}

==References==
{{reflist}}

[[Category:Cretaceous dinosaurs]]

Spinosaurus

2017-09-27T14:43:42Z

Wikkler: /* Skull */ Adding italics to scientific name.

{{pp-semi|small=yes}}
{{good article}}
{{italic title}}{{Automatic taxobox
| name = ''Spinosaurus''
| fossil_range = [[Early Cretaceous|Early]]–[[Late Cretaceous]], {{Fossilrange|112|93.5|earliest=130}}
| image = Spinosaurus in Japan Expo.jpg
| image_width = 250px
| image_caption = Reconstructed ''Spinosaurus'' skeleton in swimming posture
| authority = [[Ernst Stromer|Stromer]], 1915
| type_species = '''''Spinosaurus aegyptiacus'''''
| type_species_authority = Stromer, 1915
| synonyms =
*?''Spinosaurus maroccanus'' [[Dale Russell|Russell]], 1996
*?''Sigilmassasaurus brevicollis'' Russell, 1996
}}
'''''Spinosaurus''''' (meaning "spine lizard") is a [[genus]] of [[Theropoda|theropod]] [[dinosaur]] that lived in what now is [[North Africa]], during the upper [[Albian]] to upper [[Turonian]] [[faunal stage|stages]] of the [[Cretaceous]] [[Period (geology)|period]], about 112 to 93.5 [[annum|million years ago]]. This genus was known first from [[Egypt]]ian remains discovered in 1912 and described by [[Germans|German]] [[paleontology|paleontologist]] [[Ernst Stromer]] in 1915. The original remains were destroyed in [[World War II]], but additional material has come to light in recent years. It is unclear whether one or two species are represented in the fossils reported in the scientific literature. The best known species is ''S. aegyptiacus'' from Egypt, although a potential second species, ''S. maroccanus'', has been recovered from [[Morocco]].

''Spinosaurus'' was among the largest of all known [[carnivore|carnivorous]] dinosaurs, nearly as large or even larger than ''[[Tyrannosaurus]]'', ''[[Giganotosaurus]]'' and ''[[Carcharodontosaurus]]''. Estimates published in 2005, 2007, and 2008 suggested that it was between {{convert|12.6|-|18|m|ft|0}} in length and {{convert|7|to|20.9|t|short ton}} in weight.<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><ref name=TH07>{{cite journal |last=Therrien |first=F. |author2=Henderson, D.M. |year=2007 |title=My theropod is bigger than yours...or not: estimating body size from skull length in theropods |journal=Journal of Vertebrate Paleontology |volume=27 |issue=1 |pages=108–115 |doi=10.1671/0272-4634(2007)27[108:MTIBTY]2.0.CO;2 |issn=0272-4634}}</ref><ref name="dalsassoetal05">{{cite journal |last=dal Sasso |first=C. |author2=Maganuco, S. |author3=Buffetaut, E. |author4= Mendez, M.A. |year=2005 |title=New information on the skull of the enigmatic theropod ''Spinosaurus'', with remarks on its sizes and affinities |journal=Journal of Vertebrate Paleontology |volume=25 |issue=4 |pages=888–896|url=http://www.bioone.org/perlserv/?request=get-abstract&doi=10.1671%2F0272-4634%282005%29025%5B0888%3ANIOTSO%5D2.0.CO%3B2|doi=10.1671/0272-4634(2005)025[0888:NIOTSO]2.0.CO;2 |issn=0272-4634}}</ref> A new estimate published in 2014 and based on a more complete specimen, supported the earlier research, finding that ''Spinosaurus'' could reach lengths of over {{convert|15|m|ft|abbr=on}}.<ref name=Ibrahim_et_al_2014>{{cite journal|last1=Ibrahim|first1=Nizar|last2=Sereno|first2=Paul C.|last3=Dal Sasso|first3=Cristiano|last4=Maganuco|first4=Simone|last5=Fabri|first5=Matteo|last6=Martill|first6=David M.|last7=Zouhri|first7=Samir|last8=Myhrvold|first8=Nathan|last9=Lurino|first9=Dawid A.|title=Semiaquatic adaptations in a giant predatory dinosaur|journal=Science|date=2014|volume=345|issue=6204|doi=10.1126/science.1258750|url=http://www.sciencemag.org/content/345/6204/1613.abstract|pmid=25213375|pages=1613–6}} [http://www.sciencemag.org/content/suppl/2014/09/10/science.1258750.DC1/Ibrahim.SM.pdf Supplementary Information]</ref> The [[skull]] of ''Spinosaurus'' was long and narrow, similar to that of a modern [[crocodilia]]n. ''Spinosaurus'' is known to have eaten fish, and most scientists believe that it hunted both terrestrial and aquatic prey; evidence suggests that it lived both on land and in water as a modern crocodilian does. The distinctive [[spinous process|spines]] of ''Spinosaurus'', which were long extensions of the [[vertebra]]e, grew to at least {{convert|1.65|m|ft|sp=us}} long and were likely to have had skin connecting them, forming a [[sail (anatomy)|sail-like structure]], although some authors{{By whom|date=August 2017}} have suggested that the spines were covered in fat and formed a hump. Multiple functions have been put forward for this structure, including [[thermoregulation]] and display.

==Description==
[[File:Largesttheropods.png|thumb|left|Size comparison of selected giant theropod dinosaurs, ''S. aegyptiacus'' in red]]
Since its discovery, ''Spinosaurus'' has been a contender for the longest and largest theropod dinosaur.<ref>[http://video.nationalgeographic.com/video/magazine/140911-ngm-superjaws nationalgeographic.com 'River Monster': 50-Foot Spinosaurus]</ref> Both [[Friedrich von Huene]] in 1926<ref name = vH26>{{cite journal |last=von Huene |first=F.R. |year=1926 |title=The carnivorous saurischia in the Jura and Cretaceous formations principally in Europe |journal=Rev. Mus. La Plata |volume=29 |pages=35–167}}</ref> and [[Donald F. Glut]] in 1982 listed it as among the most massive theropods in their surveys, at {{convert|15|m|ft|sp=us}} in length and upwards of {{convert|6|t|LT ST|abbr=on}} in weight.<ref name=DFG82>{{cite book |last=Glut |first=D.F. |year=1982 |title=The New Dinosaur Dictionary |publisher=Citadel Press |location=Secaucus, NJ |pages=226–228 |isbn=0-8065-0782-9}}</ref> In 1988, [[Gregory Paul]] also listed it as the longest theropod at {{convert|15|m|ft|sp=us}}, but gave a lower mass estimate of {{convert|4|t|LT ST}}.<ref name=GSP88/>

Dal Sasso ''et al''. (2005) assumed that ''Spinosaurus'' and ''[[Suchomimus]]'' had the same body proportions in relation to their skull lengths, and thereby calculated that ''Spinosaurus'' was {{convert|16|to|18|m|ft|sp=us}} in length and {{convert|7|to|9|t|LT ST}} in weight.<ref name=dalsassoetal05/> The Dal Sasso ''et al''. estimates were criticized because the skull length estimate was uncertain, and (assuming that body mass increases as the cube of body length) scaling ''Suchomimus'' which was {{convert|11|m|ft|sp=us}} long and {{convert|3.8|t|ST}} in mass to the range of estimated lengths of ''Spinosaurus'' would produce an estimated body mass of {{convert|11.7|to|16.7|t|short ton}}.<ref name=TH07/>
[[File:Spinosaurus skull steveoc.jpg|thumb|''S. aegyptiacus'' head based on the 2005 Dal Sasso reconstruction]]
François Therrien and Donald Henderson, in a 2007 paper using scaling based on skull length, challenged previous estimates of the size of ''Spinosaurus'', finding the length too great and the weight too small.<ref name=TH07/> Based on estimated skull lengths of {{convert|1.5|to|1.75|m|ft|sp=us}}, their estimates include a body length of {{convert|12.6|to|14.3|m|ft|sp=us}} and a body mass of {{convert|12|to|20.9|t|LT ST}}.<ref name=TH07/> The lower estimates for ''Spinosaurus'' would imply that the animal was shorter and lighter than ''Carcharodontosaurus'' and ''Giganotosaurus''.<ref name=TH07/> The Therrien and Henderson study has been criticized for the choice of theropods used for comparison (e.g., most of the theropods used to set the initial equations were [[tyrannosaurid]]s and [[Carnosauria|carnosaurs]], which have a different build than spinosaurids), and for the assumption that the ''Spinosaurus'' skull could be as little as {{convert|1.5|m|ft|sp=us}} in length.<ref>{{cite web |url=http://dml.cmnh.org/2007Mar/msg00292.html |title=Comments on Therrien and Henderson's new paper |accessdate=22 September 2010 |last=Mortimer |first=M. |date=2007-03-25 |publisher=Dinosaur Mailing List}}</ref><ref>{{cite web |url=http://dml.cmnh.org/2007Mar/msg00294.html|title=Re: Comments on Therrien and Henderson's new paper|accessdate=22 September 2010|last=Harris|first=J.D. |date=2007-03-26 |publisher=Dinosaur Mailing List}}</ref> Improvement of the precision of size estimates for ''Spinosaurus'' requires the discovery of more complete remains as available for some other dinosaurs,<ref>{{Cite journal | last1 = Bates | first1 = K.T. | last2 = Manning | first2 = P.L. | last3 = Hodgetts | first3 = D. | last4 = and Sellers | first4 = W.I. | last5 = Sellers | first5 = William I. | title = Estimating Mass Properties of Dinosaurs Using Laser Imaging and 3D Computer Modelling | journal = PLoS ONE | volume = 4 | issue = 2 | pages = e4532 | year = 2009 | pmid = 19225569 | pmc = 2639725 | doi = 10.1371/journal.pone.0004532 | bibcode=2009PLoSO...4.4532B | editor1-last = Beckett | editor1-first = Ronald}}</ref> especially the limb bones of ''Spinosaurus'' which are "hitherto unknown".<ref name=dalsassoetal05/>

===Neural spines===
[[File:Reproduction of Stromer's Spinosaurus Display.jpg|thumb|left|Reconstruction of the holotype fossils]]
Very tall [[spinous process|neural spines]] growing on the back [[vertebra]]e of ''Spinosaurus'' formed the basis of what is usually called the animal's "sail". The lengths of the neural spines reached over 10 times the diameters of the vertebral bodies from which they extended.<ref name=JBB97>{{cite journal |last=Bailey |first=J.B.|year=1997 |title=Neural spine elongation in dinosaurs: sailbacks or buffalo-backs? |journal=Journal of Paleontology |volume=71 |issue=6 |pages=1124–1146|jstor= 1306608 }}</ref> The neural spines were slightly longer front to back at the base than higher up, and were unlike the thin rods seen in the [[pelycosaur]] finbacks ''[[Edaphosaurus]]'' and ''[[Dimetrodon]]'', contrasting also with the thicker spines in the iguanodontian ''[[Ouranosaurus]]''.<ref name=JBB97/>

''Spinosaurus'' sails were unusual, although other dinosaurs, namely the [[ornithopod]] ''[[Ouranosaurus]]'', which lived a few million years earlier in the same general region as ''Spinosaurus'', and the South American sauropod ''[[Amargasaurus]],'' might have developed similar structural adaptations of their vertebrae. The sail may be an [[Analogy (biology)|analog]] of the sail of the [[Permian]] [[synapsid]] ''[[Dimetrodon]]'', which lived before the dinosaurs even appeared, produced by [[convergent evolution]].<ref name=JBB97/>

The structure may also have been more hump-like than sail-like, as noted by Stromer in 1915 ("one might rather think of the existence of a large hump of fat <nowiki>[</nowiki>[[German Language|German]]: ''Fettbuckel''], to which the [neural spines] gave internal support")<ref name=Stromer15/> and by Jack Bowman Bailey in 1997.<ref name=JBB97/> In support of his "buffalo-back" hypothesis, Bailey argued that in ''Spinosaurus'', ''Ouranosaurus'', and other dinosaurs with long neural spines, the spines were relatively shorter and thicker than the spines of [[pelycosaur]]s (which were known to have sails); instead, the dinosaurs' neural spines were similar to the neural spines of extinct hump-backed mammals such as ''[[Megacerops]]'' and ''[[Bison latifrons]]''.<ref name=JBB97/>

===Skull===
[[File:Spinosaurus skull en.svg|thumb|Annotated skull diagram]]
The skull had a narrow snout filled with straight conical teeth that lacked serrations. There were six or seven teeth on each side of the very front of the upper jaw, in the [[premaxilla]]e, and another twelve in both [[maxilla]]e behind them. The second and third teeth on each side were noticeably larger than the rest of the teeth in the premaxilla, creating a space between them and the large teeth in the [[anatomical terms of location|anterior]] maxilla; large teeth in the lower jaw faced this space. The very tip of the snout holding those few large anterior teeth was expanded, and a small crest was present in front of the eyes.<ref name=dalsassoetal05/> Using the dimensions of three specimens known as MSNM V4047, UCPC-2, and BSP 1912 VIII 19, and assuming that the postorbital part of the skull of MSNM V4047 had a shape similar to the postorbital part of the skull of ''[[Irritator]]'', Dal Sasso ''et al''. (2005) estimated that the skull of ''Spinosaurus'' was {{convert|1.75|m|ft|sp=us}} long.<ref name=dalsassoetal05/> The Dal Sasso ''et al''. skull length estimate was questioned because skull shapes can vary across spinosaurid species.<ref name=TH07/>

A 2013 made study performed by scientists Andrew R. Cuff and Emily Rayfield showed that Spinosaurids like '' Spinosaurus'' had relatively poor resistance in their skulls for torsion compared to other members of this group (''Baryonyx'') and modern alligators, thus showing ''Spinosaurus'' preyed more regularly on fish than it did on land animals, although considered predators of the former too.<ref name=cuff13/>

==Discovery and naming==

===Naming of species===
[[File:Spinosaurus aegyptiacus holotype skeletal.jpg|thumb|Skeletal restoration showing the holotype elements]]
Two species of ''Spinosaurus'' have been named: ''Spinosaurus aegyptiacus'' (meaning "Egyptian spine lizard") and ''Spinosaurus maroccanus'' (meaning "Moroccan spine lizard").<ref name=Smith06/><ref name=DAR96/><ref>[http://www.ted.com/talks/nizar_ibrahim_how_we_unearthed_the_spinosaurus www.ted.com November 2014 Nizar Ibrahim How We Unearthed the Spinosaurus]</ref>

The first described remains of ''Spinosaurus'' were found and described in the early 20th century. In 1912, Richard Markgraf discovered a partial skeleton of a dinosaur in the [[Bahariya Formation]] of western Egypt. In 1915, German paleontologist [[Ernst Stromer]] published an article assigning the specimen to a new genus and species ''Spinosaurus aegyptiacus''.<ref name=Stromer15>{{cite journal |last=Stromer |first=E. |authorlink=Ernst Stromer |year=1915 |title= Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstes Cenoman). 3. Das Original des Theropoden ''Spinosaurus aegyptiacus'' nov. gen., nov. spec |journal= Abhandlungen der Königlich Bayerischen Akademie der Wissenschaften, Mathematisch-physikalische Klasse |volume=28|issue=3 |pages=1–32 |language=German|url=http://www.megaupload.com/?d=3KCCC7LS}}</ref><ref name=Smith06/>

Fragmentary additional remains from Bahariya, including vertebrae and hindlimb bones, were designated by Stromer as "''Spinosaurus B''" in 1934.<ref name=stromer34>{{cite journal |last=Stromer |first=E. |authorlink=Ernst Stromer |year=1934 |title=Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstes Cenoman). 13. Dinosauria|journal=Abhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch-naturwissenschaftliche Abteilung, Neue Folge |volume=22 |pages=1–79 |language=German }}</ref> Stromer considered them different enough to belong to another species, and this has been borne out. With the advantage of more expeditions and material, it appears that they pertain either to ''[[Carcharodontosaurus]]''<ref name="serenoetal98">{{cite journal | last1 = Sereno | first1 = P.C. | last2 = Beck | first2 = A.L. | last3 = Dutheuil | first3 = D.B. | last4 = Gado | first4 = B. | last5 = Larsson | first5 = H.C. | last6 = Lyon | first6 = G.H. | last7 = Marcot | first7 = J.D. | last8 = Rauhut | first8 = O.W.M. | last9 = Sadleir | first9 = R.W. | last10 = Sidor | first10 = C.A. | last11 = Varricchio | first11 = D.J. | last12 = Wilson | first12 = G.P. | last13 = Wilson | first13 = J.A. | year = 1998 | title = A long-snouted predatory dinosaur from Africa and the evolution of spinosaurids | url = | journal = Science | volume = 282 | issue = | pages = 1298–1302 | doi=10.1126/science.282.5392.1298 | pmid=9812890}}</ref> or to ''[[Sigilmassasaurus]]''.<ref name=DAR96>{{cite journal |last=Russell |first=D.A. |authorlink=Dale Russell |year=1996 |title=Isolated dinosaur bones from the Middle Cretaceous of the Tafilalt, Morocco |journal=Bulletin du Muséum National d'Histoire Naturelle, Paris, 4e série, section C |volume=18 |issue=2–3 |pages=349–402}}</ref>

''S. maroccanus'' was originally described by [[Dale Russell]] in 1996 as a new species based on the length of its neck vertebrae.<ref name=DAR96/> Specifically, Russell claimed that the ratio of the length of the centrum ([[body of vertebra]]) to the height of the posterior articular facet was 1.1 in ''S. aegyptiacus'' and 1.5 in ''S. maroccanus''.<ref name=DAR96/> Later authors have been split on this topic. Some authors note that the length of the vertebrae can vary from individual to individual, that the holotype specimen was destroyed and thus cannot be compared directly with the ''S. maroccanus'' specimen, and that it is unknown which cervical vertebrae the ''S. maroccanus'' specimens represent. Therefore, though some have retained the species as valid without much comment,<ref name=HMC04/><ref>{{Cite journal | last1 = Mahler | first1 = L. | title = Record of Abelisauridae (Dinosauria: Theropoda) from the Cenomanian of Morocco | journal = Journal of Vertebrate Paleontology | volume = 25 | pages = 236–239| year = 2005 | issn = 0272-4634 | doi = 10.1671/0272-4634(2005)025[0236:ROADTF]2.0.CO;2 | url= http://www.oeb.harvard.edu/faculty/losos/mahler/publications/Mahler_2005.pdf }}</ref><ref name=Hasegawa2010>{{cite journal |last1= Hasegawa |first1= Y. |last2= Tanaka |first2= G. |last3= Takakuwa |first3= Y. |last4= and Koike |first4= S. |year=2010 |title= Fine sculptures on a tooth of ''Spinosaurus'' (Dinosauria, Theropoda) from Morocco |journal= Bulletin of Gunma Museum of Natural History |volume=14 |pages=11–20 |url= http://www.gmnh.pref.gunma.jp/research/no_14/bulletin14_2.pdf }}</ref> most researchers regard ''S. maroccanus'' as a ''[[nomen dubium]]''<ref name=dalsassoetal05/><ref name = BufOua02>{{cite journal |author1=Buffetaut, E. |author2=Ouaja, M. |last-author-amp=yes |year=2002 |title= A new specimen of ''Spinosaurus'' (Dinosauria, Theropoda) from the Lower Cretaceous of Tunisia, with remarks on the evolutionary history of the Spinosauridae |journal=Bulletin de la Société Géologique de France |volume=173 |issue= 5|pages= 415–421|doi=10.2113/173.5.415|url=http://documents.irevues.inist.fr/bitstream/handle/2042/216/04.pdf }}</ref><ref name=Rauhut03>{{cite journal | author = Rauhut, O.W.M. | year = 2003 | title = The interrelationships and evolution of basal theropod dinosaurs | journal = Special Papers in Palaeontology | volume = 69 | pages = 1–213|isbn=0-901702-79-X}}</ref> or as a junior synonym of ''S. aegyptiacus''.<ref name=serenoetal98 />

===Specimens===
Six main partial specimens of ''Spinosaurus'' have been described.
[[File:Spinosaurus holotype.jpg|thumb|Plate I in Stromer (1915) showing ''S. aegyptiacus'' holotype elements]]
'''BSP 1912 VIII 19''', described by Stromer in 1915 from the [[Bahariya Formation]], was the [[holotype]].<ref name = Stromer15/><ref name=Smith06/> The material consisted of the following items, most of which were incomplete: right and left [[dentary|dentaries]] and [[splenial]]s from the lower jaw measuring {{convert|75|cm|in|sp=us}} long; a straight piece of the left [[maxilla]] that was described but not drawn; 20 teeth; 2 [[neck|cervical]] vertebrae; 7 dorsal (trunk) vertebrae; 3 sacral vertebrae; 1 [[tail|caudal]] vertebra; 4 thoracic ribs; and [[gastralium|gastralia]].<ref name = Stromer15/> Of the nine neural spines whose heights are given, the longest ("i," associated with a dorsal vertebra) was {{convert|1.65|m|ft|sp=us}} in length.<ref name = Stromer15/> Stromer claimed that the specimen was from the early Cenomanian, about 97 million years ago.<ref name = Stromer15/><ref name=Smith06/>

This specimen was destroyed in [[World War II]], specifically "during the night of 24/25 April 1944 in a British bombing raid of Munich" that severely damaged the building housing the [[Paläontologisches Museum München]] (Bavarian State Collection of Paleontology).<ref name=Smith06/> However, detailed drawings and descriptions of the specimen remain. Stromer's son donated Stromer's archives to the Paläontologische Staatssammlung München in 1995, and Smith ''et al.'' analyzed two photographs of the ''Spinosaurus'' holotype specimen BSP 1912 VIII 19 discovered in the archives in 2000.<ref name=Smith06/> On the basis of a photograph of the lower jaw and a photograph of the entire specimen as mounted, Smith concluded that Stromer's original 1915 drawings were slightly inaccurate.<ref name=Smith06/> In 2003, Oliver Rauhut suggested that Stromer's ''Spinosaurus'' [[holotype]] was a [[chimera (paleontology)|chimera]], composed of vertebrae and neural spines from a [[Carcharodontosauridae|carcharodontosaurid]] similar to ''[[Acrocanthosaurus]]'' and a [[dentary]] from ''[[Baryonyx]]'' or ''[[Suchomimus]]''.<ref name=Rauhut03/> This analysis was rejected in at least one subsequent paper.<ref name=dalsassoetal05/>

'''NMC 50791''', held by the [[Canadian Museum of Nature]], is a mid-cervical vertebra which is {{convert|19.5|cm|in|sp=us}} long from the [[Kem Kem Beds]] of [[Morocco]].<ref name=DAR96/> It is the holotype of ''Spinosaurus maroccanus'' as described by Russell in 1996.<ref name=DAR96/> Other specimens referred to ''S. maroccanus'' in the same paper were two other mid-cervical vertebrae (NMC 41768 and NMC 50790), an anterior dentary fragment (NMC 50832), a mid-dentary fragment (NMC 50833), and an anterior dorsal [[vertebral arch|neural arch]] (NMC 50813).<ref name=DAR96/> Russell stated that "only general locality information could be provided" for the specimen, and therefore it could be dated only "possibly" to the Albian.<ref name=DAR96/>
[[File:Spinosaurus marocannus.jpg|thumb|Specimen MNHN SAM 124 of ''S. maroccanus'', [[Muséum national d'Histoire naturelle|Muséum National d'Histoire Naturelle]], Paris]]
'''MNHN SAM 124''', housed at the [[Muséum National d'Histoire Naturelle]], is a snout (consisting of partial premaxillae, partial maxillae, [[vomer]]s, and a dentary fragment).<ref name=Taquet1998>{{Cite journal | last1 = Taquet | first1 = P. | last2 = and Russell | first2 = D.A. | title = New data on spinosaurid dinosaurs from the Early Cretaceous of the Sahara | journal = Comptes Rendus de l'Académie des Sciences - Series IIA - Earth & Planetary Sciences | volume = 327 | pages = 347–353 | year = 1998 | doi = 10.1016/S1251-8050(98)80054-2 | url=http://lesdinos.free.fr/spi329.pdf |accessdate=22 September 2010 | bibcode=1998CRASE.327..347T | issue = 5}}</ref> Described by Taquet and Russell in 1998, the specimen is {{convert|13.4|to(-)|13.6|cm|in|sp=us}} in width; no length was stated.<ref name=Taquet1998/> The specimen was located in Algeria, and "is of Albian age."<ref name=Taquet1998/> Taquet and Russell believed that this specimen along with a premaxilla fragment (SAM 125), two cervical vertebrae (SAM 126-127), and a dorsal neural arch (SAM 128), belonged to ''S. maroccanus''.<ref name=Taquet1998/>

'''BM231''' (in the collection of the Office National des Mines, Tunis) was described by Buffetaut and Ouaja in 2002.<ref name = BufOua02/> It consists of a partial anterior dentary {{convert|11.5|cm|in|sigfig=3}} in length from an early Albian [[stratum]] of the [[Chenini Formation]] of [[Tunisia]].<ref name = BufOua02/> The dentary fragment, which included four [[Dental alveolus|alveoli]] and two partial teeth, was "extremely similar" to existing material of ''S. aegyptiacus''.<ref name = BufOua02/>

'''UCPC-2''' in the [[University of Chicago]] Paleontological Collection consists mainly of two narrow connected [[nasal bone|nasal]]s with a "fluted crest" from the region between the eyes.<ref name=dalsassoetal05/> The specimen, which is {{convert|18.0|cm|in|sigfig=3}} long, was located in an early Cenomanian part of the Moroccan Kem Kem Beds in 1996 and described in the scientific literature in 2005 by [[Cristiano Dal Sasso]] of the [[Museo Civico di Storia Naturale di Milan|Civic Natural History Museum]] in [[Milan]] and colleagues.<ref name=dalsassoetal05/>
[[File:Spinocombo.jpg|thumb|Specimen MSNM V4047 of ''S. aegyptiacus'' in the [[Museo Civico di Storia Naturale di Milano|Civic Natural History Museum]] in [[Milan]]]]
'''MSNM V4047''' (in the [[Museo Civico di Storia Naturale di Milan|Museo di Storia Naturale di Milano]]), described by Dal Sasso ''et al.'' in 2005, consists of a snout (premaxillae, partial maxillae, and partial nasals) {{convert|98.8|cm|in|sigfig=3}} long from the Kem Kem Beds.<ref name=dalsassoetal05/> Like UCPC-2, it is thought to have come from the early Cenomanian.

'''FSAC-KK 11888''' is a partial subadult skeleton recovered from the Kem Kem beds of North Africa. Described by Ibrahim ''et al.'' (2014) and designated as the neotype specimen (although Evers et al. 2015 reject the neotype designation for FSAC-KK-11888).<ref name="Evers 2015"/> It includes cervical vertebrae, dorsal vertebrae, neural spines, a complete sacrum, femora, tibiae, pedal phalanges, caudal vertebra, several dorsal ribs, and fragments of the skull.<ref>{{cite journal|title=Ibrahim, N.; Sereno, P. C.; Dal Sasso, C.; Maganuco, S.; Fabbri, M.; Martill, D. M.; Zouhri, S.; Myhrvold, N.; Iurino, D. A. (2014). "Supplementary Materials for Semiaquatic Adaptations in a Giant Predatory Dinosaur". DOI: 10.1126/science.1258750}}</ref> The body proportions of this specimen have been debated as the hind limbs are disproportionately shorter in the specimen than in previous reconstructions. However, it has been demonstrated by multiple paleontologists that the specimen is not a chimaera, and is indeed a specimen of ''Spinosaurus'' that suggests that the animal had much smaller hind limbs than previously thought <ref>{{cite web|title=Cau, Andrea (2014; online) "Spinosaurus Revolution, Episodio IV: Una soluzione a tutti gli enigmi?"http://theropoda.blogspot.com/2014/09/spinosaurus-revolution-episodio-iv-una.html}}</ref><ref>{{cite web|title=Cau, Andrea (2014) "Spinosaurus Revolution, Episodio V: Sigilmassasaurus vs Spinosaurus: una battaglia tafonomica"http://theropoda.blogspot.com/2014/09/spinosaurus-revolution-episodio-v.html}}</ref><ref>{{cite web|title=Mortimer, Michael. (2014) "Spinosaurus surprise"http://theropoddatabase.blogspot.com/2014/09/spinosaurus-surprise.html}}</ref>
Other known specimens consist mainly of very fragmentary remains and scattered teeth. These include:
*A 1986 paper described prismatic structures in [[tooth enamel]] from two ''Spinosaurus'' teeth from Tunisia.<ref>{{Cite journal | last1 = Buffetaut | first1 = E. | last2 = Dauphin | first2 = Y. | last3 = Jaeger | first3 = J.-J. | last4 = Martin | first4 = M. | last5 = Mazin | first5 = J.-M. | last6 = and Tong | first6 = H. | title = Prismatic dental enamel in theropod dinosaurs | journal = Naturwissenschaften | volume = 73 | pages = 326–327 | year = 1986 | doi = 10.1007/BF00451481 | pmid=3748191 | bibcode=1986NW.....73..326B | issue = 6}}</ref>
* Buffetaut (1989, 1992) referred three specimens from the Institut und Museum für Geologie und Paläontologie of the [[University of Göttingen]] in Germany to ''Spinosaurus'': a right maxilla fragment IMGP 969-1, a jaw fragment IMGP 969-2, and a tooth IMGP 969-3.<ref name=Buffetaut89>{{Cite journal | last1 = Buffetaut | first1 = E. | title = New remains of the enigmatic dinosaur ''Spinosaurus'' from the Cretaceous of Morocco and the affinities between ''Spinosaurus'' and ''Baryonyx'' | journal = Neues Jahrbuch für Geologie und Paläontologie, Monatshefte | issue = 2 | pages = 79–87 | year = 1989}}</ref><ref name=Buffetaut92>{{Cite journal | last1 = Buffetaut | first1 = E. | title = Remarks on the Cretaceous theropod dinosaurs ''Spinosaurus'' and ''Baryonyx'' | journal = Neues Jahrbuch für Geologie und Paläontologie, Monatshefte | issue = 2 | pages = 88–96 | year = 1992}}</ref> These had been found in a Lower Cenomanian or Upper Albian deposit in southeastern Morocco in 1971.<ref name=Buffetaut89/>
[[File:Spinoface.jpg|thumb|Reconstructed subadult skeleton from a private collection]]
*Kellner and Mader (1997) described two unserrated spinosaurid teeth from Morocco (LINHM 001 and 002) that were "highly similar" to the teeth of the ''S. aegyptiacus'' holotype.<ref>{{cite journal |last1= Kellner |first1=A.W.A. |last2= and Mader |first2=B.J. |year= 1997 |title= Archosaur teeth from the Cretaceous of Morocco |journal= Journal of Paleontology |volume= 71 |issue= 3 |pages= 525–527 |jstor= 1306632 }}</ref>
*Teeth from the Chenini Formation in Tunisia which are "narrow, somewhat rounded in cross-section, and lack the anterior and posterior serrated edges characteristic of theropods and basal [[archosaur]]s" were assigned to ''Spinosaurus'' in 2000.<ref>{{Cite journal | last1 = Benton | first1 = M.J. | last2 = Bouaziz | first2 = S. | last3 = Buffetaut | first3 = E. | last4 = Martill | first4 = D. | last5 = Ouaja | first5 = M. | last6 = Soussi | first6 = M. | last7 = and Trueman | first7 = C. | title = Dinosaurs and other fossil vertebrates from fluvial deposits in the Lower Cretaceous of southern Tunisia | journal = Palaeogeography, Palaeoclimatology, Palaeoecology | volume = 157 | pages = 227–246 | year = 2000 | doi = 10.1016/S0031-0182(99)00167-4 | issue = 3–4}}</ref>
*Teeth from the [[Echkar Formation]] of [[Niger]] were "tentatively" referred to ''Spinosaurus'' in 2007.<ref name="brusatte&sereno2007">{{Cite journal | last1 = Brusatte | first1 = S. L. | last2 = and Sereno | first2 = P. C. | title = A new species of ''Carcharodontosaurus'' (Dinosauria: Theropoda) from the Cenomanian of Niger and a revision of the genus | journal = Journal of Vertebrate Paleontology | volume = 27 | pages = 902–916 | year = 2007 | doi = 10.1671/0272-4634(2007)27[902:ANSOCD]2.0.CO;2 | issn = 0272-4634 | issue = 4}}</ref>
* A partial tooth 8 cm long purchased at a fossil trade show, reportedly from the Kem Kem Bed of Morocco and attributed to ''Spinosaurus maroccanus'', showed 1–5 mm wide longitudinal striations and micro-structures (irregular ridges) among the striations in a 2010 paper.<ref name=Hasegawa2010/>


'''MHNM.KK374 to.KK378''' are five isolated quadrates (skull bones) of different sizes were collected by locals and acquired commercially in
the Kem Kem region of southeastern Morocco, provided by François Escuillié and are deposited in the collections of the Muséum d’Histoire Naturelle of Marrakech. The quadrates show two different morphologies suggesting the existence of two spinosaurines in Morocco.<ref>{{cite journal | last1 = Hendrickx | first1 = C. | last2 = Mateus | first2 = O. | last3 = Buffetaut | first3 = E. | year = 2016 | title = Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa | url = | journal = PLoS ONE | volume = 11 | issue = | page = e0144695 | doi=10.1371/journal.pone.0144695}}</ref>

====Possible specimens====
Possible material belonging to ''Spinosaurus'' has been reported from the [[Turkana Grits]] of [[Kenya]].<ref name= Weishampel04>{{cite book |author1=Weishampel, D.B. |author2=Barrett, P.M. |author3=Coria, R.A. |author4=Le Loeuff, J. |author5=Xu, X. |author6=Zhao, X. |author7=Sahni, A. |author8=Gomani, E.M.P. |author9=Noto, C.R. |last-author-amp=yes |editor1=Weishampel, D.B. |editor2=Dodson, P. |editor3=Osmólska, H. |title=The Dinosauria |year= 2004|publisher=University of California Press |location=Berkeley |isbn=978-0-520-25408-4 |edition= 2nd|pages=517–606 |chapter=Dinosaur distribution}}</ref>

Some scientists have considered the genus ''[[Sigilmassasaurus]]'' a junior synonym of ''Spinosaurus''. In Ibrahim et al. (2014), the specimens of ''Sigilmassasaurus'' was referred to ''Spinosaurus aegyptiacus'' together with "Spinosaurus B" as the [[neotype]] and ''Spinosaurus maroccanus'' was considered as a ''[[nomen dubium]]'' following the conclusions of the other papers.<ref name="dalsassoetal05"/><ref name=Ibrahim_et_al_2014/><ref name="serenoetal98" /> A 2015 re-description of ''Sigilmassasaurus'' disputed these conclusions, and considered the genus valid.<ref name="Evers 2015">{{Cite journal|doi=10.7717/peerj.1323|title=A reappraisal of the morphology and systematic position of the theropod dinosaur ''Sigilmassasaurus'' from the "middle" Cretaceous of Morocco|journal=PeerJ|volume=3|pages=e1323|year=2015|last1=Evers|first1=S. W.|last2=Rauhut|first2=O. W. M.|last3=Milner|first3=A. C.|last4=McFeeters|first4=B.|last5=Allain|first5=R.|pmid=26500829|pmc=4614847}}</ref>

==Classification==
[[File:Спинозавр - новая реконструкция.jpg|thumb|Restoration based on the 2014 description]]
[[File:Spinosaurus sp Marocco Cenom.JPG|thumb|Tooth from Morocco]]
''Spinosaurus'' gives its name to the [[Spinosauridae]] [[family (biology)|family]] of dinosaurs, which includes two subfamilies: Baryonychinae and Spinosaurinae. The Baryonychinae include ''[[Baryonyx]]'' from southern [[England]] and ''[[Suchomimus]]'' from [[Niger]] in central [[Africa]]. The Spinosaurinae include ''Spinosaurus'', ''[[Irritator]]'' from [[Brazil]], and ''[[Angaturama]]'' (which is probably synonymous with ''Irritator'') from Brazil.<ref name=dalsassoetal05/> The Spinosaurinae share unserrated straight teeth that are widely spaced (e.g., 12 on one side of the maxilla), as opposed to the Baryonychinae which have serrated curved teeth that are numerous (e.g., 30 on one side of the maxilla).<ref name=dalsassoetal05/><ref name=HMC04>{{cite book|title=The Dinosauria|year=2004|chapter=Basal Tetanurae|author1=Holtz, T.R., Jr. |author2=Molnar, R.E. |author3=Currie, P.J. |last-author-amp=yes |editor1=Weishampel, D.B. |editor2=Dodson, P. |editor3=Osmólska, H. |pages=71–110|edition= 2nd|publisher=University of California Press|isbn=978-0-520-25408-4}}</ref>

The following [[cladogram]] shows an analysis of [[Tetanurae]] simplified to show only Spinosauridae from Allain ''et al.'' (2012):<ref name=AXRK12>{{Cite journal | last1 = Allain | first1 = R. | last2 = Xaisanavong | first2 = T. | last3 = Richir | first3 = P. | last4 = Khentavong | first4 = B. | title = The first definitive Asian spinosaurid (Dinosauria: Theropoda) from the early cretaceous of Laos | doi = 10.1007/s00114-012-0911-7 | journal = Naturwissenschaften | year = 2012 | pmid = 22528021| pmc = | volume=99 | issue=5 | pages=369–377|bibcode = 2012NW.....99..369A }}</ref>

{{clade| style=font-size:100%;line-height:85%
|label1=[[Spinosauridae]]
|1={{clade
|label1=[[Spinosaurinae]]
|1={{clade
|1=''[[Irritator]]''[[File:Irritator Life Reconstruction.jpg|80 px]]
|2='''''Spinosaurus'''''[[File:Spinosaurus by Joschua Knüppe.png|80 px]] }}
|label2=[[Baryonychinae]]
|2={{clade
|1=''[[Ichthyovenator]]''
|2={{clade
|1=''[[Suchomimus]]'' [[File:Suchomimus2.jpg|80 px]]
|2=''[[Baryonyx]]''[[File:Baryonyx BW.jpg|80 px]]}} }} }} }}

==Paleobiology==

===Function of neural spines===
[[File:Spinosaurus vertebrae.jpg|thumb|left|upright|1915 illustration of ''S. aegyptiacus'' dorsal vertebrae]]
The function of the dinosaur's sail or hump is uncertain; scientists have proposed several [[hypothesis|hypotheses]] including heat regulation and display. In addition, such a prominent feature on its back could make it appear even larger than it was, intimidating other animals.<ref name=JBB97/>

The structure may have been used for [[thermoregulation]]. If the structure contained abundant blood vessels, the animal could have used the sail's large surface area to absorb heat. This would imply that the animal was only partly warm-blooded at best and lived in climates where nighttime temperatures were cool or low and the sky usually not cloudy. It is also possible that the structure was used to radiate excess heat from the body, rather than to collect it. Large animals, due to the relatively small ratio of surface area of their body compared to the overall volume ([[Haldane's principle]]), face far greater problems of dissipating excess heat at higher temperatures than gaining it at lower. Sails of large dinosaurs added considerably to the skin area of their bodies, with minimum increase of volume. Furthermore, if the sail was turned away from the sun, or positioned at a 90 degree angle towards a cooling wind, the animal would quite effectively cool itself in the warm climate of Cretaceous Africa.<ref name=LBH75>{{cite book |last=Halstead |first=L.B.|authorlink=Beverly Halstead|year=1975 |title=The Evolution and Ecology of the Dinosaurs |publisher=Eurobook Limited |location=London |pages=1–116 |isbn=0-85654-018-8 }}</ref> However, Bailey (1997) was of the opinion that a sail could have absorbed more heat than it radiated.<ref name=JBB97/> Bailey proposed instead that ''Spinosaurus'' and other dinosaurs with long neural spines had fatty humps on their backs for energy storage, insulation, and shielding from heat.<ref name=JBB97/>

Elaborate body structures of many modern-day animals usually serve to attract members of the opposite sex during mating. It is quite possible that the sails or humps of these dinosaurs were used for courtship, in a way similar to a [[peacock]]'s tail. Stromer speculated that males and females may have differed in the size of the neural spine.<ref name=Stromer15/>

Gimsa ''et al''. (2015) suggest that the dorsal sail of ''Spinosaurus'' was homologous to the dorsal fins of [[sailfish]] and served a hydrodynamic purpose.<ref>{{cite journal|title=Gimsa, J., Sleigh, R., Gimsa, U., (2015) : "The riddle of ''Spinosaurus aegyptiacus'' ' dorsal sail". University of Rostock, Chair for Biophysics, Gertrudenstr. 11A, 18057 Rostock, Germany}}</ref> Gimsa and others point out that more basal, long-legged spinosaurids have otherwise round or crescent-shaped dorsal sails, whereas in ''Spinosaurus'', the dorsal neural spines form a shape that is roughly rectangular and similar in shape to the dorsal fins of sailfish. They therefore argue that ''Spinosaurus'' used its dorsal neural sail in the same manner as sailfish, and that it also employed its long narrow tail to stun prey like a modern thresher shark.
Sailfish employ their dorsal fins for herding schools of fish into a "bait hall" where they cooperate to trap the fish into a certain area where the sailfish can snatch the fish with their bills.
The sail could have possibly reduced yaw rotation by counteracting the lateral force in the direction opposite to the slash as suggested by Gimsa ''et al''., (2015).
Gimsa and colleagues specifically wrote :
[[File:Spinosaurus durbed.jpg|thumb|Restoration]]
''Spinosaurus'' anatomy exhibits another feature that may have a modern homology: its long tail resembled that of the thresher shark, employed to slap the water to herd and stun shoals of fish before devouring them (Oliver ''et al''., 2013).
The strategies that sailfish and thresher sharks employ against shoaling fish are more effective when the shoal is first concentrated into a ‘bait ball’ (Helfman, Collette & Facey, 1997; Oliver ''et al''., 2013; Domenici ''et al''., 2014). Since this is difficult for individual predators to achieve, they cooperate in this effort. When herding a shoal of fish or squid, sailfish also raise their sails to make themselves appear larger. When they slash or wipe their bills through shoaling fish by turning their heads, their dorsal sail and fins are outstretched to stabilize their bodies hydrodynamically (Lauder & Drucker, 2004). Domenici et al. (2014) postulate that these fin extensions enhance the accuracy of tapping and slashing. The sail can reduce yaw rotation by counteracting the lateral force in the direction opposite to the slash. This means that prey is less likely to recognize the massive trunk as being part of an approaching predator (Marras ''et al''., 2015; Webb & Weihs 2015). Film footage available online impressively demonstrates the hunting strategies of sailfish and thresher sharks.

Interestingly, ''Spinosaurus'' exhibited the anatomical features required to combine all three hunting strategies: a sail for herding prey more efficiently, as well as flexible tail and neck to slap the water for stunning, injuring or killing prey. The submerged dorsal sail would have provided a strong centreboard-like counterforce for powerful sidewards movements of the strong neck and long tail, as performed by sailfish (Domenici ''et al''., 2014) or thresher sharks (Oliver ''et al''., 2013). While smaller dorsal sails or fins make the dorsal water volume better accessible for slashing, it can be speculated that their smaller stabilization effect makes lateral slashing less efficient (e.g. for thresher sharks). Forming a hydrodynamic fulcrum and hydrodynamically stabilizing the trunk along the dorsoventral axis, ''Spinosaurus''’ sail would also have compensated for the inertia of the lateral neck by tail movements and vice versa not only for predation but also for accelerated swimming. This behaviour might also have been one reason for ''Spinosaurus''’ muscular chest and neck reported by Ibrahim ''et al''. (2014).'' "
Finally, it is quite possible that the sail or hump combined these functions, acting normally as a heat regulator, becoming a courting aid during the mating season, being used to cool itself and, on occasions, turning into an intimidating device when an animal was feeling threatened.<ref name=JBB97/>

===Diet===
[[File:Spinosaurus new skull.jpg|thumb|left|upright|Reconstructed skull and neck]]
It is unclear whether ''Spinosaurus'' was primarily a terrestrial predator or a [[piscivore]], as indicated by its elongated jaws, conical teeth and raised nostrils. The hypothesis of spinosaurs as specialized fish eaters has been suggested before by A. J. Charig and A. C. Milner for ''[[Baryonyx]]''. They base this on the anatomical similarity with crocodilians and the presence of digestive acid-etched fish scales in the rib cage of the [[type specimen]].<ref name=CM97/> Large fish are known from the faunas containing other spinosaurids, including the ''[[Mawsonia (fish)|Mawsonia]]'', in the mid-Cretaceous of northern Africa and Brazil. Direct evidence for spinosaur diet comes from related [[Europe]]an and [[South America]]n taxa. ''Baryonyx'' was found with fish scales and bones from juvenile ''[[Iguanodon]]'' in its stomach, while a tooth embedded in a South American [[pterosaur]] bone suggests that spinosaurs occasionally preyed on pterosaurs,<ref>{{cite journal |last=Buffetaut |first=E. |author2=Martill, D. |author3= Escuillié, F. |year=2004 |title=Pterosaurs as part of a spinosaur diet |journal=Nature |volume=430 |page=33 |doi=10.1038/430033a |pmid=15229562 |issue=6995|bibcode = 2004Natur.429...33B }}</ref> but ''Spinosaurus'' was likely to have been a generalized and opportunistic predator, possibly a Cretaceous equivalent of large [[grizzly bear]]s, being biased toward fishing, though it undoubtedly [[scavenger|scavenged]] and took many kinds of small or medium-sized prey.<ref name=GSP88>{{cite book |last=Paul |first=G.S. |authorlink=Gregory S. Paul |title=Predatory Dinosaurs of the World |year=1988 |publisher=Simon & Schuster |location=New York |isbn=0-671-61946-2 |chapter=Family Spinosauridae |pages=271–274 }}</ref> A study by Cuff and Rayfield (2013) concluded that bio-mechanical data suggests that ''Spinosaurus'' was not an [[obligate]] piscivore and that its diet was more closely associated with each individual's size.<ref name=cuff13>{{Cite journal | last1 = Cuff | first1 = A. R. | last2 = Rayfield | first2 = E. J. | editor1-last = Farke | editor1-first = Andrew A | doi = 10.1371/journal.pone.0065295 | title = Feeding Mechanics in Spinosaurid Theropods and Extant Crocodilians | journal = PLoS ONE | volume = 8 | issue = 5 | pages = e65295 | year = 2013 | pmid = 23724135| pmc =3665537 }}</ref> The characteristic [[Anatomical terms of location#Other directional terms|rostral]] morphology of ''Spinosaurus'' allowed its jaws to resist bending in the vertical direction, however its jaws were poorly adapted with respect to resisting lateral bending.<ref name=cuff13/>
[[File:Feeding-Mechanics-in-Spinosaurid-Theropods-and-Extant-Crocodilians-pone.0065295.s007.ogv|thumbtime=0:00|thumb|CT scan of partial snout NHMUK 16665, used in a biomechanical study]]
In 2009, Dal Sasso ''et al.''. reported the results of [[X-ray computed tomography]] of the MSNM V4047 snout.<ref name=DalSasso2009>{{cite web |url= http://www2.mnhn.fr/hdt203/info/media/navep1/abstracts.pdf |title= A neurovascular cavity within the snout of the predatory dinosaur ''Spinosaurus'' |last=Dal Sasso |first=C. |author2=Maganuco, S. |author3= Cioffi, A. |date= 26 May 2009 |work = 1st International Congress on North African Vertebrate Palaeontology |publisher= Muséum national d'Histoire naturelle |accessdate=22 September 2010}}</ref> As the [[Foramen|foramina]] on the outside all communicated with a space on the inside of the snout, the authors speculated that ''Spinosaurus'' had [[Mechanoreceptor|pressure receptors]] inside the space that allowed it to hold its snout at the surface of the water to detect swimming prey species without seeing them.<ref name=DalSasso2009/>

A 2010 [[isotope analysis]] by Romain Amiot and colleagues found that [[isotopes of oxygen|oxygen isotope]] ratios of spinosaurid teeth, including teeth of ''Spinosaurus'', indicate semiaquatic lifestyles.<ref name=RMetal10>{{cite journal |last=Amiot |first=R. |author2=Buffetaut, E. |author3=Lécuyer, C. |author4=Wang, X. |author5=Boudad, L. |author6=Ding, Z. |author7=Fourel, F. |author8=Hutt, S. |author9=Martineau, F. |author10=Medeiros, A. |author11=Mo, J. |author12=Simon, L. |author13=Suteethorn, V. |author14=Sweetman, S. |author15=Tong, H. |author16=Zhang, F. |author17=Zhou, Z. |year=2010 |title=Oxygen isotope evidence for semi-aquatic habits among spinosaurid theropods |journal=Geology |volume=38 |issue=2 |pages=139–142 |doi=10.1130/G30402.1}}</ref> Isotope ratios from tooth enamel and from other parts of ''Spinosaurus'' (found in Morocco and Tunisia) and of other predators from the same area such as ''[[Carcharodontosaurus]]'' were compared with isotopic compositions from contemporaneous theropods, turtles, and crocodilians.<ref name=RMetal10/> The study found that ''Spinosaurus'' teeth from five of six sampled localities had oxygen isotope ratios closer to those of turtles and crocodilians when compared with other theropod teeth from the same localities.<ref name=RMetal10/> The authors postulated that ''Spinosaurus'' switched between terrestrial and aquatic habitats to compete for food with large crocodilians and other large theropods respectively.<ref name=RMetal10/>

===Posture===
{{multiple image
| direction = vertical
| width =
| header =
| image1 = Spinosaurus in Japan.jpg
| alt1 = Old hip region
| caption1 = Hip region of an older reconstruction
| image2 = Spinosaurus hip.jpg
| alt2= New hip region
| caption2 = Hip region of the new reconstruction
}}
Although traditionally depicted as a [[Bipedalism|biped]], it has been suggested since the mid-1970s that ''Spinosaurus'' was at least an occasional [[Quadrupedalism|quadruped]].<ref name=DFG82/><ref name=LBH75/> This has been bolstered by the discovery of ''[[Baryonyx]]'', a relative with robust arms.<ref name=DFG00>{{cite book |last=Glut |first=D.F. |authorlink=Donald F. Glut |title=Dinosaurs: The Encyclopedia. 1st Supplement |year=2000 |publisher=McFarland & Company, Inc |location=Jefferson, North Carolina |isbn=0-7864-0591-0 |pages=329–333 |chapter=Spinosaurus}}</ref> Because of the mass of the hypothesized fatty dorsal humps of ''Spinosaurus'', Bailey (1997) was open to the possibility of a quadrupedal posture,<ref name=JBB97/> leading to new restorations of it as such.<ref name=DFG00/> The hypothesis that ''Spinosaurus'' had a typical quadrupedal gait has fallen out of favor, though spinosaurids may have crouched in a quadrupedal posture.<ref name=CM97>{{cite journal |last=Charig |first=A.J. |author2=Milner, A.C. |year=1997 |title=''Baryonyx walkeri'', a fish-eating dinosaur from the Wealden of Surrey |journal=Bulletin of the Natural History Museum, Geology Series |volume=53|pages=11–70}}</ref>

Theropods, including spinosaurids, could not pronate their hands (rotate the forearm so the palm faced the ground),<ref name="carpenter2002">{{Cite journal | doi = 10.1007/BF03043773 | last1 = Carpenter | first1 = K. | year = 2002 | title = Forelimb biomechanics of nonavian theropod dinosaurs in predation | url = | journal = [[Senckenbergiana Lethaea]] | volume = 82 | issue = 1| pages = 59–76 }}</ref> but a resting position on the side of the hand was possible, as shown by fossil prints from an Early Jurassic theropod.<ref name=AMetal09>{{cite journal |last=Milner |first=A.R.C. |author2=Harris, J.D. |author3=Lockley, M.G. |author4=Kirkland, J.I. |author5= Matthews, N.A. |title=Bird-like anatomy, posture, and behavior revealed by an Early Jurassic theropod dinosaur resting trace|journal=PLoS ONE |year=2009 |volume=4 |issue=3 |page=e4591 |doi=10.1371/journal.pone.0004591|url=http://www.plosone.org/article/info:doi/10.1371/journal.pone.0004591 |editor1-last=Harpending |editor1-first=Henry |pmid=19259260 |pmc=2645690|bibcode = 2009PLoSO...4.4591M }}</ref> A 2014 paper describing new material of ''Spinosaurus'', proposed that its legs were too short for it to move effectively on land. The reconstruction used in the study was an extrapolation based on different sized individuals, scaled to what was assumed to be the correct proportions.<ref name="Spinosaurus 2014">{{Cite journal | doi = 10.1126/science.1258750| title = Semiaquatic adaptations in a giant predatory dinosaur| journal = Science| volume = 345| issue = 6204| pages = 1613–1616| year = 2014| last1 = Ibrahim | first1 = N.| last2 = Sereno | first2 = P. C.| last3 = Dal Sasso | first3 = C.| last4 = Maganuco | first4 = S.| last5 = Fabbri | first5 = M.| last6 = Martill | first6 = D. M.| last7 = Zouhri | first7 = S.| last8 = Myhrvold | first8 = N.| last9 = Iurino | first9 = D. A. | pmid=25213375
}}</ref> Palaeontologist John Hutchinson of the Royal Veterinary College of the University of London has expressed scepticism to the new reconstruction, and cautioned that using different specimens can result in inaccurate chimaeras.<ref>{{Cite journal | doi = 10.1038/nature.2014.15901| title = Swimming dinosaur found in Morocco| journal = Nature| year = 2014| last1 = Witze | first1 = A. }}</ref> Scott Hartman also expressed criticism because he believes the legs and the pelvis were inaccurately scaled (27% too short) and don't match the published lengths.<ref name="shartman">{{Cite web|author=Scott Hartman|title=There's something fishy about Spinosaurus|date=12 September 2014|accessdate=20 September 2014|work=skeletaldrawing.com|url=http://www.skeletaldrawing.com/home/theres-something-fishy-about-spinosaurus9112014}}</ref> However, responses from Ibrahim et al. to Mark Witton have been positively received as reliable.<ref>{{Cite web|author=Mark Witton|title=The Spinosaurus hindlimb controversy: a detailed response from the authors|date=22 September 2014|accessdate=22 September 2014|work=markwitton-com.blogspot.de|url=http://markwitton-com.blogspot.de/2014/09/the-spinosaurus-hindlimb-controversy.html?spref=tw}}</ref> The 2015 re-description of ''Sigilmassasaurus''<ref name="Evers 2015"/> Evers et al. 2015 doubted whether the material assigned to ''Spinosaurus'' by Ibrahim et. al. belonged to it.<ref name="Evers 2015"/>

==Paleoecology==
[[File:Spinosaurus with contemporaneous taxa.jpg|thumb|''Spinosaurus'' with contemporaneous aquatic animals]]
The environment inhabited by ''Spinosaurus'' is only partially understood, and covers a great deal of what is now northern Africa. The region of Africa ''Spinosaurus'' is preserved in dates from 112 to 93.5 million years ago.<ref>{{cite book |title=A Geologic Time Scale 2004 |editor1=Gradstein, F.M. |editor2=Ogg, J.G. |editor3=Smith, A.G. |year=2004 |publisher= Cambridge University Press |location=Cambridge and New York |isbn=0-521-78673-8 |page=380}}</ref><ref name=Smith06>{{Cite journal | last1 = Smith | first1 = J.B. | last2 = Lamanna | first2 = M.C. | last3 = Mayr | first3 = H. | last4 = and Lacovara | first4 = K.J. | title = New information regarding the holotype of ''Spinosaurus aegyptiacus'' Stromer, 1915 | journal = Journal of Paleontology | volume = 80 | pages = 400–406 | year = 2006 | doi = 10.1666/0022-3360(2006)080[0400:NIRTHO]2.0.CO;2 |url= | issue = 2 | issn = 0022-3360}}</ref><ref name="Holtz2008"/> A 1996 study concluded from Moroccan fossils that ''Spinosaurus'', ''[[Carcharodontosaurus]]'', and ''[[Deltadromeus]]'' "ranged across north Africa during the late Cretaceous (Cenomanian)."<ref name="serenoetal96">{{cite journal | last1 = Sereno | first1 = PC | last2 = Dutheil | first2 = DB | last3 = Iarochene | first3 = M | last4 = Larsson | first4 = HCE | last5 = Lyon | first5 = GH | last6 = Magwene | first6 = PM | last7 = Sidor | first7 = CA | last8 = Varricchio | first8 = DJ | last9 = Wilson | first9 = JA | year = 1996 | title = Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation | url = | journal = Science | volume = 272 | issue = | pages = 986–991 | doi=10.1126/science.272.5264.986 | pmid=8662584}}</ref> Those ''Spinosaurus'' that lived in the Bahariya Formation of what is now [[Egypt]] may have contended with shoreline conditions on [[mudflat|tidal flats]] and channels, living in [[mangrove]] forests alongside similarly large dinosaurian predators ''[[Bahariasaurus]]'' and ''Carcharodontosaurus'', the [[titanosaur]] [[sauropods]] ''[[Paralititan]]'' and ''[[Aegyptosaurus]]'', [[Crocodylomorpha|crocodylomorphs]], bony and cartilaginous fish, turtles, lizards, and [[plesiosaur]]s.<ref name=JSetal01>{{cite journal |last=Smith |first=J.B. |author2=Lamanna, M.C. |author3=Lacovara, K.J. |author4=Dodson, P. |author5=Smith, J.R. |author6=Poole, J.C. |author7=Giegengack, R. |author8= Attia, Y. |year=2001 |title=A giant sauropod dinosaur from an Upper Cretaceous mangrove deposit in Egypt |journal=Science |volume=292 |issue=5522 |pages=1704–1706 |doi=10.1126/science.1060561 |pmid=11387472|bibcode = 2001Sci...292.1704S }}</ref> In the dry season it might have resorted to preying on [[pterosaur]]s.<ref name="naish, 2012">{{cite book|last=Naish|first=Darren|title=Planet Dinosaur : The Next Generation of Killer Giants|year=2012|publisher=Firefly Books|isbn=978-1-77085-049-1|pages=80–85}}</ref> This situation resembles that in the Late [[Jurassic]] [[Morrison Formation]] of [[North America]], which boasts up to five theropod genera over one tonne in weight, as well as several smaller genera (Henderson, 1998; Holtz ''et al.'', 2004). Differences in head shape and body size among the large North African theropods may have been enough to allow [[Ecological niche|niche]] partitioning as seen among the many different predator species found today in the [[Africa]]n [[savanna]] (Farlow & Pianka, 2002).

==In popular culture==
[[File:Spinosaurus - Museu Blau - 2016 - 01.jpg|thumb|left|Sculpture based on the 2014 reconstruction, [[Museu Blau]], Barcelona]]
''Spinosaurus'' appeared in the 2001 film ''[[Jurassic Park III]]'', replacing ''[[Tyrannosaurus]]'' as the main antagonist.<ref name=variety>{{cite news|title=Jurassic Park III |work=[[Variety (magazine)|Variety]] |url=http://www.variety.com/review/VE1117798505.html?categoryid=31&cs=1 |accessdate=July 9, 2007 | first=Derek | last=Elley | date=July 17, 2001}}</ref> The film's consulting paleontologist [[John R. Horner]] was quoted as saying: "If we base the ferocious factor on the length of the animal, there was nothing that ever lived on this planet that could match this creature [''Spinosaurus'']. Also my hypothesis is that T-rex was actually a scavenger rather than a killer. ''Spinosaurus'' was really the predatory animal."<ref>{{cite news |title= Spinosaurus makes ''T. Rex'' look like a pussycat: When it comes to Jurassic Park III, size does matter |author= Portman, J. |newspaper= Ottawa Citizen |date= 11 July 2001}}</ref> (He has since retracted the statement about T. Rex being a scavenger.) In the film, ''Spinosaurus'' was portrayed as larger and more powerful than ''[[Tyrannosaurus]]'': in a scene depicting a battle between the two resurrected predators, ''Spinosaurus'' emerges victorious by biting the tyrannosaur's neck.<ref>{{cite news|last=Chandler|first=G.|title=A bite-size guide to the dinosaurs of the new movie Jurassic Park III|work=National Geographic World|date=August 2001}}</ref> In the fourth film ''[[Jurassic World]]'', there is a nod to this fight where the T-Rex smashes through the skeleton of a Spinosaurus in the climatic fight near the end of the film.<ref>{{cite news|last=Romano|first=Nick|title=Jurassic World Snuck in A Sweet Nod to Jurassic Park 3|url= http://www.cinemablend.com/new/Jurassic-World-Snuck-Sweet-Nod-Jurassic-Park-3-72074.html| work=CinemaBlend|date=June 2015}}</ref>

''Spinosaurus'' has long been depicted in popular books about dinosaurs, although only recently has there been enough information about spinosaurids for an accurate depiction. After an influential 1955 skeletal reconstruction by Lapparent and Lavocat<ref name=Lapparent>Lapparent, A.F. de; and Lavocat, R. (1955). "Dinosauriens." In: Piveteau, J., editor. ''Traité de Paléontologie. Tome V. La Sortie des Eaux. Naissance de la Tétrapodie. L'Exubérance de la Vie Végétative. La Conquête de l'Air. Amphibiens. Reptiles. Oiseaux.'' Paris: Masson et Cie, pp. 785-962.</ref> based on a 1936 diagram by Stromer,<ref name=Stromer36>{{cite journal |last=Stromer |first=E. |authorlink=Ernst Stromer |year=1936 |title= Ergebnisse der Forschungsreisen Prof. E. Stromers in den Wüsten Ägyptens. VII. Baharije-Kessel und -Stufe mit deren Fauna und Flora. Eine ergänzende Zusammenfassung |journal= Abhandlungen der Bayerischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Abteilung, Neue Folge |volume=33|pages=1–102 |language=German}}</ref> it has been treated as a generalized upright theropod, with a skull similar to that of other large theropods and a sail on its back, even having four-fingered hands.<ref name=DFG00/>

In addition to films, action figures, video games, and books, ''Spinosaurus'' has been depicted on postage stamps such as ones from [[Angola]], [[The Gambia]], and [[Tanzania]].<ref>{{cite news |title=From the past |author= Khatri, V.S. |newspaper=The Hindu |date=9 June 2006 |url=http://www.hindu.com/yw/2006/06/09/stories/2006060900450500.htm |accessdate=12 September 2010}}</ref><ref>{{Cite book |editor1-last= Farlow |editor1-first= J.O. |editor2-last= and Walters |editor2-first= R.H | last1 = Glut | first1 = D.F.| last2 =and Brett-Surman | first2 = M.K.| chapter=Dinosaurs and the media |chapterurl= http://si-pddr.si.edu/dspace/bitstream/10088/8043/1/paleo_1997e_MBS__part2.pdf |title = The Complete Dinosaur | year =2000 | publisher = Indiana University Press | location = Bloomington, IN | isbn = 0-253-21313-4 | pages = 673–706 |accessdate=12 September 2010}}</ref>

==References==
{{Reflist|2}}

==Further reading==
* Glut, D.F. "In search of ''Spinosaurus''." In: ''Jurassic classics: a collection of saurian essays and Mesozoic musings'', pp. 77–85. Jefferson, NC: McFarland, 2001. {{ISBN|0-7864-0961-4}}.
* Nothdurft, W.; and Smith, J. ''The Lost Dinosaurs of Egypt.'' New York: Random House, 2002. {{ISBN|0-375-50795-7}}.
*A Tribute to Ernst Stromer: Hundred Years of the Discovery of ''Spinosaurus aegypticus'': Saubhik Ghosh

==External links==
{{Commons}}
{{wikiquote|Spinosaurus}}
{{wikispecies}}
{{Portal|Dinosaurs}}
*Hartman, Scott. [http://www.skeletaldrawing.com/psgallery/gallery.htm Spinosaur Comparison.] ''SkeletalDrawing.com'', 2006.
*Lloyd, Robin. [https://www.webcitation.org/5mqv3WNdR The Biggest Carnivore: Dinosaur History Rewritten.] ''LiveScience'', 1 March 2006.
*Mortimer, Mickey. [http://archosaur.us/theropoddatabase/Megalosauroidea.htm#Spinosaurusaegyptiacus ''Spinosaurus'' Stromer, 1915.] (List of specimens from The Theropod Database.)
*[[Natural History Museum, London|Natural History Museum]]. [http://www.nhm.ac.uk/nature-online/life/dinosaurs-other-extinct-creatures/dino-directory/detail.dsml?Genus=Spinosaurus Dino Directory: Spinosaurus.]
*Prendergast, John. [http://www.upenn.edu/gazette/0701/prendergast.html Dinosaurs Lost and Found.] ''The Pennsylvania Gazette, the Alumni Magazine of the University of Pennsylvania'', July/Aug 2001.
*[http://discovermagazine.com/2007/jan/paleontology/article_view?b_start:int=1&-C= The Top 8 Paleontology Stories of 2006: 40 New Dinosaur King Rears Its Head.] ''Discover Magazine'', 29 December 2006.

{{Megalosauroidea}}

{{taxonbar}}

[[Category:Early Cretaceous dinosaurs of Africa]]
[[Category:Late Cretaceous dinosaurs of Africa]]
[[Category:Spinosaurids]]
[[Category:Articles containing video clips]]
[[Category:Fossil taxa described in 1915]]
[[Category:Albian genus first appearances]]
[[Category:Cenomanian genus extinctions]]
[[Category:Taxa named by Ernst Stromer]]
[[Category:Paleontology in Egypt]]
[[Category:Paleontology in Morocco]]
[[Category:Lost fossils]]

Triceratops

2017-09-21T16:43:25Z

Wikkler: /* Depiction in popular media */

{{pp-move-indef}}
{{Automatic taxobox
| fossil_range = [[Late Cretaceous]], {{fossil range|68|66|earliest=70}}
| image = LA-Triceratops_mount-2.jpg
| image_width = 250px
| image_caption = Skeletal mount of a ''T. prorsus'' specimen at the [[Natural History Museum of Los Angeles County|Natural History Museum of Los Angeles]]
| authority = [[Othniel Charles Marsh|Marsh]], 1889
| type_species = {{extinct}}'''''Triceratops horridus'''''
| type_species_authority = Marsh, 1889
| subdivision_ranks = [[Species]]
| subdivision =
{{extinct}}''T. horridus'' Marsh, 1889 
{{extinct}}''T. prorsus'' Marsh, 1890
| synonyms ={{collapsible list|bullets = true|title=List
|''[[Agathaumas]]''? [[Edward Drinker Cope|Cope]], 1872
|''[[Polyonax]]''? Cope, 1874
|''Bison alticornis'' Marsh, 1887
|''Sterrholophus'' Marsh, 1891
|''[[Claorhynchus]]''? Cope, 1892
|''Ugrosaurus'' Cobabe & Fastovsky, 1987
|''[[Nedoceratops]]''? Ukrainsky, 2007
|''Diceratus''? [[Octávio Mateus|Mateus]], 2008
|''[[Ojoceratops]]''? Sullivan & Lucas, 2010
|''[[Tatankaceratops]]''? Ott & Larson, 2010}}
}}
[[File:Triceratops BWMK.jpg|thumb|''Triceratops'' life reconstruction]]
[[File:Triceratops-four-skulls-04w.jpg|thumb|''Triceratops'' skulls]]
'''''Triceratops''''' is a [[genus]] of [[herbivorous]] [[Ceratopsidae|ceratopsid]] [[dinosaur]] that first appeared during the late [[Maastrichtian]] stage of the late [[Cretaceous]] [[Period (geology)|period]], about 68 [[mya (unit)|million years ago]] (mya) in what is now [[North America]]. It is one of the last known non-avian dinosaur genera, and became extinct in the [[Cretaceous–Paleogene extinction event]] 66 million years ago.<ref name="scannella2014">{{Cite journal|authors=Scannella, J.B., & Fowler, D.W.|year=2014|title=A stratigraphic survey of ''Triceratops'' localities in the Hell Creek Formation, northeastern Montana (2006–2010)|journal=Geological Society of America Special Papers|volume=503|pages=313–332|doi=10.1130/2014.2503(12) |series=Geological Society of America Special Papers|isbn=978-0-8137-2503-1}}</ref> The term ''Triceratops'', which literally means "three-horned face", is derived from the [[Ancient Greek|Greek]] τρί- (''tri-'') meaning "three", [[wikt:κέρας|κέρας]] (''kéras'') meaning "horn", and [[wikt:ὤψ|ὤψ]] (''ops'') meaning "face".<ref>{{cite book|author1=Liddell, H.G. |author2=R. Scott |lastauthoramp=yes |year=1980|title=Greek-English Lexicon, Abridged Edition |publisher=Oxford University Press, Oxford, UK|isbn=0-19-910207-4}}</ref><ref name="OnlineEtDict">{{cite web|title=triceratops|url=http://www.etymonline.com/index.php?term=triceratops&allowed_in_frame=0|publisher=[[Online Etymology Dictionary]]}}</ref>

Bearing a large bony [[neck frill|frill]] and three [[horn (anatomy)|horns]] on its large four-legged body, and possessing similarities with the modern [[rhinoceros]], ''Triceratops'' is one of the most recognizable of all dinosaurs and the best known ceratopsid. It shared the landscape with and was probably [[predation|preyed upon]] by ''[[Tyrannosaurus]]'',<ref name="erickson1996">{{cite journal|doi=10.1080/02724634.1996.10011297|last1=Erickson|first1=GM|last2=Olson |first2=KH|year=1996|title=Bite marks attributable to ''Tyrannosaurus rex'': preliminary description and implications|journal=Journal of Vertebrate Paleontology|volume=16|issue=1|pages=175–178}}</ref> though it is less certain that the two did battle in the manner often depicted in traditional museum displays and popular images.

The exact placement of the genus ''Triceratops'' within the ceratopsid group has been debated by [[paleontology|paleontologists]]. Two [[species]], ''T. horridus'' and ''T. prorsus'', are considered valid, although many other species have been named. Research published in 2010 suggested that the contemporaneous ''[[Torosaurus]]'', a ceratopsid long regarded as a separate genus, represents ''Triceratops'' in its mature form.<ref name="ScanHorn2010">{{cite journal |last1=Scannella |first1=J. |last2=Horner |first2=J.R. |year=2010 |title=''Torosaurus'' Marsh, 1891, is ''Triceratops'' Marsh, 1889 (Ceratopsidae: Chasmosaurinae): synonymy through ontogeny |journal=Journal of Vertebrate Paleontology |volume=30 |issue=4 |pages=1157–1168 |doi=10.1080/02724634.2010.483632}}</ref><ref>{{cite web|last=Switek|first=Brian|title=New Study Says Torosaurus=Triceratops|url=http://blogs.smithsonianmag.com/dinosaur/2010/07/new-study-says-torosaurustriceratops/|work=Dinosaur Tracking|publisher=Smithsonian.com|accessdate=2 March 2011}}</ref> The view was immediately disputed<ref name="AF2011" /><ref name="longrichfieldstudy">[http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0032623#abstract0], Longrich NR, Field DJ (2012) Torosaurus Is Not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy. ''PLoS ONE'' 7(2): e32623. {{DOI|10.1371/journal.pone.0032623}} {{open access}}</ref><ref name="bbcTriNotToro">[http://www.bbc.co.uk/news/science-environment-17192624], Bowdler, Neil(1 March 2012).''<nowiki>Triceratops and Torosaurus dinosaurs 'two species, not one'</nowiki>''. Retrieved July 29, 2013 from http://www.bbc.co.uk/news/science-environment-17192624</ref> and examination of more fossil evidence is expected to settle the debate.

''Triceratops'' has been documented by numerous remains collected since the genus was first described in 1889, including at least one complete individual skeleton.<ref name="fujiwara2009" /> Paleontologist John Scannella observed: "It is hard to walk out into the [[Hell Creek Formation]] and not stumble upon a ''Triceratops'' weathering out of a hillside." Forty-seven complete or partial skulls were discovered in just that area from 2000 to 2010.<ref name="newsci">{{cite web|url=https://www.newscientist.com/article/mg20727713.500-morphosaurs-how-shapeshifting-dinosaurs-deceived-us.html?full=true |title=Morph-osaurs: How shape-shifting dinosaurs deceived us - life - 28 July 2010 |doi=10.1080/02724634.2010.483632 |publisher=New Scientist |accessdate=2010-08-03}}</ref> Specimens representing life stages from hatchling to adult have been found.<ref name="Lambert93">{{cite book|title=The Ultimate Dinosaur Book|year=1993|author=Lambert, D.|pages=152–167|publisher=Dorling Kindersley, New York|isbn=1-56458-304-X}}</ref>

The functions of the frills and three distinctive facial horns on its head have long inspired debate. Traditionally, these have been viewed as defensive weapons against predators. More recent theories, noting the presence of blood vessels in the skull bones of ceratopsids, find it more probable that these features were primarily used in identification, [[mating|courtship]] and dominance displays, much like the [[antler]]s and horns of modern [[reindeer]], [[mountain goat]]s, or [[rhinoceros beetle]]s.<ref name="Dodhorned">{{cite book|title=The Horned Dinosaurs|year=1996|author=Dodson, P.|publisher=Princeton University Press, Princeton, New Jersey|isbn=0-691-02882-6}}</ref> The theory would find additional support if ''Torosaurus'' was found to be the mature form of ''Triceratops'', as this would mean the frill also developed holes ([[Fenestra (anatomy)|fenestrae]]) as individuals reached maturity, rendering the structure more useful for display than defense.<ref name="ScanHorn2010" />

As the archetypal ceratopsid, ''Triceratops'' is one of the most popular dinosaurs, and has been featured in film, postal stamps, and many other types of media.

==Description==
[[File:Triceratops scale.png|thumb|left|Size of ''T. prorsus'' (orange) and ''T. horridus'' (green) compared to a human.]]
Individual ''Triceratops'' are estimated to have reached about 7.9 to 9.0 [[metre|m]] (25.9–29.5 [[foot (length)|ft]]) in length, {{convert|2.9|to|3.0|m|ft|abbr=on}} in height,<ref name="dinodict">{{cite web|url=http://www.dinodictionary.com/dinos_tpg2.asp|title=T Dinosaurs Page 2 |publisher=DinoDictionary.com |accessdate=2010-08-03}}</ref><ref name="nhm">{{cite web|url=http://internt.nhm.ac.uk/jdsml/nature-online/dino-directory/detail.dsml?Genus=Triceratops |title=Triceratops in The Natural History Museum's Dino Directory |publisher=Internt.nhm.ac.uk |accessdate=2010-08-03}}</ref> and 6.1–12.0 [[tonne]]s (13,000–26,000 [[pound (mass)|lb]]) in [[weight]].<ref name="Alexander">{{cite journal | doi = 10.1111/j.1096-3642.1985.tb00871.x | last1 = Alexander | first1 = R.M. | year = 1985 | title = Mechanics of posture and gait of some large dinosaurs | url = | journal = Zoological Journal of the Linnean Society | volume = 83 | issue = | pages = 1–25}}</ref> The most distinctive feature is their large [[skull]], among the largest of all land animals. The largest known skull (specimen [[Museum of Western Colorado|MWC]] 7584, formerly [[Brigham Young University|BYU]] 12183) is estimated to have been {{convert|2.5|m|ft}} in length when complete,<ref name="ScanHorn2010" /> and could reach almost a third of the length of the entire animal.<ref name="Lambert93" /> A specimen of ''T. horridus'' named Kelsey measured {{convert|24|ft|m|order=flip}} long with a {{convert|6.5|ft|m|order=flip|sigfig=3}} skull, stood about {{convert|7.5|ft|m|order=flip}} tall, and was estimated by the Black Hills institute to weight nearly {{convert|6|t}}.<ref>https://www.bhigr.com/pages/info/info_klsy.htm</ref> A ''Triceratops'' {{convert|8|m|ft}} long has been estimated by [[Gregory S. Paul]] to have massed {{convert|9.3|tonnes}}.<ref name="paul2010" /> It bore a single horn on the snout, above the [[nostril]]s, and a pair of horns approximately {{convert|1|m|ft|abbr=on}} long, with one above each eye.<ref>{{cite news|url=https://news.google.com/newspapers?nid=894&dat=20031118&id=OCNTAAAAIBAJ&sjid=MYIDAAAAIBAJ&pg=6720,2967799|title=Denver museum unveils 7-foot-long, 1,000-pound Triceratops skull |publisher=The Daily Courier |date= November 18, 2003|accessdate=2013-12-26}}</ref> In 2010, paleontologists revealed a fossil (named "Yoshi's Trike," MOR 3027) with {{convert|115|cm|ft|adj=mid|-long}} horn cores, housed and displayed at the [[Museum of the Rockies]] in [[Montana]].<ref>http://msuexponent.com/news/2012/10/04/yoshis-trike-on-display-at-museum-of-the-rockies/</ref><ref>John B. Scannella, Denver W. Fowlera,b, Mark B. Goodwinc, and John R. Horner, [https://www.researchgate.net/publication/278242897_PNAS-2014-Scannella_et_al-Triceratops_strat Evolutionary trends in Triceratops from the Hell Creek Formation, Montana], June 2015.</ref> To the rear of the skull was a relatively short, bony frill, adorned with [[epoccipitals]] in some specimens. Most other ceratopsids had large [[Fenestra (anatomy)|fenestrae]] in their frills, while those of ''Triceratops'' were noticeably solid.<ref>{{cite news|url=https://news.google.com/newspapers?nid=2249&dat=19011024&id=U5I-AAAAIBAJ&sjid=XFoMAAAAIBAJ&pg=5758,3411357 |title=Making A Triceratops. Science Supplies Missing Part! Of Skeleton |publisher=Boston Evening Transcript |date= October 24, 1901|accessdate=2013-12-26}}</ref> ''T. horridus'' can be distinguished from ''T. prorsus'' by having a shallower snout.<ref name="paul2010">{{cite book | url=http://press.princeton.edu/titles/9287.html | title=The Princeton Field Guide to Dinosaurs | publisher=Princeton University Press | last=Paul | first=G. S. | authorlink=Gregory S. Paul | year=2010 | pages=265–267 | isbn=978-0-691-13720-9}}</ref>
[[File:Triceratops by Tom Patker.png|thumb|Life reconstruction of a subadult ''T. horridus'']]
''Triceratops'' species possessed a sturdy build, with strong limbs, short hands with three hooves each, and short feet with four hooves each.<ref name="fujiwara2009">{{cite journal | doi = 10.1671/039.029.0406 | last1 = Fujiwara | first1 = S.-I. | year = 2009 | title = A Reevaluation of the manus structure in ''Triceratops'' (Ceratopsia: Ceratopsidae) | url = | journal = Journal of Vertebrate Paleontology | volume = 29 | issue = 4| pages = 1136–1147 }}</ref> Although certainly [[quadruped]]al, the posture of these dinosaurs has long been the subject of some debate. Originally, it was believed that the front legs of the animal had to be [[Terrestrial locomotion#Posture|sprawling]] at angles from the [[thorax]] in order to better bear the weight of the head.<ref name="Dodhorned" /> This stance can be seen in paintings by [[Charles R. Knight|Charles Knight]] and [[Rudolph F. Zallinger|Rudolph Zallinger]]. [[ichnology|Ichnological]] evidence in the form of [[fossil trackway|trackways]] from horned dinosaurs and recent reconstructions of skeletons (both physical and digital) seem to show that ''Triceratops'' and other ceratopsids maintained an upright stance during normal locomotion, with the elbows flexed and slightly bowed out, in an intermediate state between fully upright and fully sprawling (as in the modern rhinoceros).<ref name="fujiwara2009" /><ref name="CP01">{{cite journal | last1 = Christiansen | first1 = P. | last2 = Paul | first2 = G.S. | year = 2001 | title = Limb bone scaling, limb proportions, and bone strength in neoceratopsian dinosaurs | url =http://gspauldino.com/GaiaNeoceratopsian.pdf | journal = Gaia | volume = 16 | issue = | pages = 13–29 }}</ref><ref name="thompsonholmes2007">{{cite journal |last1=Thompson |first1=S. |last2=Holmes |first2=R. |year=2007 |title=Forelimb stance and step cycle in ''Chasmosaurus irvinensis'' (Dinosauria: Neoceratopsia) |journal=Palaeontologia Electronica |volume=10 |issue=1 |pages=17 p. |url=http://palaeo-electronica.org/2007_1/step/index.html}}</ref><ref name="regaetal2010">{{Cite book |last1=Rega |first1=E. |last2=Holmes |first2=R. |last3=Tirabasso |first3=A. |year=2010 |chapter=Habitual locomotor behavior inferred from manual pathology in two Late Cretaceous chasmosaurine ceratopsid dinosaurs, ''Chasmosaurus irvinensis'' (CMN 41357) and ''Chasmosaurus belli'' (ROM 843) |editors=Ryan, Michael J.; Chinnery-Allgeier, Brenda J.; and Eberth, David A. (editors.) |title=New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium |location=Bloomington and Indianapolis |publisher=Indiana University Press |pages=340–354 |isbn=978-0-253-35358-0}}</ref>

The hands and forearms of ''Triceratops'' retained a fairly primitive structure compared to other quadrupedal dinosaurs such as [[thyreophora]]ns and many [[Sauropoda|sauropods]]. In those two groups, the forelimbs of quadrupedal species were usually rotated so that the hands faced forward with palms backward ("pronated") as the animals walked. ''Triceratops'', like other ceratopsians and the related quadrupedal [[Ornithopoda|ornithopods]], walked with most of their fingers pointing out and away from the body, the primitive condition for dinosaurs also retained by bipedal forms like the [[Theropoda|theropods]]. In ''Triceratops'', the weight of the body was carried by only the first three fingers of the hand, while digits 4 and 5 were vestigial and lacked claws or hooves.<ref name="fujiwara2009" /> The phalangeal formula is 2-3-4-3-1, meaning that the innermost finger of the forelimb has two bones, the next has three, etc.<ref>Martin, A.J. (2006). Introduction to the Study of Dinosaurs. Second Edition. Oxford, Blackwell Publishing. 560 pp. {{ISBN|1-4051-3413-5}}.</ref>

==Discovery and identification==
[[Image:Triceratops alticornis.jpg|thumb|Illustration of specimen YPM 1871E, the horn cores that were erroneously attributed to ''Bison alticornis'', the first named specimen of ''Triceratops'']]
The first named specimen now attributed to ''Triceratops'' is a pair of brow horns attached to a skull roof, found near [[Denver, Colorado]] in the spring of 1887.<ref name="KC06">Carpenter, K. (2006). "''Bison''" ''alticornis'' and O.C. Marsh's early views on ceratopsians. In: Carpenter, K. (ed.). ''Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs'', Indiana University Press, Bloomington and Indianapolis, pp. 349–364. {{ISBN|0-253-34817-X}}.</ref> This specimen was sent to [[Othniel Charles Marsh]], who believed that the [[geologic formation|formation]] from which it came dated from the [[Pliocene]], and that the bones belonged to a particularly large and unusual [[bison]], which he named ''Bison alticornis''.<ref name="KC06" /><ref name="OCM87">{{cite journal | last1 = Marsh | first1 = O.C. | year = 1887 | title = Notice of new fossil mammals | url = http://ajs.library.cmu.edu/books/pages.cgi?call=AJS_1887_034_1887&layout=vol0/part0/copy0&file=00000332 | journal = American Journal of Science | volume = 34 | issue = | pages = 323–331 }}</ref> He realized that there were horned dinosaurs by the next year, which saw his publication of the genus ''[[Ceratops]]'' from fragmentary remains,<ref name="OCM88">{{cite journal | last1 = Marsh | first1 = O.C. | year = 1888 | title = A new family of horned Dinosauria, from the Cretaceous | url = http://ajs.library.cmu.edu/books/pages.cgi?call=AJS_1888_036_1888&layout=vol0/part0/copy0&file=00000493 | journal = American Journal of Science | volume = 36 | issue = | pages = 477–478 }}</ref> but he still believed ''B. alticornis'' to be a Pliocene [[mammal]]. It took a third and much more complete skull to change his mind. The specimen, collected in 1888 by [[John Bell Hatcher]] from the [[Lance Formation]] of [[Wyoming]], was initially described as another species of ''Ceratops''.<ref name="OCM89a">{{cite journal | last1 = Marsh | first1 = O.C. | year = 1889a | title = Notice of new American Dinosauria | url = http://ajs.library.cmu.edu/books/pages.cgi?call=AJS_1889_037_1889&layout=vol0/part0/copy0&file=00000339 | journal = American Journal of Science | volume = 37 | issue = | pages = 331–336 }}</ref> After reflection, Marsh changed his mind and gave it the generic name ''Triceratops'', accepting his ''Bison alticornis'' as another species of ''Ceratops''<ref name="OCM89b">{{cite journal | last1 = Marsh | first1 = O.C. | year = 1889b | title = Notice of gigantic horned Dinosauria from the Cretaceous | url = http://ajs.library.cmu.edu/books/pages.cgi?call=AJS_1889_038_1889&layout=vol0/part0/copy0&file=00000183 | journal = American Journal of Science | volume = 38 | issue = | pages = 173–175 }}</ref> (it would later be added to ''Triceratops''<ref name="HML07" />). The sturdy nature of the animal's skull has ensured that many examples have been preserved as [[fossil]]s, allowing variations between [[species]] and individuals to be studied. ''Triceratops'' remains have subsequently been found in the American states of [[Montana]] and [[South Dakota]] (in addition to Colorado and Wyoming), and in the provinces of [[Saskatchewan]] and [[Alberta]], Canada.

An earlier specimen, also recovered from the Lance Formation, was named ''[[Agathaumas sylvestris]]'' by [[Edward Drinker Cope]] in 1872. Originally identified as a hadrosaur, this specimen consists only of post-cranial remains and is only provisionally considered an example of ''Triceratops''.<ref name="gillette1999">Breithaupt, B.H. (1999). "First Discovery of Dinosaurs in the American West." Pp. 59-65 in Gillette, D.D. (ed.), ''Vertebrate Paleontology In Utah''. Utah Geological Survey. {{ISBN|1-55791-634-9}}, {{ISBN|978-1-55791-634-1}}</ref>

===Species===
[[File:Triceratops holotype.jpg|thumb|left|Type specimen of the type species, ''T. horridus'']]
Within the first decades after ''Triceratops'' was described, various skulls were collected, which varied to a lesser or greater degree from the original ''Triceratops'', named ''T. horridus'' by Marsh (from the [[Latin]] ''horridus''; "rough, rugose", suggesting the roughened texture of those bones belonging to the type specimen, later identified as an aged individual). This variation is unsurprising, given that ''Triceratops'' skulls are large three-dimensional objects, coming from individuals of different ages and both sexes, and which were subjected to different amounts and directions of pressure during fossilization.<ref name="Dodhorned" /> Discoverers would name these as separate species (listed below), and came up with several [[phylogeny|phylogenetic]] schemes for how they were related to each other.

In the first attempt to understand the many species, Lull found two groups, although he did not say how he distinguished them: one composed of ''T. horridus'', ''T. prorsus'', and ''T. brevicornus''; the other of ''T. elatus'' and ''T. calicornis''. Two species (''T. serratus'' and ''T. flabellatus'') stood apart from these groups.<ref name="HML07" /> By 1933, and his revision of the landmark 1907 Hatcher-Marsh-Lull [[monograph]] of all known ceratopsians, he retained his two groups and two unaffiliated species, with a third lineage of ''T. obtusus'' and ''T. hatcheri'' that was characterized by a very small nasal horn.<ref name="RSL33" /> ''T. horridus''-''T. prorsus''-''T. brevicornus'' was now thought to be the most conservative lineage, with an increase in skull size and a decrease in nasal horn size, and ''T. elatus''-''T. calicornis'' was defined by large brow horns and small nasal horn.<ref name="RSL33" /><ref>{{cite journal | last1 = Goussard | first1 = F | year = 2006 | title = The skull of Triceratops in the palaeontology gallery, Muséum national d'Histoire naturelle, Paris | url = | journal = Geodiversitas | volume = 28 | issue = 3| pages = 467–476 }}</ref> C. M. Sternberg made one modification, adding ''T. eurycephalus'' and suggesting that it linked the second and third lineages closer together than they were to the ''T. horridus'' lineage.<ref name="CMS49" /> This pattern was followed until the major studies of the 1980s and 1990s.
[[File:Triceratops prorsus old.jpg|thumb|1896 skeletal restoration of ''T. prorsus'' by [[O.C. Marsh]], based on the holotype skull and referred elements]]
With time, the idea that the differing skulls might be representative of individual variation within one (or two) species gained popularity. In 1986, Ostrom and Wellnhofer published a paper in which they proposed that there was only one species, ''Triceratops horridus''.<ref>{{cite journal | last1 = Ostrom | first1 = J. H. | last2 = Wellnhofer | first2 = P. | year = 1986 | title = The Munich specimen of ''Triceratops'' with a revision of the genus | url = | journal = Zitteliana | volume = 14 | issue = | pages = 111–158 }}</ref> Part of their rationale was that generally there are only one or two species of any large animal in a region (modern examples being the [[African elephant|elephant]] and the [[giraffe]] in modern Africa). To their findings, Lehman added the old Lull-Sternberg lineages combined with maturity and [[sexual dimorphism]], suggesting that the ''T. horridus''-''T. prorsus''-''T. brevicornus'' lineage was composed of females, the ''T.calicornis''-''T.elatus'' lineage was made up of males, and the ''T. obtusus''-''T. hatcheri'' lineage was of [[pathology|pathologic]] old males.<ref name="TML90" /> His reasoning was that males had taller, more erect horns and larger skulls, and females had smaller skulls with shorter, forward-facing horns.

These findings were contested a few years later by Catherine Forster, who reanalyzed ''Triceratops'' material more comprehensively and concluded that the remains fell into two species, ''T. horridus'' and ''T. prorsus'', although the distinctive skull of ''T.'' ("''Nedoceratops''") ''hatcheri'' differed enough to warrant a separate genus.<ref>{{cite journal | doi = 10.1080/02724634.1996.10011313 | last1 = Forster | first1 = C.A. | year = 1996 | title = Species resolution in ''Triceratops'': cladistic and morphometric approaches | url = | journal = Journal of Vertebrate Paleontology | volume = 16 | issue = 2| pages = 259–270 }}</ref> She found that ''T. horridus'' and several other species belonged together, and ''T. prorsus'' and ''T. brevicornus'' stood alone, and since there were many more specimens in the first group, she suggested that this meant the two groups were two species. It is still possible to interpret the differences as representing a single species with sexual dimorphism.<ref name="Dodhorned" /><ref name="TML98">{{cite journal | last1 = Lehman | first1 = T. M. | year = 1998 | title = A gigantic skull and skeleton of the horned dinosaur ''Pentaceratops sternbergi'' from New Mexico | journal = Journal of Paleontology | volume = 72 | issue = 5| pages = 894–906 | jstor = 1306666 }}</ref>

In 2009, John Scannella and Denver Fowler supported the separation of ''T. prorsus'' and ''T. horridus'', and noted that the two species are also separated stratigraphically within the Hell Creek Formation, indicating that they did not live together at the same time.<ref name="scannella&fowler2009">Scannella, J.B. and Fowler, D.W. (2009). "Anagenesis in ''Triceratops'': evidence from a newly resolved stratigraphic framework for the Hell Creek Formation." Pp. 148–149 in ''9th North American Paleontological Convention Abstracts''. Cincinnati Museum Center Scientific Contributions 3.</ref>

====Valid species====
[[File:Triceratops prorsus - IMG 0697.jpg|thumb|''T. prorsus'', [[Carnegie Museum of Natural History]]]]
[[File:Triceratops AMNH 01.jpg|thumb|The skull (AMNH 5116) of this ''T. horridus'' composite specimen was formerly assigned to ''T. elatus'']]
* ''T. horridus'' (Marsh, 1889) (originally ''[[Ceratops]]'') ([[type species]])
* ''T. prorsus'' (Marsh, 1890)

====Synonyms and doubtful species====
The following species are considered ''[[nomen dubium|nomina dubia]]'' ("dubious names"), and are based on remains that are too poor or incomplete to be distinguished from pre-existing ''Triceratops'' species.

* ''T. albertensis'' ([[Charles Mortram Sternberg|C. M. Sternberg]], 1949)
* ''T. alticornis'' ([[Othniel Charles Marsh|Marsh]], 1887 [originally ''[[Bison]]''])
* ''T. brevicornus'' (Hatcher, 1905) (=''T. prorsus'')
* ''T. calicornis'' (Marsh, 1898) (=''T. horridus'')
* ''T. elatus'' (Marsh, 1891) (=''T. horridus'')
* ''T. eurycephalus'' ([[Erich Maren Schlaikjer|Schlaikjer]], 1935)
* ''T. flabellatus'' (Marsh, 1889) (=''T. horridus'')
* ''T. galeus'' (Marsh, 1889)
* ''T. hatcheri'' (Lull, 1907) (contentious; see ''Nedoceratops'' below)
* ''T. ingens'' ([[R. S. Lull|Lull]], 1915)
* ''T. maximus'' ([[Barnum Brown|Brown]], 1933)
* ''T. mortuarius'' ([[Edward Drinker Cope|Cope]], 1874) (''nomen dubium''; originally ''[[Polyonax mortuarius]]'')
* ''T. obtusus'' (Marsh, 1898) (=''T. horridus'')
* ''T. serratus'' (Marsh, 1890) (=''T. horridus'')
* ''T. sulcatus'' (Marsh, 1890)
* ''T. sylvestris'' (Cope, 1872) (''nomen dubium''; originally ''[[Agathaumas sylvestris]]'')

==Classification==
[[File:Triceratops Specimen at the Houston Museum of Natural Science.JPG|thumb|left|Specimen nicknamed "Lane", the most complete specimen known]]
''Triceratops'' is the best known genus of the [[Ceratopsidae]], a family of large North American [[Ceratopsia|horned dinosaurs]]. The exact location of ''Triceratops'' among the ceratopsians has been debated over the years. Confusion stemmed mainly from the combination of short, solid frills (similar to that of [[Centrosaurinae]]), and the long brow horns (more akin to [[Ceratopsidae|Ceratopsinae]], also known as Chasmosaurinae).<ref>{{cite web|url=http://dinosaurios.org/triceratops/|title=What is special about the Triceratops?|publisher=Dinosaurios.org|accessdate=2013-12-26}}</ref> In the first overview of horned dinosaurs, [[R. S. Lull]] hypothesized two lineages, one of ''[[Monoclonius]]'' and ''[[Centrosaurus]]'' leading to ''Triceratops'', the other with ''[[Ceratops]]'' and ''[[Torosaurus]]'', making ''Triceratops'' a centrosaurine as the group is understood today.<ref name="HML07">Hatcher, J. B., Marsh, O. C., and Lull, R. S. (1907) ''The Ceratopsia''. Government Printing Office, Washington, D.C. {{ISBN|0-405-12713-8}}.</ref> Later revisions supported this view, formally describing the first, short-frilled group as Centrosaurinae (including ''Triceratops''), and the second, long-frilled group as Chasmosaurinae.<ref name="RSL33">{{cite journal | last1 = Lull | first1 = R. S. | year = 1933 | title = A revision of the Ceratopsia or horned dinosaurs | url =https://archive.org/details/revisionofcerato33lull | journal = Memoirs of the Peabody Museum of Natural History | volume = 3 | issue = 3| pages = 1–175 |accessdate=20 November 2010 | doi=10.5962/bhl.title.5716}}</ref><ref name="LL15">{{cite journal | last1 = Lambe | first1 = L.M. | year = 1915 | title = On ''Eoceratops canadensis'', gen. nov., with remarks on other genera of Cretaceous horned dinosaurs | url =http://isbndb.com/d/book/on_eoceratops_canadensis_gen_nov_with_remarks_on_other_gener.html | journal = Canada Department of Mines Geological Survey Museum Bulletin | volume = 12 | issue = | pages = 1–49 | isbn= 0-665-82611-7 }}</ref>

In 1949, [[Charles Mortram Sternberg|C. M. Sternberg]] was the first to question this and favoured instead that ''Triceratops'' was more closely related to ''[[Arrhinoceratops]]'' and ''[[Chasmosaurus]]'' based on skull and horn features, making ''Triceratops'' a ceratopsine (chasmosaurine of his usage) genus.<ref name="CMS49">{{cite journal | last1 = Sternberg | first1 = C. M. | year = 1949 | title = The Edmonton fauna and description of a new ''Triceratops'' from the Upper Edmonton member; phylogeny of the Ceratopsidae | url = | journal = National Museum of Canada Bulletin | volume = 113 | issue = | pages = 33–46 }}</ref> He was largely ignored, with [[John Ostrom]],<ref name="Ostrom66">{{cite journal|author=Ostrom, J. H.|year=1966| title=Functional morphology and evolution of the ceratopsian dinosaurs| journal=[[Evolution (journal)|Evolution]]| volume=20| issue=3 | pages = 290–308 | doi=10.2307/2406631|jstor=2406631}}</ref> and later David Norman both placing ''Triceratops'' within Centrosaurinae.<ref>{{cite book |last=Norman |first=David |title=The Illustrated Encyclopaedia of Dinosaurs |year=1985 |publisher=Salamander Books |location=London |isbn=0-517-46890-5}}</ref>
[[File:Smithsonian Museum of Natural History Triceratops.jpg|thumb|First mounted ''T. horridus'' skeleton (the holotype of ''T. "obtusus"''), nicknamed "Hatcher", [[Smithsonian Museum]]]]
[[File:Yoshi-Trike-0006.jpg|thumb|Yoshi's Trike, a specimen with 115 cm long horns cores, and juvenile on display in the [[Museum of the Rockies]] in [[Montana]], USA.]]
Subsequent discoveries and analyses upheld Sternberg's view on the position of ''Triceratops'', with Lehman defining both subfamilies in 1990 and diagnosing ''Triceratops'' as ceratopsine (chasmosaurine of his usage) on the basis of several morphological features. In fact, it fits well into the ceratopsine subfamily, apart from its one feature of a shortened frill.<ref name="TML90">Lehman, T. M. (1990). The ceratopsian subfamily Chasmosaurinae: sexual dimorphism and systematics. in: Carpenter, K., and Currie, P. J. (eds.). ''Dinosaur Systematics: Perspectives and Approaches''. Cambridge University Press, Cambridge, pp. 211–229. {{ISBN|0-521-36672-0}}.</ref> Further research by [[Peter Dodson]], including a 1990 [[cladistics|cladistic]] analysis<ref>Dodson, P., and Currie, P. J. (1990). Neoceratopsia. 593–618. in Weishampel, D. B., Dodson, P., & Osmólska, H. (eds.). ''The Dinosauria''. University of California Press, Berkeley, pp. 593–618. {{ISBN|0-520-06727-4}}.</ref> and a 1993 study using RFTRA (resistant-fit theta-rho analysis),<ref>{{cite journal | last1 = Dodson | first1 = P. | year = 1993 | title = Comparative craniology of the Ceratopsia | url = http://earth.geology.yale.edu/~ajs/1993/11.1993.07Dodson.pdf | journal = American Journal of Science | volume = 293 | issue = | pages = 200–234 | doi = 10.2475/ajs.293.A.200 }}</ref> a [[morphometrics|morphometric technique]] which systematically measures similarities in skull shape, reinforces ''Triceratops''' placement in the ceratopsine subfamily.

The below cladogram follows Longrich (2015), who named a new species of ''Pentaceratops'', and included nearly all species of chasmosaurine.<ref name="longrich15">{{Cite journal | doi = 10.1016/j.cretres.2014.06.011| title = The horned dinosaurs Pentaceratops and Kosmoceratops from the upper Campanian of Alberta and implications for dinosaur biogeography| journal = Cretaceous Research| volume = 51| pages = 292–308| year = 2014| last1 = Longrich | first1 = N. R. }}</ref>
{{clade| style=font-size:85%;line-height:85%
|label1=[[Chasmosaurinae]]
|1={{clade
|1=''[[Mercuriceratops]]''
|2={{clade
|1=''[[Judiceratops]]''
|2={{clade
|1={{clade
|1=''[[Chasmosaurus]]''
|2=''[[Mojoceratops]]'' }}
|2={{clade
|1=''[[Agujaceratops]]''
|2={{clade
|1={{clade
|1=''[[Pentaceratops aquilonius]]''
|2=Williams Fork chasmosaur
|3={{clade
|1=''[[Pentaceratops sternbergii]]''
|2=''[[Utahceratops]]'' }} }}
|2={{clade
|1=''[[Kosmoceratops]]''
|2={{clade
|1={{clade
|1=''[[Anchiceratops]]''
|2=Almond Formation chasmosaur }}
|2={{clade
|1={{clade
|1=''[[Bravoceratops]]''
|2=''[[Coahuilaceratops]]'' }}
|2={{clade
|1=''[[Arrhinoceratops]]''
|label2=[[Triceratopsini]]
|2={{clade
|1=''[[Titanoceratops]]''
|2={{clade
|1=''[[Torosaurus]]''
|2='''''Triceratops'''''}} }} }} }} }} }} }} }} }} }} }} }}
[[File:Laramie-skull.jpg|thumb|left|Skull of specimen DMNH 48617 from the [[Laramie Formation]] of eastern [[Colorado]]. Based on the age of the formation, it may be the oldest ''Triceratops'' known.]]
For many years after its discovery, the evolutionary origins of ''Triceratops'' remained largely obscure. In 1922, the newly discovered ''[[Protoceratops]]'' was seen as its ancestor by [[Henry Fairfield Osborn]],<ref name="Dodhorned" /> but many decades passed before additional findings came to light. Recent years have been fruitful for the discovery of several dinosaurs related to ancestors of ''Triceratops''. ''[[Zuniceratops]]'', the earliest known ceratopsian with brow horns, was described in the late 1990s, and ''[[Yinlong]]'', the first known [[Jurassic]] ceratopsian, in 2005.

These new finds have been vital in illustrating the origins of horned dinosaurs in general, suggesting an [[Asia]]n origin in the Jurassic, and the appearance of truly horned ceratopsians by the beginning of the late Cretaceous in North America.<ref name="Dino2">Dodson, P.; Forster, C.A.; and Sampson, S.D. (2004) ''Ceratopsidae''. In: Weishampel, D. B.; Dodson, P.; and Osmólska, H. (eds.), ''The Dinosauria'' (second edition). University of California Press, Berkeley, pp. 494–513. {{ISBN|0-520-24209-2}}.</ref> As ''Triceratops'' is increasingly shown to be a member of the long-frilled Ceratopsinae subfamily, a likely ancestor may have resembled ''[[Chasmosaurus]]'', which thrived some 5 million years earlier.

In [[phylogenetics|phylogenetic taxonomy]], the genus ''Triceratops'' has been used as a reference point in the definition of Dinosauria; dinosaurs have been designated as all descendants of the [[most recent common ancestor]] of ''Triceratops'' and [[modern birds|Neornithes]] (i.e. modern [[bird]]s).<ref>{{cite journal | last1 = Gauthier | first1 = J. A. | year = 1986 | title = Saurischian monophyly and the origin of birds. The Origin of Birds and the Evolution of Flight, K. Padian (ed.) | url = | journal = Memoirs of the California Academy of Sciences | volume = 8 | issue = | pages = 1–55 }}</ref> Furthermore, the bird-hipped dinosaurs, [[Ornithischia]], have all been designated dinosaurs with a more recent common ancestor to ''Triceratops'' than modern birds.<ref>{{cite journal | last1 = Sereno | first1 = P. C. | year = 1998 | title = A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria | url = | journal = Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen | volume = 210 | issue = 1| pages = 41–83 }}</ref>

==Paleobiology==
[[File:Pasta - triceratops brain.jpg|thumb|A 1905 chart showing the relatively small brain of a ''Triceratops'' (top) and an ''[[Edmontosaurus]]'' (bottom).]]
Although ''Triceratops'' are commonly portrayed as [[herd]]ing animals, there is currently little evidence that they lived in herds. While several other genera of horned dinosaurs are known from [[bonebed]]s preserving bones from two to hundreds or thousands of individuals, to date there is only one documented bonebed dominated by ''Triceratops'' bones: a site in southeastern Montana with the remains of three juveniles. It may be significant that only juveniles were present.<ref name="JCMetal09">{{cite journal |doi=10.1080/02724634.2009.10010382 |last1=Mathews |first1=Joshua C. |last2=Brusatte |first2=Stephen L. |last3=Williams |first3=Scott A. |last4=Henderson |first4=Michael D. |year=2009 |title=The first ''Triceratops'' bonebed and its implications for gregarious behavior |journal=Journal of Vertebrate Paleontology |volume=29 |issue=1 |pages=286–290}}</ref> Another, more recent find may reveal that ''Triceratops'' lived in small family groups. In 2012, a group of three ''Triceratops'' in relatively complete condition, each of varying sizes from a full-grown adult to a small juvenile, were found in Wyoming, near Newcastle. The remains are currently under excavation by paleontologist Peter Larson and a team from the Black Hills Institute. It is believed that the animals were traveling as a family unit, but it remains unknown if the group consists of a mated pair and their offspring, or two females and a juvenile they were caring for. The remains also show signs of predation or scavenging from ''[[Tyrannosaurus]]'', particularly on the largest specimen, with the bones of the front limbs showing breakage and puncture wounds from ''Tyrannosaurus'' teeth.<ref>[http://www.cnn.com/2013/06/03/us/triceratops-found/index.html Triceratops trio unearthed in Wyoming]. CNN.com. Retrieved on 2013-08-25.</ref>

For many years, ''Triceratops'' finds were known only from solitary individuals.<ref name="JCMetal09" /> These remains are very common; for example, [[Bruce Erickson (paleontologist)|Bruce Erickson]], a paleontologist of the [[Science Museum of Minnesota]], has reported having seen 200 specimens of ''T. prorsus'' in the [[Hell Creek Formation]] of [[Montana]].<ref>{{cite journal | last1 = Erickson | first1 = B.R. | year = 1966 | title = Mounted skeleton of ''Triceratops prorsus'' in the Science Museum | url = | journal = Scientific Publications of the Science Museum | volume = 1 | issue = | pages = 1–16 }}</ref> Similarly, [[Barnum Brown]] claimed to have seen over 500 skulls in the field.<ref name="Dodhorned" />{{rp|79}} Because ''Triceratops'' teeth, horn fragments, frill fragments, and other skull fragments are such abundant fossils in the [[Lancian|Lancian faunal stage]] of the late [[Maastrichtian]] ([[late Cretaceous]], 66 mya) Period of western North America, it is regarded as among the dominant herbivores of the time, if not the most dominant herbivore. In 1986, [[Robert Bakker]] estimated it as making up 5/6ths of the large dinosaur fauna at the end of the Cretaceous.<ref name="RTB86">Bakker, R.T. (1986). ''The Dinosaur Heresies: New Theories Unlocking The Mystery of the Dinosaurs and Their Extinction''. William Morrow, New York. {{ISBN|0-14-010055-5}}.</ref>{{rp|438}} Unlike most animals, skull fossils are far more common than [[postcrania]]l bones for ''Triceratops'', suggesting that the skull had an unusually high preservation potential.<ref name="KD94">{{cite book |author=Derstler, K. |year=1994 |editor=Nelson, G. E. |title=The Dinosaurs of Wyoming |series=Wyoming Geological Association Guidebook, 44th Annual Field Conference |chapter=Dinosaurs of the Lance Formation in eastern Wyoming |publisher=Wyoming Geological Association |pages=127–146}}</ref>

''Triceratops'' was one of the last ceratopsian genera to appear before the [[Cretaceous–Paleogene extinction event]]. The related ''[[Torosaurus]]'', and the more distantly related diminutive ''[[Leptoceratops]]'', were also present, though their remains have been rarely encountered.<ref name="Dodhorned" />

===Dentition and diet===
[[File:Tric1.JPG|thumb|Close up of the jaws and teeth]]
''Triceratops'' were [[herbivore|herbivorous]], and because of their low head, their primary food was probably low growth, although they may have been able to knock down taller [[plant]]s with their horns, [[beak]], and bulk.<ref name="Dino2" /><ref>{{cite journal | last1 = Tait | first1 = J. | last2 = Brown | first2 = B. | year = 1928 | title = How the Ceratopsia carried and used their head | url = | journal = Transactions of the Royal Society of Canada | volume = 22 | issue = | pages = 13–23 }}</ref> The [[jaw]]s were tipped with a deep, narrow beak, believed to have been better at grasping and plucking than biting.<ref name="Ostrom66" />

''Triceratops'' [[tooth|teeth]] were arranged in groups called batteries, of 36 to 40 tooth columns, in each side of each jaw with 3 to 5 stacked teeth per column, depending on the size of the animal.<ref name="Dino2" /> This gives a range of 432 to 800 teeth, of which only a fraction were in use at any given time (tooth replacement was continuous and occurred throughout the life of the animal).<ref name="Dino2" /> They functioned by shearing in a vertical to near-vertical orientation.<ref name="Dino2" /> The great size and numerous teeth of ''Triceratops'' suggests that they ate large volumes of fibrous plant material,<ref name="Dino2" /> with some suggesting [[Arecaceae|palms]] and [[cycad]]s,<ref>{{cite journal | last1 = Ostrom | first1 = J. H. | year = 1964 | title = A functional analysis of jaw mechanics in the dinosaur ''Triceratops'' | url = http://www.peabody.yale.edu/scipubs/bulletins_postillas/ypmP088_1964.pdf | format=PDF | accessdate=20 November 2010 | journal = Postilla | volume = 88 | issue = | pages = 1–35 }}</ref><ref>{{cite journal | last1 = Weishampel | first1 = D. B. | year = 1984 | title = Evolution of jaw mechanisms in ornithopod dinosaurs | url = | journal = Advances in Anatomy, Embryology, and Cell Biology | volume = 87 | issue = | pages = 1–110 | pmid = 6464809 | doi=10.1007/978-3-642-69533-9| series = Advances in Anatomy Embryology and Cell Biology | isbn = 978-3-540-13114-4 }}</ref> and others suggesting [[fern]]s, which then grew in prairies.<ref>Coe, M. J.; Dilcher, D. L.; Farlow, J. O.; Jarzen, D. M.; and Russell, D. A. (1987). Dinosaurs and land plants. In: Friis, E. M.; Chaloner, W. G.; and Crane, P. R. (eds.) ''The Origins of Angiosperms and their Biological Consequences'' Cambridge University Press, pp. 225–258. {{ISBN|0-521-32357-6}}.</ref>

===Functions of the horns and frill===
[[File:Triceratops skull frills.jpg|thumb|left|upright|Front view of skull with a prominent [[epoccipital]] fringe, [[Houston Museum of Natural Science]]]]
There has been much speculation over the functions of ''Triceratops''' head adornments. The two main theories have revolved around use in combat, or display in courtship, with the latter thought now to be the most likely primary function.<ref name="Dino2" />

Early on, Lull postulated that the frills may have served as anchor points for the jaw muscles to aid chewing by allowing increased size and thus power for the muscles.<ref>{{cite journal | last1 = Lull | first1 = R. S. | year = 1908 | title = The cranial musculature and the origin of the frill in the ceratopsian dinosaurs | url = | journal = American Journal of Science | volume = 4 | issue = 25| pages = 387–399 |doi = 10.2475/ajs.s4-25.149.387 }}</ref> This has been put forward by other authors over the years, but later studies do not find evidence of large muscle attachments on the frill bones.<ref name="Forster90">Forster, C. A. (1990). The cranial morphology and systematics of ''Triceratops'', with a preliminary analysis of ceratopsian phylogeny. Ph.D. Dissertation. University of Pennsylvania, Philadelphia. 227 pp.</ref>

''Triceratops'' were long thought to have possibly used their horns and frills in combat with predators such as ''[[Tyrannosaurus]]'', the idea being discussed first by [[Charles Hazelius Sternberg|C. H. Sternberg]] in 1917 and 70 years later by Robert Bakker.<ref name="RTB86" /><ref>Sternberg, C. H. (1917). ''Hunting Dinosaurs in the Badlands of the Red Deer River, Alberta, Canada''. Published by the author, San Diego, California, 261 pp.</ref> There is evidence that ''Tyrannosaurus'' did have aggressive head-on encounters with ''Triceratops'', based on partially healed tyrannosaur tooth marks on a ''Triceratops'' brow horn and [[squamosal]]; the bitten horn is also broken, with new bone growth after the break. Which animal was the aggressor is not known.<ref name="JH08">{{cite book |author=Happ, J. |chapter=An analysis of predator-prey behavior in a head-to-head encounter between ''Tyrannosaurus rex'' and ''Triceratops'' |editors=Larson, P.; and Carpenter, K. (editors) |title=Tyrannosaurus rex, the Tyrant King (Life of the Past) |publisher=Indiana University Press |location=Bloomington |year=2008 |pages=355–368 |isbn=0-253-35087-5}}</ref> Since the ''Triceratops'' wounds healed, it is most likely that the ''Triceratops'' survived the encounter and managed to overcome the ''Tyrannosaurus''. Paleontologist [[Peter Dodson]] estimates that if ''Tyrannosaurus'' attacked a bull ''Triceratops'', the ''Triceratops'' had the upper hand and would successfully defend itself by inflicting fatal wounds to the ''Tyrannosaurus'' using its sharp horns.<ref>Dodson, Peter, ''The Horned Dinosaurs'', Princeton Press. p.19</ref> ''Tyrannosaurus'' is also known to have fed on ''Triceratops''. Evidence for this includes a heavily tooth-scored ''Triceratops'' [[ilium (bone)|ilium]] and [[sacrum]].<ref name="erickson1996" />
[[File:Triceratops lesions.jpg|thumb|Examples of [[Periosteal reaction|periosteal reactive]] bone in selected specimens of ''Triceratops'']]
In addition to combat with predators using horns, ''Triceratops'' are classically shown engaging each other in combat with horns locked. While studies show that such activity would be feasible, if unlike that of present-day horned animals,<ref>{{cite journal | last=Farke |first= A. A. |year=2004 |title= Horn Use in ''Triceratops'' (Dinosauria: Ceratopsidae): Testing Behavioral Hypotheses Using Scale Models |url=http://www.nhm.ac.uk/hosted_sites/pe/2004_1/horn/horn.pdf | format=PDF | accessdate=20 November 2010 |journal = Palaeo-electronica | volume=7 |issue= 1|pages= 1–10 }}</ref> there is disagreement about whether they did so. Although pitting, holes, lesions, and other damage on ''Triceratops'' skulls (and the skulls of other ceratopsids) are often attributed to horn damage in combat, a 2006 study finds no evidence for horn thrust injuries causing these forms of damage (for example, there is no evidence of infection or healing). Instead, non-pathological [[bone resorption]], or unknown bone diseases, are suggested as causes.<ref name="TF06">Tanke, D. H, and Farke, A. A. (2006). Bone resorption, bone lesions, and extracranial fenestrae in ceratopsid dinosaurs: a preliminary assessment. in: Carpenter, K. (ed.). ''Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs'' Indiana University Press: Bloomington. pp. 319–347. {{ISBN|0-253-34817-X}}.</ref> A newer study compared incidence rates of skull lesions and [[periosteal reaction]] in ''Triceratops'' and ''[[Centrosaurus]]'' and showed that these were consistent with ''Triceratops'' using its horns in combat and the frill being adapted as a protective structure, while lower pathology rates in ''Centrosaurus'' may indicate visual rather than physical use of cranial ornamentation, or a form of combat focused on the body rather than the head.<ref>{{cite journal | last1 = Farke | first1 = A.A. | last2 = Wolff | first2 = E.D.S. | last3 = Tanke | first3 = D.H. | last4 = Sereno | first4 = Paul| year = 2009 | title = Evidence of Combat in ''Triceratops'' | journal = PLoS ONE | volume = 4 | issue = 1| page = e4252 | doi = 10.1371/journal.pone.0004252 | editor1-last = Sereno | editor1-first = Paul | bibcode = 2009PLoSO...4.4252F | pmid=19172995 | pmc=2617760}} {{open access}}</ref> The frequency of injury was found to be 14% in ''Triceratops''.<ref>{{cite journal | last1 = Peterson | first1 = JE | last2 = Dischler | first2 = C | last3 = Longrich | first3 = NR | year = 2013 | title = Distributions of Cranial Pathologies Provide Evidence for Head-Butting in Dome-Headed Dinosaurs (Pachycephalosauridae) | journal = PLoS ONE | volume = 8 | issue = 7| page = e68620 | doi = 10.1371/journal.pone.0068620 | pmid=23874691 | pmc=3712952}} {{open access}}</ref> The researchers also concluded that the damage found on the specimens in the study was often too localized to be caused by bone disease.<ref>{{cite news|url=https://www.wired.com/wiredscience/2009/01/dinofight/ |title=Scars Reveal How Triceratops Fought – |publisher=Wired.com |date= 2009-01-27|accessdate=2010-08-03 |first=Michael |last=Wall}}</ref> Histological examination reveals that the frill of ''Triceratops'' is composed of fibrolamellar bone<ref>Reid REH (1997) Histology of bones and teeth. In: Currie, PJ and Padian, K, editors. Encyclopedia of Dinosaurs. Academic Press, San Diego, CA. 329–339.</ref> which contains [[fibroblasts]] that play a critical role in wound healing, and are capable of rapidly depositing bone during remodeling.<ref>{{cite journal | last1 = Horner | first1 = JR | last2 = Goodwin | first2 = MB | year = 2009 | title = Extreme Cranial Ontogeny in the Upper Cretaceous Dinosaur Pachycephalosaurus | journal = PLoS ONE | volume = 4 | issue = 10| page = e7626 | doi = 10.1371/journal.pone.0007626 | pmid = 19859556 | pmc=2762616}}</ref><ref>{{cite journal | last1 = Horner | first1 = JR | last2 = Lamm | first2 = E | year = 2011 | title = Ontogeny of the parietal frill of Triceratops: a preliminary histological analysis | url = | journal = Comptes Rendus Palevol | volume = 10 | issue = | pages = 439–452 | doi=10.1016/j.crpv.2011.04.006}}</ref>
[[File:UCMP Triceratops right.JPG|left|thumb|Juvenile and adult skulls — the juvenile skull is about the size of an adult human head]]
One skull, assigned to ''Triceratops'', was observed to have a hole in the [[Jugal bone#In dinosaurs|jugal]] which appears to be a puncture wound that was sustained while this individual was still alive. This is supported by signs of healing that are present in the bone around the supposed wound. When examined closely, the hole in the bone has a diameter that is very similar to diameter of the distal end of a ''Triceratops'' horn. This, and other apparent healed wounds in the skulls of ceratopsians, has been cited as evidence of non-fatal intraspecific competition in these dinosaurs.<ref>{{cite journal | last1 = Farlow | first1 = J. O. | last2 = Dodson | first2 = P. | year = 1975 | title = The behavioral significance of frill and horn morphology in ceratopsian dinosaurs | url = | journal = Evolution | volume = 29 | issue = 2| pages = 353–361 | doi=10.2307/2407222| jstor = 2407222 }}</ref><ref>Martin, A.J. (2006). Introduction to the Study of Dinosaurs. Second Edition. Oxford, Blackwell Publishing. pg. 299-300. {{ISBN|1-4051-3413-5}}.</ref>

The large frill also may have helped to increase body area to [[thermoregulation|regulate body temperature]].<ref>{{cite journal|author=Wheeler, P.E.|year=1978|title=Elaborate CNS cooling structures in large dinosaurs|journal=[[Nature (journal)|Nature]]|issue= 5679|pages=441–443|doi=10.1038/275441a0 | volume = 275|pmid=692723|bibcode=1978Natur.275..441W}}</ref> A similar theory has been proposed regarding the plates of ''[[Stegosaurus]]'',<ref>{{cite journal|author=Farlow, J. O., Thompson, C. V., and Rosner, D. E.|year=1976| title=Plates of the dinosaur ''Stegosaurus'': Forced convection heat loss fins?| journal=[[Science (magazine)|Science]]| volume=192 | pages = 1123–5| doi=10.1126/science.192.4244.1123| pmid=17748675|issue=4244|bibcode=1976Sci...192.1123F}}</ref> although this use alone would not account for the bizarre and extravagant variation seen in different members of the [[Ceratopsidae]].<ref name="Dino2" /> This observation is highly suggestive of what is now believed to be the primary function, display.

The theory of their use in sexual display was first proposed by Davitashvili in 1961 and has gained increasing acceptance since.<ref name="TML90" /><ref name="Forster90" /><ref name="Davitashvili61">{{cite book|title=Teoriya Polovogo Otbora (Theory of Sexual Selection)|year=1961|author=Davitashvili, L. Sh.|page=538|publisher=Izdatel'stvo Akademii nauk SSSR}}</ref> Evidence that visual display was important, either in courtship or in other social behavior, can be seen in the fact that horned dinosaurs differ markedly in their adornments, making each species highly distinctive. Also, modern living creatures with such displays of horns and adornments use them in similar behavior.<ref>{{cite journal|author1=Farlow, J.O. |author2=Dodson, P. |lastauthoramp=yes |year=1975|title=The behavioral significance of frill and horn morphology in ceratopsian dinosaurs|journal=[[Evolution (journal)|Evolution]]|volume=29|issue=2 | page = 353|doi=10.2307/2407222|jstor=2407222}}</ref> A 2006 study of the smallest ''Triceratops'' skull, ascertained to be a juvenile, shows the frill and horns developed at a very early age, predating sexual development and thus probably important for visual communication and species recognition in general.<ref>{{cite journal|author1=Goodwin, M.B. |author2=Clemens, W.A. |author3=Horner, J.R. |author4=Padian, K. |last-author-amp=yes | url=http://www.ucmp.berkeley.edu/people/mbg/Goodwin_et_al_2006.pdf|format=PDF| title=The smallest known ''Triceratops'' skull: new observations on ceratopsid cranial anatomy and ontogeny| journal=Journal of Vertebrate Paleontology| volume=26| issue=1| pages = 103–112 | doi = 10.1671/0272-4634(2006)26[103:TSKTSN]2.0.CO;2| year=2006|issn=0272-4634}}</ref> The use of the exaggerated structures in dinosaurs as species identification has been questioned, as no such function exists for structures in modern species.<ref>{{cite journal|author=Hone, D. W. E.; Naish, D.|year=2013|title=The ‘species recognition hypothesis’ does not explain the presence and evolution of exaggerated structures in non-avialan dinosaurs|journal=Journal of Zoology|volume=290|issue=3|pages=172–180|doi=10.1111/jzo.12035}}</ref>

===Growth and ontogeny===
[[File:Triceratops ontogeny.jpg|thumb|Skull growth series]]
In 2006, the first extensive ontogenetic study of ''Triceratops'' was published in the journal [[Proceedings of the Royal Society]]. The study, by [[John R. Horner]] and [[Mark Goodwin]], found that individuals of ''Triceratops'' could be divided into four general ontogenetic groups, babies, juveniles, subadults, and adults. With a total number of 28 skulls studied, the youngest was only {{convert|38|cm|in|abbr=on}} long. 10 of the 28 skulls could be placed in order in a growth series with one representing each age. Each of the four growth stages were found to have identifying features. Multiple ontogenetic trends were discovered, including the size reduce of the epoccipitals, development and reorientation of postorbital horns, and hollowing out of the horns.<ref name="horner2006">{{cite journal|last=Horner|first=J.R.|last2=Goodwin|first2=M.B.|year=2006|title=Major cranial changes during ''Triceratops'' ontogeny|journal=Proceedings of the Royal Society B: Biological Sciences|volume=273|issue=1602|pages=2757–2761|doi=10.1098/rspb.2006.3643|pmc=1635501|pmid=17015322}}</ref>

====''Torosaurus'' as growth stage of ''Triceratops''====

{{Main article|Torosaurus}}
''[[Torosaurus]]'' is a ceratopsid genus first identified from a pair of skulls in 1891, two years after the identification of ''Triceratops''. The ''Torosaurus'' genus resembles ''Triceratops'' in geological age, distribution, anatomy and physical size and it has been recognised as a close relative.<ref>{{cite book | last1=Farke |first1= A. A. |year=2006 |chapter= Cranial osteology and phylogenetic relationships of the chasmosaurine ceratopsid ''Torosaurus latus'' |editor1-last= Carpenter |editor1-first= K. |title=Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs |publisher= Indiana University Press |location= Bloomington |pages=235–257 | isbn=978-0-253-34817-3 }}</ref> Its distinguishing features are an elongated skull and the presence of two fenestrae, or holes, in the frill. Paleontologists investigating dinosaur [[ontogeny]] (growth and development of individuals over the life span) in the [[Hell Creek Formation]], Montana, US, have recently presented evidence that the two represent a single genus.
[[File:Torosaurus and Triceratops.tif|thumb|upright|A, ''Triceratops prorsus'' holotype YPM 1822 and B, ''Torosaurus latus'' ANSP 15192]]
John Scannella, in a paper presented in [[Bristol|Bristol, UK]] at the conference of the [[Society of Vertebrate Paleontology]] (25 September 2009) reclassified ''Torosaurus'' as especially mature ''Triceratops'' individuals, perhaps representing a single sex. Jack Horner, Scannella's mentor at Bozeman Campus, [[Montana State University]], noted that ceratopsian skulls consist of metaplastic bone. A characteristic of metaplastic bone is that it lengthens and shortens over time, extending and resorbing to form new shapes. Significant variety is seen even in those skulls already identified as ''Triceratops'', Horner said, "where the horn orientation is backwards in juveniles and forward in adults". Approximately 50% of all subadult ''Triceratops'' skulls have two thin areas in the frill that correspond with the placement of "holes" in ''Torosaurus'' skulls, suggesting that holes developed to offset the weight that would otherwise have been added as maturing ''Triceratops'' individuals grew longer frills.<ref name="growth09">{{cite web|url=http://www.sciencedaily.com/releases/2009/10/091031002314.htm|title=New Analyses Of Dinosaur Growth May Wipe Out One-third Of Species|date=2009-10-31|work=Science News|publisher=ScienceDaily.com|accessdate=2009-11-03}}</ref> A paper describing these findings in detail was published in July 2010 by Scannella and Horner. It formally argues that ''Torosaurus'' and the similar contemporary ''Nedoceratops'' are synonymous with ''Triceratops''.<ref name="ScanHorn2010" />

The assertion ignited debate. Andrew Farke had in 2006 stressed that, apart from the frill, no systematic differences could be found between ''Torosaurus'' and ''Triceratops''.<ref name="Farke2006">Farke, A. A. "Cranial osteology and phylogenetic relationships of the chasmosaurine ceratopsid ''Torosaurus latus''", pp. 235-257. In K. Carpenter (ed.). ''Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs''. Indiana Univ. Press (Bloomington), 2006.</ref> He nevertheless disputed Scannella's conclusion by arguing in 2011 that the proposed morphological changes required to "age" a ''Triceratops'' into a ''Torosaurus'' would be without precedent among ceratopsids. Creatures would require the growth of [[epoccipital]]s, reversion of bone texture from adult to immature forms back to adult, and growth of frill holes at a later stage than usual.<ref name="AF2011" /> A study by Nicholas Longrich and Daniel Field analyzed 35 specimens of both ''Triceratops'' and ''Torosaurus''. The authors concluded that ''Triceratops'' individuals too old to be considered immature forms are represented in the fossil record, as are ''Torosaurus'' individuals too young to be considered fully mature adults. The synonymy of ''Triceratops'' and ''Torosaurus'' cannot be supported, they said, without more convincing intermediate forms than Scannella and Horner initially produced. Scannella's ''Triceratops'' specimen with a hole on its frill, they argued, could represent a diseased or malformed individual rather than a transitional stage between an immature ''Triceratops'' and mature ''Torosaurus'' form.<ref name="longrichfieldstudy" /><ref name="bbcTriNotToro" />

Given the abundance of fossils, particularly of ''Triceratops'', additional field discoveries are expected to settle the debate in time.

====Other genera as growth stages of ''Triceratops''====

{{Main article|Nedoceratops}}
[[File:Nedoceratops skull, PLoS ONE.png|thumb|Comparisons between the skulls of ''Triceratops'' and ''[[Nedoceratops]]'']]
Opinion has varied on the validity of a separate genus for ''[[Nedoceratops]]''. John Scannella and [[Jack Horner (paleontologist)|Jack Horner]] regarded it as an intermediate growth stage between ''Triceratops'' and ''Torosaurus''.<ref name="ScanHorn2010" /><ref>{{Cite journal | last1 = Scannella | first1 = J. B. | last2 = Horner | first2 = J. R. | editor1-last = Claessens | editor1-first = Leon | title = 'Nedoceratops': An Example of a Transitional Morphology | journal = PLoS ONE | volume = 6 | issue = 12 | pages = e28705 | year = 2011 | doi = 10.1371/journal.pone.0028705| pmid = 22194891 | bibcode = 2011PLoSO...628705S | pmc=3241274}} {{open access}}</ref> Andrew Farke, in his 2011 redescription of the only known skull, concluded that it was an aged individual of its own valid [[taxon]], ''Nedoceratops hatcheri''.<ref name="AF2011">{{cite journal |last=Farke |first=Andrew A. |year=2011 |title= Anatomy and taxonomic status of the chasmosaurine ceratopsid ''Nedoceratops hatcheri'' from the Upper Cretaceous Lance Formation of Wyoming, U.S.A |url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0016196 |journal=PLoS ONE |volume=6 |issue=1 |pages=e16196 |doi=10.1371/journal.pone.0016196 |editor1-last=Claessens |editor1-first=Leon |pmid=21283763 |pmc=3024410|bibcode=2011PLoSO...616196F }} {{open access}}</ref> Nicholas Longrich and Daniel Fields also did not consider it a transition between ''Torosaurus'' and ''Triceratops'', suggesting that the frill holes were pathological.<ref name="bbcTriNotToro" />

As described above, John Scannella had argued in 2010 that ''Nedoceratops'' should be considered a synonym of ''Triceratops''.<ref name="ScanHorn2010" /> Andrew Farke (2011) maintained that it represents a valid distinct genus.<ref name="AF2011" /> Nick Longrich agreed with Scannella about ''Nedoceratops'' and made a further suggestion: that the recently described ''[[Ojoceratops]]'' was likewise a synonym. The fossils, he argued, are indistinguishable from the ''T. horridus'' specimens that were previously attributed to the defunct species ''T. serratus''.

Longrich observed that another newly described genus, ''[[Tatankaceratops]]'', displayed a strange mix of characteristics already found in adult and juvenile ''Triceratops''. Rather than representing a distinct genus, ''Tatankaceratops'' could as easily represent a dwarf ''Triceratops'' or a ''Triceratops'' individual with a developmental disorder that caused it to stop growing prematurely.<ref name="Longrich">{{Cite journal|author=Nicholas R. Longrich |year=2011 |title=''Titanoceratops ouranous'', a giant horned dinosaur from the Late Campanian of New Mexico |url=http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6WD3-51TPVM1-1&_user=10&_coverDate=12%2F29%2F2010&_rdoc=1&_fmt=high&_orig=browse&_origin=browse&_zone=rslt_list_item&_srch=doc-info(%23toc%236755%239999%23999999999%2399999%23FLA%23display%23Articles)&_cdi=6755&_sort=d&_docanchor=&_ct=21&_acct=C000050221&_version=1&_urlVersion=0&_userid=10&md5=8f5e3845b1c44a1dc08c04b09952a44a&searchtype=a |journal=Cretaceous Research |volume=32 |issue= 3|pages= 264–276|doi=10.1016/j.cretres.2010.12.007}}</ref>

==Paleoecology==
[[File:Upper Cretaceous Hell Creek dinosaur census.png|thumb|upright|left|Pie chart of the time averaged census for large-bodied dinosaurs from the entire Hell Creek Formation in the study area]]
''Triceratops'' lived during the Late Cretaceous of North America, its fossils have come from the [[Evanston Formation]], [[Scollard Formation]], [[Laramie Formation]], [[Lance Formation]], [[Denver Formation]], and [[Hell Creek Formation]].<ref name="weishampel"/> These fossil formations date back to the time of the [[Cretaceous-Paleogene Extinction Event]], and has been dated to 66 ± 0.07 million years ago.<ref name="hcfage">{{Cite journal | doi = 10.1016/j.epsl.2011.03.008| title = Astronomical calibration of the Maastrichtian (Late Cretaceous)| journal = Earth and Planetary Science Letters| volume = 305| issue = 3–4| pages = 328–340| year = 2011| last1 = Husson | first1 = D. E. | last2 = Galbrun | first2 = B. | last3 = Laskar | first3 = J. | last4 = Hinnov | first4 = L. A. | last5 = Thibault | first5 = N. | last6 = Gardin | first6 = S. | last7 = Locklair | first7 = R. E. | bibcode = 2011E&PSL.305..328H}}</ref> Many animals and plants have been found in these formations, but mostly from the Lance Formation and Hell Creek Formation.<ref name="weishampel"/>

Theropods from these formations include genera of [[tyrannosauridae|tyrannosaurids]], [[ornithomimidae|ornithomimids]], [[troodontidae|troodontids]],<ref name="weishampel">{{cite book|last=Weishampel|first=D.B.|last2=Dodson|first2=Peter|last3=Osmólska|first3=H.|year=2004|title=The Dinosauria (Second ed.)|location=Berkeley|publisher=University of California Press|pages=861|isbn=0-520-24209-2}}</ref> [[Avialae|avialans]],<ref name="pnas">{{Cite journal | doi = 10.1073/pnas.1110395108| title = Mass extinction of birds at the Cretaceous-Paleogene (K-Pg) boundary| journal = Proceedings of the National Academy of Sciences| volume = 108| issue = 37| pages = 15253–15257| year = 2011| last1 = Longrich | first1 = N. R.| last2 = Tokaryk | first2 = T.| last3 = Field | first3 = D. J.| bibcode = 2011PNAS..10815253L | pmid=21914849 | pmc=3174646}}</ref> [[caenagnathidae|caenagnathids]],<ref name=Anzu>{{Cite journal | doi = 10.1371/journal.pone.0092022| pmid = 24647078| title = A New Large-Bodied Oviraptorosaurian Theropod Dinosaur from the Latest Cretaceous of Western North America| journal = PLoS ONE| volume = 9| issue = 3| pages = e92022| year = 2014| last1 = Lamanna | first1 = M. C. | last2 = Sues | first2 = H. D. | last3 = Schachner | first3 = E. R. | last4 = Lyson | first4 = T. R. | bibcode = 2014PLoSO...992022L | pmc=3960162}} {{open access}}</ref> and [[dromaeosauridae|dromaeosaurids]]. ''[[Acheroraptor]]'' and ''[[Dakotaraptor]]'' are dromaeosaurids from the Hell Creek Formation. Indeterminate dromaeosaurs are known from other fossil formations. Common teeth previously referred to ''[[Dromaeosaurus]]'' and ''[[Saurornitholestes]]'' later were considered to be ''Acheroraptor''.<ref name=Acheroraptor>{{Cite journal | last1 = Evans | first1 = D. C. | last2 = Larson | first2 = D. W. | last3 = Currie | first3 = P. J. | doi = 10.1007/s00114-013-1107-5 | title = A new dromaeosaurid (Dinosauria: Theropoda) with Asian affinities from the latest Cretaceous of North America | journal = Naturwissenschaften | volume = 100 | issue = 11 | pages = 1041–1049 | year = 2013 | pmid = 24248432| pmc = | bibcode = 2013NW....100.1041E }}</ref> The tyrannosaurids from the formation are ''[[Nanotyrannus]]'' and ''Tyrannosaurus'', although the former might be a junior synonym of the latter. Among ornithomimids are the genera ''[[Struthiomimus]]'' as well as ''[[Ornithomimus]]'',<ref name="weishampel"/> although an undescribed animal named "[[Orcomimus]]" could be from the formation.<ref name="tb1997">{{cite journal|last=Triebold|first=M.|year=1997|title=The Sandy site: Small dinosaurs from the Hell Creek Formation of South Dakota|editor-last=Wolberg|editor-first=D.|editor2-last=Stump|editor2-first=E.|editor3-last=Rosenberg|editor3-first=G.|journal=Dinofest International: Proceedings of a Symposium|publisher=Arizona State University Academy of Natural Science|pages=245–248}}</ref> Troodontids are only represented by ''[[Pectinodon]]'' and ''[[Paronychodon]]'' in the Hell Creek Formation; with a possible species of ''[[Troodon]]'' from the Lance Formation. One species of coelurosaur is known from Hell Creek and similar formations by a single species, ''[[Richardoestesia]]''. Only three [[oviraptorosauria|oviraptorosaurs]] are from the Hell Creek Formation, ''[[Anzu (dinosaur)|Anzu]]'', ''[[Leptorhynchos (dinosaur)|Leptorhynchos]]''<ref name=Anzu/> and a giant species of caenagnathid, very similar to ''[[Gigantoraptor]]'', from South Dakota. However, only fossilized foot prints were discovered.<ref>{{cite web|url=http://rmdrc.blogspot.com/2013/12/giant-oviraptor-tracks-from-hell-creek.html|last1=Maltese|first1=Anthony|title=Giant Oviraptor Tracks from the Hell Creek|publisher=RMDRC paleo lab|accessdate=17 December 2013}}</ref> The avialans known from the formation are ''[[Avisaurus]]'',<ref name="weishampel"/> multiple species of ''[[Brodavis]]'',<ref name="brodavis">{{Cite journal | doi = 10.1016/j.palwor.2012.02.005| title = A new evolutionary lineage of diving birds from the Late Cretaceous of North America and Asia| journal = [[Palaeoworld]]| volume = 21| pages = 59–63| year = 2012| last1 = Martin | first1 = L. D. | last2 = Kurochkin | first2 = E. N. | last3 = Tokaryk | first3 = T. T. }}</ref> and several other species of [[Hesperornithiform|hesperornithoforms]], as well as several species of true [[bird]]s including ''[[Cimolopteryx]]''.<ref name="pnas"/>
[[File:Hell Creek dinosaurs and pterosaurs by durbed.jpg|thumb|''Triceratops'' and other animals of the Hell Creek Formation]]

[[Ornithischia]]ns are abundant in the Scollard Lance, Laramie, Lance, Denver, and Hell Creek Formation. The main groups of ornithischians are [[ankylosauria]]ns, [[ornithopoda|ornithopods]], [[ceratopsia]]ns, and [[pachycephalosauria]]ns. Three ankylosaurians are known, ''[[Ankylosaurus]]'', ''[[Denversaurus]]'', and possibly a species of ''[[Edmontonia]]'' or an undescribed genus. Multiple genera of ceratopsians are known from the formation other than ''Triceratops'', the [[leptoceratopsidae|leptoceratopsid]] ''[[Leptoceratops]]'', and the [[chasmosaurine]] [[ceratopsidae|ceratopsids]] ''[[Torosaurus]]'',<ref name="weishampel" />''[[Nedoceratops]]'' and ''[[Tatankaceratops]]''.<ref name="tatanka">{{cite book|last=Ott|first=C.J.|last2=Larson|first2=P.L.|year=2010|chapter=A New, Small Ceratopsian Dinosaur from the Latest Cretaceous Hell Creek Formation, Northwest South Dakota, United States: A Preliminary Description|editor-last=Ryan|editor-first=M.J.|editor2-last=Chinnery-Allgeier|editor2-first=B.J.|editor3-last=Eberth|editor3-first=D.A.|title=New Perspectives on Horned Dinosaurs: The Royal Tyrrell Museum Ceratopsian Symposium|location=Bloomington|publisher=Indiana University Press|pages=656}}</ref> Ornithopods are common in the Hell Creek Formation, and are known from several species of the ornithopod ''[[Thescelosaurus]]'', and the [[hadrosauridae|hadrosaurids]] ''[[Edmontosaurus]]'',<ref name="weishampel"/><ref name="hcfhadro">{{Cite journal | doi = 10.1371/journal.pone.0025186| pmid = 21969872| title = Cranial Growth and Variation in Edmontosaurs (Dinosauria: Hadrosauridae): Implications for Latest Cretaceous Megaherbivore Diversity in North America| journal = PLoS ONE| volume = 6| issue = 9| pages = e25186| year = 2011| last1 = Campione | first1 = N. S. E. | last2 = Evans | first2 = D. C. | pmc=3182183| bibcode = 2011PLoSO...625186C}} {{open access}}</ref> and a possible species of ''[[Parasaurolophus]]''. Several pachycephalosaurians have been found in the Hell Creek Formation and in similar formations. Among them are the derived [[pachycephalosauridae|pachycephalosaurids]] ''[[Stygimoloch]]'',<ref name="weishampel"/> ''[[Dracorex]]'',<ref name="bakker">{{cite journal|last=Bakker|first=R.T.|last2=Sullivan|first2=R.M.|last3=Porter|first3=V.|last4=Larson|first4=P.|last5=Saulsbury|first5=S.J.|year=2006|title=''Dracorex hogwartsia'', n. gen., n. sp., a spiked, flat-headed pachycephalosaurid dinosaur from the Upper Cretaceous Hell Creek Formation of South Dakota|editor-last=Lucas|editor-first=S.G.|editor2-last=Sullivan|editor2-first=R.M.|journal=Late Cretaceous vertebrates from the Western Interior|series=New Mexico Museum of Natural History and Science Bulletin|volume=35|pages=331–345|url=http://econtent.unm.edu/cdm/ref/collection/bulletins/id/710}}</ref> ''[[Pachycephalosaurus]]'',<ref name="weishampel"/> ''[[Sphaerotholus]]'', and an undescribed specimen from North Dakota. The first two might be junior synonyms of ''Pachycephalosaurus''.

Mammals are plentiful in the Hell Creek Formation. Groups represented include [[multituberculata|multituberculates]], [[metatheria]]ns, and [[eutheria]]ns. The multituberculates represented include ''[[Paracimexomys]]'',<ref name=kj2004>[[Zofia Kielan-Jaworowska]], Richard L. Cifelli, and Zhe-Xi Luo, ''Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure'', Columbia University Press, New York, 2004 {{ISBN|0-231-11918-6}}, p. 98-99</ref> the [[cimolomyidae|cimolomyids]] ''[[Paressonodon]]'',<ref name="mammal">{{Cite journal | doi = 10.1666/12041| title = Mammals across the K/Pg boundary in northeastern Montana, U.S.A.: Dental morphology and body-size patterns reveal extinction selectivity and immigrant-fueled ecospace filling| journal = Paleobiology| volume = 39| issue = 3| pages = 429–469| year = 2013| last1 = Wilson | first1 = G. P. }}</ref> ''[[Meniscoessus]]'', ''[[Essonodon]]'', ''[[Cimolomys]]'', ''[[Cimolodon]]'', and ''[[Cimexomys]]''; and the [[neoplagiaulacidae|neoplagiaulacids]] ''[[Mesodma]]'', and ''[[Neoplagiaulax]]''. The [[alphadontidae|alphadontids]] ''[[Alphadon]]'', ''[[Protalphodon]]'', and ''[[Turgidodon]]'', [[pediomyidae|pediomyids]] ''[[Pediomys]]'',<ref name=kj2004/> ''[[Protolambda]]'', and ''[[Leptalestes]]'',<ref name="mammalshcf">{{Cite journal | doi = 10.1007/s10914-011-9162-1| title = Protungulatum, Confirmed Cretaceous Occurrence of an Otherwise Paleocene Eutherian (Placental?) Mammal| journal = Journal of Mammalian Evolution| volume = 18| issue = 3| pages = 153–161| year = 2011| last1 = Archibald | first1 = J. D. | last2 = Zhang | first2 = Y. | last3 = Harper | first3 = T. | last4 = Cifelli | first4 = R. L. }}</ref> the [[stagodontidae|stagodontid]] ''[[Didelphodon]]'',<ref name=kj2004/> the [[Deltatheridiidae|deltatheridiid]] ''[[Nanocuris]]'', the [[herpetotheridiidae|herpetotheriid]] ''[[Nortedelphys]]'',<ref name="mammal"/> and the [[glasbiidae|glasbiid]] ''[[Glasbius]]'' all represent metatherians of the Hell Creek Formation. A few eutherians are known, being represented by ''[[Alostera]]'',<ref name=kj2004/> ''[[Protungulatum]]'',<ref name="mammalshcf"/> the [[cimolestidae|cimolestids]] ''[[Cimolestes]]'' and ''[[Batodon]]'', the [[Gypsonictopsidae|gypsonictopsid]] ''[[Gypsonictops]]'', and the possible [[Nyctitheriidae|nyctitheriid]] ''[[Paranyctoides]]''.<ref name=kj2004/>

==Depiction in popular media==
{{See also|List of cloned animals in Jurassic Park|Cultural depictions of dinosaurs}}
[[File:Knight Triceratops.jpg|thumb|1901 illustration by [[Charles R. Knight]]]]
''Triceratops'' (the species are not identified) is the official [[state fossil]] of [[South Dakota]],<ref>{{cite web|author=State of South Dakota |title=Signs and Symbols of South Dakota..... |url=http://www.state.sd.us/state/sdsym.htm |accessdate=2007-01-20 |deadurl=yes |archiveurl=https://web.archive.org/web/20080220004837/http://www.state.sd.us/state/sdsym.htm |archivedate=2008-02-20 |df= }}</ref> and the official state dinosaur of [[Wyoming]].<ref>{{cite web | author = State of Wyoming | title = State of Wyoming – General Information | url= http://wyoming.gov/general/general.asp | accessdate = 2007-01-20 |archiveurl = https://web.archive.org/web/20070210015835/http://wyoming.gov/general/general.asp |archivedate = February 10, 2007|deadurl=yes}}</ref>
The distinctive appearance of ''Triceratops'' has led to them being frequently depicted in films, computer games and documentaries, such as the 1993 film ''[[Jurassic Park (film)|Jurassic Park]]'' and the 1999 [[BBC]] television documentary ''[[Walking with Dinosaurs]]''. A recurring theme, especially in children's dinosaur books, is a climactic showdown or battle between ''Triceratops'' and ''[[Tyrannosaurus]]''. In 1942, [[Charles R. Knight]] painted a mural incorporating a confrontation between the two dinosaurs in the [[Field Museum of Natural History]] for the [[National Geographic Society]], establishing them as enemies in popular thought.<ref name="Bakker1986" /> Paleontologist [[Bob Bakker]] said of the imagined rivalry between ''Tyrannosaurus'' and ''Triceratops'', "No matchup between predator and prey has ever been more dramatic. It's somehow fitting that those two massive antagonists lived out their co-evolutionary belligerence through the [[K-T Boundary|very last days]] of the [[Maastrichtian|very last epoch]] of the Age of Dinosaurs."<ref name="Bakker1986">Bakker, R.T. 1986. ''The Dinosaur Heresies''. New York: Kensington Publishing, p. 240. On that page, Bakker has his own ''T. rex''/''Triceratops'' fight.</ref>

==See also==
{{Portal|Dinosaurs}}
* [[Timeline of ceratopsian research]]
{{clear}}
{{clear}}

==References==
{{reflist|30em}}

==External links==
{{Spoken Wikipedia|En-Triceratops-article.ogg|2007-03-19}}
{{Commons|Triceratops}}
{{Wikibooks|Wikijunior Dinosaurs/Triceratops}}
{{Wikisource|Notice of Gigantic Horned Dinosauria from the Cretaceous}}
{{Wikispecies}}
* [http://dinosaurpictures.org/Triceratops-pictures ''Triceratops''] at The Dinosaur Picture Database
* [http://www.livescience.com/24011-triceratops-facts.html LiveScience: Facts about ''Triceratops''] at LiveScience.com
* [https://www.youtube.com/watch?v=jjwhEx4LwlE Clash of the Dinosaurs: The Defenders - The Triceratops Threat], [[Discovery Channel]] (video)
* [http://dml.cmnh.org/2002Jul/msg00898.html Dinosaur Mailing List post on ''Triceratops'' stance]
* [http://www.mnh.si.edu/exhibits/triceratops/index.html Smithsonian Exhibit]
* [https://archive.org/details/en.wikipedia.org_wiki_Triceratops_2007_March_19 ''Triceratops'' at the Internet Archive]
* [http://www.nhm.ac.uk/nature-online/life/dinosaurs-other-extinct-creatures/dino-directory/triceratops.html ''Triceratops'' in the Dino Directory]
* [http://www.cbv.ns.ca/marigold/history/dinosaurs/datafiles/triceratops.html ''Triceratops''] (short summary and good color illustration)
* [http://www.enchantedlearning.com/subjects/dinosaurs/dinos/Triceratops.shtml ''Triceratops'' For Kids] (a fact sheet about the ''Triceratops'' with activities for kids)
* ''[http://www.bbc.co.uk/nature/life/Triceratops Triceratops]'', [[BBC]] Dinosaurs
* {fr} [http://www.dinosauria.ca/Dinos/triceratops.php Triceratops] - [http://www.dinosauria.ca/liste.php Liste] de [http://www.dinosauria.ca/ Dinosauria] et [http://www.dinosauria.ca/extinction.php Extinction]

{{featured article}}

{{Marginocephalia|T.}}

{{Authority control}}

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[[Category:Late Cretaceous dinosaurs of North America]]
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[[Category:Fossil taxa described in 1889]]
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[[Category:Maastrichtian genus extinctions]]

Pachycephalosauria

2017-09-13T13:23:12Z

Wikkler:

{{Automatic taxobox
| name = Pachycephalosaurs
| fossil_range = [[Late Cretaceous]], {{fossilrange|90|66|earliest=156}}
| image = Pachycephalosaurus skull.JPG
| image_caption = Cast of a ''[[Pachycephalosaurus wyomingensis]]'' skull, [[Oxford University Museum of Natural History]]
| taxon = Pachycephalosauridae
| parent_authority = [[Teresa Maryańska|Maryańska]] & [[Halszka Osmólska|Osmólska]], 1974
| authority = [[Charles Hazelius Sternberg|Sternberg]], 1945 ([[conserved name]])
| type_species = {{extinct}}''[[Pachycephalosaurus wyomingensis]]''
| type_species_authority = [[Charles Whitney Gilmore|Gilmore]], 1931 (conserved name)
| subdivision_ranks = [[Genus|Genera]]
| subdivision =
[[Pachycephalosauria#Taxonomy|See text]]
| synonyms =
'''Homalocephalidae''' Dong, 1978
}}

'''Pachycephalosauria''' ({{IPAc-en|ˌ|p|æ|k|i|ˌ|s|ɛ|f|əl|ə|ˈ|s|ɔːr|i|ə|,_|-|ˌ|k|ɛ|f|-}};{{refn|{{OxfordDictionaries.com|accessdate=2016-01-21|Pachycephalosauria}}}} from [[Ancient Greek|Greek]] παχυκεφαλόσαυρος for 'thick headed lizards') is a [[clade]] of [[ornithischian]] [[dinosaur]]s. Along with [[Ceratopsia]], it makes up the clade [[Marginocephalia]]. Genera include ''[[Pachycephalosaurus]]'', ''[[Stegoceras]]'', and ''[[Prenocephale]]''. With the exception of two species, most pachycephalosaurs lived during the Late [[Cretaceous]] Period, dating between about 85.8 and 65.5 million years ago.<ref name=":0">{{Cite book|url=http://www.geol.umd.edu/~tholtz/dinoappendix/HoltzappendixWinter2010.pdf|title=Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages|last=Holtz|first=Thomas R. Jr.|publisher=|year=2011|isbn=|location=|pages=}}</ref> They are exclusive to the [[Northern Hemisphere]], with the majority being found in what is now [[North America]] and [[Asia]]. They were all [[bipedal]], [[herbivorous]]/[[omnivorous]] animals with thick skulls. Skulls can be domed, flat, or wedge-shaped depending on the species, and are all heavily ossified. The domes were often surrounded by nodes and/or spikes. Partial skeletons have been found of several pachycephalosaur species, but to date no complete skeletons have been discovered. Often isolated skull fragments are the only bones that are found.<ref name=":1">{{Cite book|url=https://www.worldcat.org/oclc/493366196|title=The Dinosauria|last=...).,|first=Weishampel, David B. (1952-|last2=Peter,|first2=Dodson,|last3=...).,|first3=Osmólska, Halszka, (1930-|date=2007|publisher=University of California Press|isbn=0520242092|oclc=493366196}}</ref>

Candidates for the earliest known pachycephalosaur include [[Ferganocephale|''Ferganocephale adenticulatum'']] from [[Middle Jurassic]] [[Period (geology)|Period]] strata of [[Kyrgyzstan]] and [[Stenopelix|''Stenopelix valdensis'']] from [[Early Cretaceous]] strata of [[Germany]], although R.M. Sullivan has doubted that either of these species are pachycephalosaurs.[[Pachycephalosauria#cite note-sullivan2006-2|[2]]]

==Paleobiology==

=== Anatomy ===
Pachycephalosaurs were [[Bipedalism|bipedal]] [[ornithischia]]ns characterized by their thickened skulls. They had a bulky torso with an expanded gut cavity and broad hips, short forelimbs, long legs, a short, thick neck, and a heavy tail. Large orbits and a large [[optic nerve]] point to pachycephalosaurs having good vision, and uncharacteristically large [[Olfactory bulb|olfactory lobes]] indicate that they had a good sense of smell relative to other dinosaurs.<ref name=":1">{{Cite book|url=https://www.worldcat.org/oclc/493366196|title=The Dinosauria|last=...).,|first=Weishampel, David B. (1952-|last2=Peter,|first2=Dodson,|last3=...).,|first3=Osmólska, Halszka, (1930-|date=2007|publisher=University of California Press|isbn=0520242092|oclc=493366196}}</ref> They were fairly small dinosaurs, with most falling in the range of 2–3 meters (6.6-9.8 feet) in length and the largest, ''[[Pachycephalosaurus|Pachycephalosaurus wyomingensis]],'' estimated to measure 4.5 meters (14.8 feet) long and weigh 450 kilograms (990 pounds).<ref name=":0">{{Cite book|url=http://www.geol.umd.edu/~tholtz/dinoappendix/HoltzappendixWinter2010.pdf|title=Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages|last=Holtz|first=Thomas R. Jr.|publisher=|year=2011|isbn=|location=|pages=}}</ref><ref>{{Cite book|url=https://www.worldcat.org/oclc/930852339|title=The Princeton field guide to dinosaurs|last=S.|first=Paul, Gregory|date=2010|publisher=Princeton University Press|isbn=9780691137209|oclc=930852339}}</ref> The characteristic skull of pachycephalosaurs is a result of the fusion and thickening of the [[Frontal bone#Dinosaurs|frontals]] and [[Parietal bone#In dinosaurs|parietals]], accompanied by the closing of the [[Skull#Temporal fenestrae|supratemporal fenestra]]. In some species this takes the form of a raised dome; in others, the skull is flat or wedge-shaped. While the flat-headed pachycephalosaurs are traditionally regarded as distinct species or even families, they may represent juveniles of dome-headed adults.[[Pachycephalosauria#cite note-sullivan2006-2|[2]]][[Pachycephalosauria#cite note-longrichetal2010-3|[3]]] All display highly ornamented [[Jugal bone|jugals]], [[Squamosal bone|squamosals]], and [[Postorbital bone|postorbitals]] in the form of blunt horns and nodes. Many species are only known from skull fragments, and a complete pachycephalosaur skeleton is yet to be found.<ref name=":1" />

===Head-butting behavior===
[[File:Pachycephalosaurus head butting.png|thumb|left|Restoration of head butting ''Pachycephalosaurus'']]
[[File:Pachycephalosauridae head butting.png|thumb|Hypothetical examples of pachycephalosaur combat behavior, varying per species: ''[[Pachycephalosaurus]]'' (A), ''[[Prenocephale]]'' (B), ''[[Stygimoloch]]'' (C).]]The adaptive significance of the skull dome has been heavily debated. The popular hypothesis among the general public that the skull was used in head-butting, as sort of a dinosaurian battering ram, was first proposed by {{harvnb|Colbert|1955}}. This view was popularized in the 1956 science fiction story "[[A Gun for Dinosaur]]" by [[L. Sprague de Camp]]. Many paleontologists have since argued for the head-butting hypothesis, including {{harvnb|Galton|1970}} and {{harvnb|Sues|1978}}. In this hypothesis, pachycephalosaurs rammed each other head-on, as do modern-day [[mountain goat]]s and [[musk oxen]].
[[File:Pachycephalosaurus Clean.png|thumb|left|''Pachycephalosaurus'' based on the "Sandy" specimen in the [[Rocky Mountain Dinosaur Resource Center]], Woodland Park, Colorado]]
Anatomical evidence for combative behavior includes vertebral articulations providing spinal rigidity, and the shape of the back indicating strong neck musculature.<ref name="snively">{{harvnb|Snively|Cox|2008}}</ref> It has been suggested that pachycephalosaurs could make their head, neck, and body horizontally straight, in order to transmit stress during ramming. However, in no known dinosaur can the head, neck, and body be oriented in such a position. Instead, the cervical and anterior dorsal vertebrae of pachycephalosaurs show that the neck was carried in an "S"- or "U"-shaped curve.<ref name="KC97">{{harvnb|Carpenter|1997}}</ref>

Also, the rounded shape of the skull would lessen the contacted surface area during head-butting, resulting in glancing blows. Other possibilities include flank-butting, defense against predators, or both. The relatively wide build of pachycephalosaurs (which would protect vital internal organs from harm during flank-butting) and the squamosal horns of the ''[[Stygimoloch]]'' (which would have been used to great effect during flank-butting) add credence to the flank-butting hypothesis.

A histological study conducted by {{harvnb|Goodwin|Horner|2004}} argued against the battering ram hypothesis. They argued that the dome was "an ephemeral ontogenetic stage", the spongy bone structure could not sustain the blows of combat, and the radial pattern was simply an effect of rapid growth.<ref name="goodwin">{{harvnb|Goodwin|Horner|2004}}</ref> Later biomechanical analyses by {{harvnb|Snively|Cox|2008}} and {{harvnb|Snively|Theodor|2011}} concluded, however, that the domes could withstand combat stresses.<ref name="snively"/> {{harvnb|Lehman|2010}} argued that the growth patterns discussed by Goodwin and Horner are not inconsistent with head-butting behavior.<ref>{{harvnb|Lehman|2010}}</ref>

{{harvnb|Goodwin|Horner|2004}} instead argued that the dome functioned for species recognition. There is evidence that the dome had some form of external covering, and it is reasonable to consider the dome may have been brightly covered, or subject to change color seasonally.<ref name="goodwin"/> Due to the nature of the fossil record, however, it cannot be observed whether or not color played a role in dome function.

{{harvnb|Longrich|Sankey|Tanke|2010}} argued that species recognition is an unlikely evolutionary cause for the dome because dome forms are not notably different between species. Because of this general similarity, several genera of Pachycephalosauridae have sometimes been incorrectly lumped together. This is unlike the case in [[ceratopsia]]ns and [[hadrosaur]]ids, which had much more distinct cranial ornamentation. Longrich ''et al.''. argued that instead the dome had a mechanical function, such as combat, one which was important enough to justify the resource investment.<ref name="longrichetal2010"/>

New findings published in the Journal ''Plos One'' on July 16, 2013 further evidence head-butting in species of pachycephalosaurs. Of the 100 domes studied, 20 percent showed signs of healed injuries, all consistent with combative behavior. The pathologies included pitting where the bone had become infected from wounds that originated in the skin. These findings further imply that pachycephalosaurids may have been using their heads for both display and defense as many modern animals do.<ref>{{cite web|url=http://www.nbcnews.com/science/boneheaded-dinos-butted-heads-combat-6C10680037|title=Boneheaded dinos butted heads in combat|work=NBC News}}</ref>

===Diet===
The small size of most pachycephalosaur species and lack of skeletal adaptation indicates that they were not climbers and primarily ate food close to the ground. Mallon et al. (2013) examined herbivore coexistence on the island continent of [[Laramidia]], during the Late Cretaceous and concluded that pachycephalosaurids were generally restricted to feeding on vegetation at, or below, the height of 1 meter.<ref>{{cite journal|last=Mallon|first=Jordan C|author2=David C Evans |author3=Michael J Ryan |author4=Jason S Anderson |title=Feeding height stratification among the herbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta, Canada|journal=BMC Ecology|year=2013|volume=13|url=https://link.springer.com/article/10.1186/1472-6785-13-14 |doi=10.1186/1472-6785-13-14 |pages=14 |pmid=23557203 |pmc=3637170}}</ref> They exhibit [[heterodont]]y, having different tooth morphology between the [[premaxilla]]ry teeth and [[maxilla]]ry teeth. Front teeth are small and peg-like with an ovular cross section and were most likely used for grabbing food. In some species, the last premaxillary tooth was enlarged and canine-like. Back teeth are small and triangular with [[Denticle (tooth feature)|denticles]] on the front and back of the crown, used for mouth processing. In species in which the [[Mandible|dentary]] has been found, mandibular teeth are similar in size and shape to those in the upper jaw. Wear patterns on the teeth vary by species, indicating a range of food preferences which could include seeds, stems, leaves, fruits, and possibly insects. A very wide rib cage and large gut cavity extending all the way to the base of the tail suggests the use of fermentation to digest food.<ref name=":1" />

===Paleopathology===
Peterson et al. (2013) studied cranial pathologies among the Pachycephalosauridae and found that 22% of all domes examined had lesions that are consistent with [[osteomyelitis]], an infection of the bone resulting from penetrating trauma, or trauma to the tissue overlying the skull leading to an infection of the bone tissue. This high rate of pathology lends more support to the hypothesis that pachycephalosaurid domes were employed in intra-specific combat.<ref name=Peterson13>{{cite journal | last1 = Peterson | first1 = JE | last2 = Dischler | first2 = C | last3 = Longrich | first3 = NR | year = 2013 | title = Distributions of Cranial Pathologies Provide Evidence for Head-Butting in Dome-Headed Dinosaurs (Pachycephalosauridae) | journal = PLoS ONE | volume = 8 | issue = 7| page = e68620 | doi = 10.1371/journal.pone.0068620 | pmid=23874691 | pmc=3712952}}</ref> The frequency of trauma was comparable across the different genera in this [[Family (biology)|family]], despite the fact that these genera vary with respect to the size and architecture of their domes, and fact that they existed during varying geologic periods.<ref name=Peterson13/> These findings were in stark contrast with the results from analysis of the relatively flat-headed pachycephalosaurids, where there was an absence of pathology. This would support the hypothesis that these individuals represent either females or juveniles,<ref>{{cite journal | last1 = Longrich | first1 = NR | last2 = Sankey | first2 = J | last3 = Tanke | first3 = D | year = 2010 | title = Texacephale langstoni, a new genus of pachycephalosaurid (Dinosauria: Ornithischia) from the upper Campanian Aguja Formation, southern Texas, USA | url = | journal = Cretaceous Research | volume = 31 | issue = 2| pages = 274–284 | doi=10.1016/j.cretres.2009.12.002}}</ref> where intra-specific combat behavior is not expected.

Histological examination reveals that pachycephalosaurid domes are composed of a unique form of fibrolamellar bone<ref>Reid REH (1997) Histology of bones and teeth. In: Currie, PJ and Padian, K, editors. Encyclopedia of Dinosaurs. Academic Press, San Diego, CA. 329–339.</ref> which contains [[fibroblasts]] that play a critical role in wound healing, and are capable of rapidly depositing bone during remodeling.<ref>{{cite journal | last1 = Horner | first1 = JR | last2 = Goodwin | first2 = MB | year = 2009 | title = Extreme Cranial Ontogeny in the Upper Cretaceous Dinosaur Pachycephalosaurus | url = | journal = PLoS ONE | volume = 4 | issue = 10| page = e7626 | doi = 10.1371/journal.pone.0007626 | pmid=19859556 | pmc=2762616}}</ref> Peterson et al. (2013) concluded that, taken together, the frequency of lesion distribution and the bone structure of frontoparietal domes, lends strong support to the hypothesis that pachycephalosaurids used their unique cranial structures for [[agonistic behavior]].<ref name=Peterson13/>

== Biogeography ==

=== Distribution ===
Pachycephalosaurs lived exclusively in [[Laurasia]]. One species, ''[[Stenopelix|Stenopelix valdensis]]'', has been found in [[Europe]], with the rest being found in western [[North America]] and central [[Asia]]. Pachycephalosaurs originated in Asia and had two major dispersal events, resulting in the two separate waves of pachycephalosaur evolution observed in Asia. The first, occurring before the late [[Santonian]] or early [[Campanian]], involved a migration from Asia to North America, most likely by way of the [[Beringia|Bering Land Bridge]]. This migration was by a common ancestor of ''[[Ornatotholus]], [[Stygimoloch]], [[Stegoceras]], [[Tylocephale]], [[Prenocephale]],'' and ''[[Pachycephalosaurus]].'' The second event occurred before the middle [[Campanian]], and involved a migration back into Asia from North America by a common ancestor of ''[[Prenocephale]]'' and ''[[Tylocephale]].'' A smaller dispersal event occurring sometime before the [[Berriasian]], prior the first major event, resulted in the European species. Two species originally reported to be pachycephalosaurs discovered outside this range, ''[[Yaverlandia|Yaverlandia bitholus]]'' of [[England]] and ''[[Majungasaurus|Majungatholus atopus]]'' of [[Madagascar]], have recently been shown to actually be [[Theropoda|therapods]].<ref name=":1" /><ref>{{Cite journal|last=NAISH|first=D.|last2=MARTILL|first2=D. M.|title=Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: Ornithischia|url=http://mr.crossref.org/iPage?doi=10.1144%2F0016-76492007-154|journal=Journal of the Geological Society|volume=165|issue=3|pages=613–623|doi=10.1144/0016-76492007-154}}</ref>

=== Environment ===
The Asian and North American species of pachycephalosaurs lived in markedly different environments. Asian specimens are normally more of intact, indicating they were not transported far from their place of death before fossilization. They likely lived in a large desert region in central Asia with a hot and [[Desert climate|arid climate]]. North American specimens are typically found in rocks that were formed by erosion from the [[Rocky Mountains]]. Specimens are far less intact; usually only skull caps are recovered, and those found regularly exhibit surface exfoliation and other signs that they were transported long distances by water before fossilization. It is assumed that they lived in the mountains in a [[temperate climate]] and were carried by erosion after death to their final resting place.<ref>{{Cite book|url=https://www.worldcat.org/oclc/476234422|title=Dinosaurs : a concise natural history|last=1952-|first=Weishampel, David B.,|date=2009|publisher=Cambridge University Press|isbn=9780511479410|oclc=476234422}}</ref>

==Classification==
[[File:Pachycephalosaurus ontogeny.png|thumb|left|Diagram showing ''Dracorex'' and ''Stygimoloch'' as growth stages of ''Pachycephalosaurus'']]
Most pachycephalosaurid remains are not complete, usually consisting of portions of the frontoparietal bone that forms the distinctive dome. This can make [[taxonomic]] identification a difficult task, as the classification of genera and species within ''Pachycephalosauria'' relies almost entirely on cranial characteristics. Consequently, improper species have historically been appointed to the clade. For instance, ''[[Majungatholus]]'', once thought to be a pachycephalosaur, is now recognized as a specimen of the [[Abelisauridae|abelisaurid]] [[Theropoda|theropod]] ''[[Majungasaurus]]''. And ''[[Yaverlandia]]'', another dinosaur initially described as a pachycephalosaurid, has also recently been reclassified as a [[coelurosaur]] (Naish in {{harvnb|Sullivan|2006}}). Further complicating matters are the diverse interpretations of [[ontogenetic]] and [[Sexual selection|sexual features]] in pachycephalosaurs.

A 2009 paper proposed that ''Dracorex'' and ''Stygimoloch'' were just early growth stages of ''Pachycephalosaurus'', rather than distinct genera.<ref>{{harvnb|Horner|Goodwin|2009}}</ref>

===Taxonomy===
[[File:Royal_Tyrrell_Museum_Stegoceras.jpg|thumb|right|Two mounted ''[[Stegoceras]]'' skeletons.]]
The Pachycephalosauria was first named as a [[Order (biology)|suborder]] of the order [[Ornithischia]] by {{harvnb|Maryańska|Osmólska|1974}}. They included within it only one [[Family (biology)|family]], the Pachycephalosauridae.<ref name="maryanska&osmolska1974">{{harvnb|Maryańska|Osmólska|1974}}</ref> Later researchers, such as Michael Benton, have ranked it as an [[Order (biology)|infraorder]] of a suborder [[Cerapoda]], which unites the [[ceratopsia]]ns and [[ornithopod]]s.<ref name="benton2004">{{harvnb|Benton|2004| pp=472}}</ref> In 2006, Robert Sullivan published a re-evaluation of pachycephalosaur taxonomy. Sullivan considered attempts by Maryańska and Osmólska to restrict the definition of Pachycephalosauria redundant with their Pachycephalosauridae, since they were diagnosed by the same anatomical characters. Sullivan also rejected attempts by {{harvnb|Sereno|1986}}, in his [[phylogenetic]] studies,<ref>{{harvnb|Sereno|1986}}</ref> to re-define Pachycephalosauridae to include only "dome-skulled" species (including ''Stegoceras'' and ''Pachycephalosaurus''), while leaving more "basal" species outside that family in Pachycephalosauria. Therefore, Sullivan's use of Pachycephalosauridae is equivalent to Sereno and Benton's use of Pachycephalosauria.

Sullivan diagnosed the Pachycephalosauridae-based only on characters of the skull, with the defining character being a dome-shaped ''frontoparietal'' skull bone. According to Sullivan, the absence of this feature in some species assumed to be primitive led to the split in classification between domed and non-domed pachycephalosaurs; however, discovery of more advanced and possibly juvenile pachycephalosaurs with flat skulls (such as ''Dracorex hogwartsia'') show this distinction to be incorrect. Sullivan also pointed out that the original diagnosis of Pachycephalosauridae centered around "flat to dome-like" skulls, so the flat-headed forms should be included in the family.<ref name="sullivan2006">{{harvnb|Sullivan|2006}}</ref>

The following [[Taxonomy (biology)|taxonomy]] follows Sullivan's 2006 classification unless otherwise noted.

* '''Family Pachycephalosauridae'''
** ''[[Acrotholus]]''
** ''[[Alaskacephale]]''
** ''[[Amtocephale]]''<ref name=Amtocephale>{{Harvnb|Watabe|Tsogtbaatar|Sullivan|2011}}</ref>
** ''[[Colepiocephale]]''
** ''[[Foraminacephale]]''
** ''[[Goyocephale]]''
** ''[[Gravitholus]]''<ref name=longrichetal2010 />
** ''[[Hanssuesia]]''
** ''[[Homalocephale]]'' - possible juvenile form of ''Prenocephale''<ref name=longrichetal2010 />
** ''[[Prenocephale]]''
** ''[[Sphaerotholus]]''<ref name=longrichetal2010 />
** ''[[Stegoceras]]'' (incl. ''Ornatotholus'')
** ''[[Texacephale]]''<ref name="longrichetal2010">{{harvnb|Longrich|Sankey|Tanke|2010}}</ref>
** ''[[Tylocephale]]''
** ''[[Wannanosaurus]]''<ref name=longrichetal2010 />
** '''Tribe Pachycephalosaurini'''
*** ''[[Dracorex]]'' - possible juvenile form of ''Pachycephalosaurus''<ref name="juv" />
*** ''[[Pachycephalosaurus]]''
*** ''[[Stygimoloch]]'' - possible juvenile form of ''Pachycephalosaurus''<ref name="juv" >{{harvnb|Stokstad|2007}}</ref>
* ''Nomina dubia'' (doubtful names)
** ''[[Ferganocephale]]''
** "Stegoceras" ''bexelli''

Note that Butler ''et al.'', 2011 reassigned ''[[Stenopelix]]'' and ''[[Micropachycephalosaurus]]'' to the [[Ceratopsia]].

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from: 1880 till: 1890 color:1800syears text:[[1880s in paleontology|80s]]
from: 1890 till: 1900 color:1800syears text:[[1890s in paleontology|90s]]
from: 1900 till: 1910 color:1900syears text:[[1900s in paleontology|00s]]
from: 1910 till: 1920 color:1900syears text:[[1910s in paleontology|10s]]
from: 1920 till: 1930 color:1900syears text:[[1920s in paleontology|20s]]
from: 1930 till: 1940 color:1900syears text:[[1930s in paleontology|30s]]
from: 1940 till: 1950 color:1900syears text:[[1940s in paleontology|40s]]
from: 1950 till: 1960 color:1900syears text:[[1950s in paleontology|50s]]
from: 1960 till: 1970 color:1900syears text:[[1960s in paleontology|60s]]
from: 1970 till: 1980 color:1900syears text:[[1970s in paleontology|70s]]
from: 1980 till: 1990 color:1900syears text:[[1980s in paleontology|80s]]
from: 1990 till: 2000 color:1900syears text:[[1990s in paleontology|90s]]
from: 2000 till: 2010 color:2000syears text:[[2000s in paleontology|00s]]
from: 2010 till: 2020 color:2000syears text:[[2010s in paleontology|10s]]
from: 2020 till: 2030 color:2000syears text:[[2020s in paleontology|20s]]
from: 2030 till: 2040 color:2000syears text:[[2030s in paleontology|30s]]
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from: 2070 till: 2080 color:2000syears text:[[2070s in paleontology|70s]]
from: 2080 till: 2090 color:2000syears text:[[2080s in paleontology|80s]]
from: 2090 till: 2100 color:2000syears text:[[2090s in paleontology|90s]]

bar:eratop
from: 1824 till: 1900 color:1800s text:[[19th century in paleontology|19th]]
from: 1900 till: 2000 color:1900s text:[[20th century in paleontology|20th]]
from: 2000 till: 2100 color:2000s text:[[21st century in paleontology|21st]]

PlotData=
align:left fontsize:M mark:(line,white) width:5 anchor:till align:left

color:1900s bar:NAM8 at:2013 mark:(line,black) text:[[Acrotholus]]
color:1900s bar:NAM4 at:2006 mark:(line,black) text:[[Alaskacephale]]
color:1900s bar:NAM7 at:2011 mark:(line,black) text:[[Amtocephale]]
color:1900s bar:NAM2 at:2003 mark:(line,black) text:[[Colepiocephale]]
color:1900s bar:NAM6 at:1982 mark:(line,black) text:[[Goyocephale]]
color:1800s bar:NAM5 at:1979 mark:(line,black) text:[[Gravitholus]]
color:1900s bar:NAM3 at:2003 mark:(line,black) text:[[Hanssuesia]]
color:1900s bar:NAM1 at:1974 mark:(line,black) text:[[Homalocephale]]
color:1800s bar:NAM2 at:1974 mark:(line,black) text:[[Prenocephale]]
color:1800s bar:NAM1 at:2002 mark:(line,black) text:[[Sphaerotholus]]
color:1900s bar:NAM1 at:1902 mark:(line,black) text:[[Stegoceras]]
color:1800s bar:NAM6 at:2010 mark:(line,black) text:[[Texacephale]]
color:1900s bar:NAM3 at:1974 mark:(line,black) text:[[Tylocephale]]
color:1900s bar:NAM4 at:1977 mark:(line,black) text:[[Wannanosaurus]]
color:1800s bar:NAM5 at:2006 mark:(line,black) text:[[Dracorex]]
color:1900s bar:NAM1 at:1943 mark:(line,black) text:[[Pachycephalosaurus]]
color:1900s bar:NAM7 at:1983 mark:(line,black) text:[[Stygimoloch]]

PlotData=
align:center textcolor:black fontsize:M mark:(line,black) width:25

bar:period
from: 1824 till: 1830 color:1800syears text:[[1820s in paleontology|20s]]
from: 1830 till: 1840 color:1800syears text:[[1830s in paleontology|30s]]
from: 1840 till: 1850 color:1800syears text:[[1840s in paleontology|40s]]
from: 1850 till: 1860 color:1800syears text:[[1850s in paleontology|50s]]
from: 1860 till: 1870 color:1800syears text:[[1860s in paleontology|60s]]
from: 1870 till: 1880 color:1800syears text:[[1870s in paleontology|70s]]
from: 1880 till: 1890 color:1800syears text:[[1880s in paleontology|80s]]
from: 1890 till: 1900 color:1800syears text:[[1890s in paleontology|90s]]
from: 1900 till: 1910 color:1900syears text:[[1900s in paleontology|00s]]
from: 1910 till: 1920 color:1900syears text:[[1910s in paleontology|10s]]
from: 1920 till: 1930 color:1900syears text:[[1920s in paleontology|20s]]
from: 1930 till: 1940 color:1900syears text:[[1930s in paleontology|30s]]
from: 1940 till: 1950 color:1900syears text:[[1940s in paleontology|40s]]
from: 1950 till: 1960 color:1900syears text:[[1950s in paleontology|50s]]
from: 1960 till: 1970 color:1900syears text:[[1960s in paleontology|60s]]
from: 1970 till: 1980 color:1900syears text:[[1970s in paleontology|70s]]
from: 1980 till: 1990 color:1900syears text:[[1980s in paleontology|80s]]
from: 1990 till: 2000 color:1900syears text:[[1990s in paleontology|90s]]
from: 2000 till: 2010 color:2000syears text:[[2000s in paleontology|00s]]
from: 2010 till: 2020 color:2000syears text:[[2010s in paleontology|10s]]
from: 2020 till: 2030 color:2000syears text:[[2020s in paleontology|20s]]
from: 2030 till: 2040 color:2000syears text:[[2030s in paleontology|30s]]
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from: 2060 till: 2070 color:2000syears text:[[2060s in paleontology|60s]]
from: 2070 till: 2080 color:2000syears text:[[2070s in paleontology|70s]]
from: 2080 till: 2090 color:2000syears text:[[2080s in paleontology|80s]]
from: 2090 till: 2100 color:2000syears text:[[2090s in paleontology|90s]]

bar:era
from: 1824 till: 1900 color:1800s text:[[19th century in paleontology|19th]]
from: 1900 till: 2000 color:1900s text:[[20th century in paleontology|20th]]
from: 2000 till: 2100 color:2000s text:[[21st century in paleontology|21st]]
</timeline>

===Phylogeny===
The [[cladogram]] presented here follows an analysis by {{Harvnb|Williamson|Carr|2002}}.<ref name="williamson&carr2002" >{{harvnb|Williamson|Carr|2002}}</ref>
{{clade| style=font-size:100%;line-height:80%
|label1='''Pachycephalosauria'''
|1={{clade
|1=''[[Stenopelix]]''
|2=''[[Wannanosaurus]]''
|3=''[[Yaverlandia]]'' (now thought to be a [[theropod]] instead)
|label4='''Goyacephala'''
|4={{clade
|1=''[[Goyocephale]]''
|label2=unnamed
|2={{clade
|1=''[[Homalocephale]]''
|label2='''Pachycephalosauridae'''
|2={{clade
|1=''[[Stegoceras]]''
|label2='''Pachycephalosaurinae'''
|2={{clade
|label1=unnamed
|1={{clade
|1=''[[Tylocephale]]''
|2=''[[Prenocephale]]''
}}
|label2= unnamed
|2={{clade
|1=''[[Sphaerotholus]]''
|label2= unnamed
|2={{clade
|1=''[[Stygimoloch]]''
|2=''[[Pachycephalosaurus]]''
}}
}}
}}
}}
}}
}}
}}
}}

[[Cladogram]] after Longrich, Sankey and Tanke (2010).<ref name="Longrich''et al.''">{{cite journal|author= Longrich, N.R., Sankey, J., and Tanke, D. |year= 2010 |title= ''Texacephale langstoni'', a new genus of pachycephalosaurid (Dinosauria: Ornithischia) from the upper Campanian Aguja Formation, southern Texas, USA |journal= Cretaceous Research |volume=31 |issue=2 |pages=274–284 |doi= 10.1016/j.cretres.2009.12.002}}</ref>

{{clade| style=font-size:90%;line-height:85%
|label1= [[Pachycephalosauridae]] 
|1={{clade
|1={{clade
|1=''[[Stegoceras]]''
|2={{clade
|1=''[[Colepiocephale]]''
|2=''[[Gravitholus]]'' }} }}
|2=''[[Texacephale]]''
|3=''[[Hanssuesia]]''
|4={{clade
|1=''[[Foraminacephale|Foraminacephale brevis]]''
|2={{clade
|1=''[[Sphaerotholus|S. goodwini]]''
|2={{clade
|1=''[[Sphaerotholus|S. edmontonensis]]''
|2=''[[Sphaerotholus|S. buchholtzae]]'' }} }}
|3={{clade
|1={{clade
|1=''[[Alaskacephale]]''
|2=''[[Pachycephalosaurus]]''
|3={{clade
|1=''[[Stygimoloch]]''
|2=''[[Dracorex]]'' }} }}
|2={{clade
|1=''[[Tylocephale]]''
|2=''[[Prenocephale]]''
|3={{clade
|1=''[[Homalocephale]]''
|2=''[[Goyocephale]]''
|3=''[[Wannanosaurus]]'' }} }} }} }} }} }}

Below is a [[cladogram]] modified from Evans ''et al.'', 2013.<ref name=Acrotholus>{{Cite journal | last1 = Evans | first1 = D. C. | last2 = Schott | first2 = R. K. | last3 = Larson | first3 = D. W. | last4 = Brown | first4 = C. M. | last5 = Ryan | first5 = M. J. | title = The oldest North American pachycephalosaurid and the hidden diversity of small-bodied ornithischian dinosaurs | doi = 10.1038/ncomms2749 | journal = Nature Communications | volume = 4 | pages = 1828 | year = 2013 | pmid = 23652016| pmc = | bibcode = 2013NatCo...4E1828E }}</ref>

{{clade| style=font-size:85%;line-height:85%
|label1= Pachycephalosauria 
|1={{clade
|1=''[[Wannanosaurus yansiensis]]''
|label2= [[Pachycephalosauridae]] 
|2={{clade
|1={{clade
|1=''[[Colepiocephale lambei]]''
|2=''[[Hanssuesia sternbergi]]''
|3=''[[Stegoceras novomexicanum]]''
|4=''[[Stegoceras validum]]''}}
|2={{clade
|1=''[[Goyocephale lattimorei]]''
|2={{clade
|1=''[[Homalocephale calathocercos]]''
|2={{clade
|1=''[[Tylocephale gilmorei]]''
|2={{clade
|1=''[[Foraminacephale brevis]]''
|2={{clade
|1=''[[Amtocephale gobiensis]]''
|2={{clade
|1={{clade
|1=''[[Acrotholus audeti]]''
|2=''[[Prenocephale prenes]]''}}
|2={{clade
|1={{clade
|1=''[[Alaskacephale gangloffi]]''
|2=''[[Pachycephalosaurus wyomingensis]]''}}
|2={{clade
|1=''[[Sphaerotholus buchholtzae]]''
|2=''[[Sphaerotholus goodwini]]'' }} }} }} }} }} }} }} }} }} }} }}

==See also==
{{Portal|Dinosaurs}}
* [[Timeline of pachycephalosaur research]]

==Notes==
{{Reflist|30em}}

==References==
{{Refbegin|30em}}
* {{Cite book | ref = harv
| last = Benton | first = Michael J | authorlink = Michael Benton
| title = Vertebrate Palaeontology | edition = 3rd
| publisher = Blackwell Publishing | year = 2004
| isbn = 978-0-632-05637-8
}}
* {{Cite journal | ref = harv
| last=Carpenter | first=Kenneth | authorlink=Kenneth Carpenter
| title=Agonistic behavior in pachycephalosaurs (Ornithischia:Dinosauria): a new look at head-butting behavior
| journal=Contributions to Geology |year=1997 |volume=32 |issue=1 |pages= 19–25
| url = http://www.le-monde-des-dinosaures.net/pachycephalosaures.pdf
}}
* {{Cite book | ref = harv
| last = Colbert | first = Edwin | authorlink = Edwin Harris Colbert
| title = Evolution of the Vertebrates
| publisher = John wiley | location = New York | year = 1955
}}
* {{Cite journal | ref = harv
| last = Galton | first = P | authorlink = Peter Galton
| title = Pachycephalosaurids — dinosaurian battering rams
| journal = Discovery | year = 1970
| publisher = Yale Peabody Museum | location = London
| volume = 6 | pages = 23–32
}}
* {{Cite journal | ref = harv
| last1 = Goodwin | first1 = MB
| last2 = Horner | first2 = JR
| title = Cranial histology of pachycephalosaurs (Ornithischia: Marginocephalia) reveals transitory structures inconsistent with head-butting behavior
| journal = Paleobiology |date=June 2004 | volume = 30 | issue = 2 | pages = 253–267
| doi = 10.1666/0094-8373(2004)030<0253:CHOPOM>2.0.CO;2
}}
* {{Cite journal | ref = harv
| last1 = Horner | first1 = JR
| last2 = Goodwin | first2 = MB
| title = Extreme Cranial Ontogeny in the Upper Cretaceous Dinosaur Pachycephalosaurus
| journal = PLoS ONE | year = 2009 | volume = 4 | issue = 10 | page = e7626
| doi = 10.1371/journal.pone.0007626
| url = http://www.plosone.org/article/info:doi/10.1371/journal.pone.0007626 | pmid=19859556 | pmc=2762616 | editor1-last = Sereno | editor1-first = Paul
}}
* {{Cite journal | ref = harv
| last = Lehman | first = TM
| title = Pachycephalosauridae from the San Carlos and Aguja Formations (Upper Cretaceous) of west Texas, and observations of the frontoparietal dome
| journal = Journal of Vertebrate Paleontology | year = 2010 | issue = 3 | pages = 786–798
| issn = 0272-4634
| doi = 10.1080/02724631003763532 | volume = 30
}}
* {{Cite journal | ref = harv
| last1 = Longrich | first1 = NR
| last2 = Sankey | first2 =J
| last3 = Tanke | first3 = D
| title = ''Texacephale langstoni'', a new genus of pachycephalosaurid (Dinosauria: Ornithischia) from the upper Campanian Aguja Formation, southern Texas, USA
| journal = Cretaceous Research | year = 2010
| doi = 10.1016/j.cretres.2009.12.002 | volume = 31 | issue = 2 | pages = 274–284
}}
* {{Cite journal | ref = harv
| last1 = Maryańska | first1 = T | author1-link = Teresa Maryańska
| last2 = Osmólska | first2 = H | author2-link = Halszka Osmólska
| title = Pachycephalosauria, a new suborder of ornithischian dinosaurs
| journal = Palaeontologica Polonica | year = 1974 | issue = 30 | pages = 45–102
}}
* {{Cite journal | ref = harv
| last = Sereno | first = PC | authorlink = Paul Sereno
| title = Phylogeny of the bird-hipped dinosaurs (Order Ornithischia)
| journal = National Geographic Research | year = 1986 | volume = 2 | pages = 234–256
}}
* {{Cite journal | ref = harv
| last1 = Snively | first1 = E
| last2 = Cox | first2 = A
| title = Structural mechanics of pachycephalosaur crania permitted head-butting behavior
| journal = Palaeontologia Electronica | issue = 11 | year = 2008
| url = http://palaeo-electronica.org/2008_1/140/index.html
}}
* {{Cite journal | ref = harv
| last1 = Snively | first1 = E
| last2 = Theodor | first2 = JM
| title = Common Functional Correlates of Head-Strike Behavior in the Pachycephalosaur Stegoceras validum (Ornithischia, Dinosauria) and Combative Artiodactyls
| journal = PLoS ONE | year = 2011 | volume = 6 | issue = 6 | page = e21422
| doi = 10.1371/journal.pone.0021422 | editor1-last = Carpenter | editor1-first = Kenneth | pmid=21738658 | pmc=3125168
}}
* {{Cite journal | ref = harv
| last = Stokstad | first = Erik
| title = Society of Vertebrate Paleontology Meeting: Did Horny Young Dinosaurs Cause Illusion of Separate Species?
| journal = Science | volume = 318 |date=November 2007 | page = 1236
| url = http://www.sciencemag.org/cgi/content/full/318/5854/1236 | doi=10.1126/science.318.5854.1236 | pmid=18033861 | issue=5854
}}
* {{Cite journal | ref = harv
| last = Sues | first = H-D
| title = Functional morphology of the dome in pachycephalosaurid dinosaurs
| journal = Neues Jahrbuch für Geologie und Paläontologie | year = 1978 | pages = 459–472
}}
* {{Cite journal | ref = harv
| last = Sullivan | first =RM
| title = A taxonomic review of the Pachycephalosauridae (Dinosauria: Ornithischia)
| journal = New Mexico Museum of Natural History and Science Bulletin | year = 2006 | issue = 35 | pages = 347–365
| url = http://www.robertmsullivanphd.com/uploads/130_Sullivan__2006__-Pachycephalosauridae.pdf
}}
* {{cite journal | ref = harv
| last1 = Watabe | first1 = Mahito
| last2 = Tsogtbaatar | first2 = Khishigjaw
| last3 = Sullivan | first3 = Robert M.
| title = A new pachycephalosaurid from the Baynshire Formation (Cenomanian-late Santonian), Gobi Desert, Mongolia
| url = http://www.robertmsullivanphd.com/uploads/174.Watabe_et_al__Mongolian_pachy_.pdf
| journal = Fossil Record 3. New Mexico Museum of Natural History and Science, Bulletin | year = 2011 | volume = 53 | pages = 489–497
}}
* {{Cite journal | ref = harv
| last1 = Williamson | first1 = TE
| last2 = Carr | first2 = TD
| title = A new genus of derived pachycephalosaurian from western North America
| journal = Journal of Vertebrate Paleontology | year = 2002 | volume = 22 | issue = 4 | pages = 779–801
| doi = 10.1671/0272-4634(2002)022[0779:ANGODP]2.0.CO;2 | issn = 0272-4634
}}
{{Refend}}

==External links==
* {{Youtube|xYbMXzBwpIo|TEDx talk by [[John R. Horner|Jack Horner]] on shape-shifting dinosaur skulls and dinosaur misclassification}}

{{Marginocephalia|P.}}

{{taxonbar}}

[[Category:Marginocephalians]]
[[Category:Pachycephalosaurs| ]]
[[Category:Turonian first appearances]]
[[Category:Maastrichtian extinctions]]

Troodontidae

2017-09-01T16:21:54Z

Wikkler: /* Relationships */ Corrected spelling of Avialae.

{{Automatic taxobox
| name = Troodontids
| fossil_range = [[Late Jurassic]]–[[Late Cretaceous]], {{Fossil range|150|66|earliest=160}}
| image_caption = Mounted skeletal cast of an unnamed Alaskan troodontid, [[Perot Museum]]
| image = Alaskan troodont.jpg
| authority = [[Charles W. Gilmore|Gilmore]], 1924
| type_species = ''[[Troodon formosus]]''
| type_species_authority = [[Joseph Leidy|Leidy]], 1856
| subdivision_ranks = Subfamilies and Genera
| subdivision =
*{{extinct}}''[[Albertavenator]]''
*{{extinct}}''[[Anchiornis]]''?
*{{extinct}}''[[Aurornis]]''?
*{{extinct}}''[[Eosinopteryx]]''?
*{{extinct}}''[[Geminiraptor]]''
*{{extinct}}''[[Jianianhualong]]''
*{{extinct}}''[[Liaoningvenator]]''?
*{{extinct}}''[[Sinornithoides]]''
*{{extinct}}''[[Talos (dinosaur)|Talos]]''
*{{extinct}}''[[Xiaotingia]]''?
*{{extinct}}''[[Xixiasaurus]]''
*{{extinct}}[[Jinfengopteryx|Jinfengopteryginae]]?
*{{extinct}}[[Sinovenatorinae]]
*{{extinct}}[[Troodontinae]]
| synonyms =
Saurornithoididae [[Rinchen Barsbold|Barsbold]], 1974
}}

'''Troodontidae''' is a [[family]] of bird-like [[Theropoda|theropod]] [[dinosaur]]s. During most of the 20th century, troodontid fossils were few and scrappy and they have therefore been allied, at various times, with many [[dinosaur]]ian lineages. More recent fossil discoveries of complete and articulated specimens (including specimens which preserve [[feather]]s, [[Egg (biology)|eggs]] and [[embryos]], and complete juveniles), 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]].

==Physical characteristics==
Troodontids are a group of small, bird-like, [[wikt:gracile|gracile]] [[maniraptora]]ns. All troodontids have unique features of the skull, such as large numbers of closely spaced [[teeth]] in the lower jaw. Troodontids have sickle-claws and [[Velociraptor|raptor]]ial [[hands]], and some of the highest non-[[Bird|avian]] [[encephalization quotient]]s, suggesting that they were behaviourally advanced and had keen senses.<ref name="Lüetal">{{cite journal|author1=Junchang Lü |author2=Li Xu |author3=Yongqing Liu |author4=Xingliao Zhang |author5=Songhai Jia |author6=Qiang Ji |last-author-amp=yes |year= 2010 |title= A new troodontid (Theropoda: Troodontidae) from the Late Cretaceous of central China, and the radiation of Asian troodontids. |journal= Acta Palaeontologica Polonica |volume=55 |issue=3 |pages=381–388 |doi= 10.4202/app.2009.0047 |url=http://www.app.pan.pl/archive/published/app55/app20090047.pdf}}</ref> The largest troodontid was ''[[Troodon]]'', and the smallest was ''[[Anchiornis]]''. They had unusually long legs compared to other theropods, with a large, curved claw on their retractable second toes, similar to the "sickle-claw" of the [[Dromaeosauridae|dromaeosaurids]]. However, the sickle-claws of troodontids were not as large or recurved as in their relatives, and in some instances could not be held off the ground and "retracted" to the same degree. In at least one troodontid, ''[[Borogovia]]'', the second toe could not be held far off the ground at all and the claw was straight, not curved or sickle-like.
[[File:Saurornithoides mongoliensis.jpg|thumb|left|Skull of the troodontid ''[[Saurornithoides|Saurornithoides mongoliensis]]''.]]
Troodontids had unusually large brains among dinosaurs, comparable to those of living flightless birds. Their eyes were also large, and pointed forward, indicating that they had good [[binocular vision]]. The ears of troodontids were also unusual among theropods, having enlarged middle ear cavities, indicating acute hearing ability. The placement of this cavity near the eardrum may have aided in the detection of low-frequency sounds.<ref name ="currie1985">{{cite journal | last1 = Currie | first1 = P. J. | year = 1985 | title = Cranial anatomy of ''Stenonychosaurus inequalis'' (Saurischia, Theropoda) and its bearing on the origin of birds | url = | journal = Canadian Journal of Earth Sciences | volume = 22 | issue = | pages = 1643–1658 | doi=10.1139/e85-173}}</ref> In some troodontids, ears were also asymmetrical, with one ear placed higher on the skull than the other, a feature shared only with some [[owl]]s. The specialization of the ears may indicate that troodontids hunted in a manner similar to owls, using their hearing to locate small prey.<ref name="castanhinha&mateus2006">{{cite journal | last1 = Castanhinha | first1 = R. | last2 = Mateus | first2 = O. | year = 2006 | title = On the left-right asymmetry in dinosaurs | url = | journal = Journal of Vertebrate Paleontology | volume = 26 | issue = Supp. 3| page = 48A | doi = 10.1080/02724634.2006.10010069}}</ref>

[[File:Unnamed troodontid.jpg|thumb|Skeleton of an unnamed troodontid]]
Although most [[Paleontology|paleontologists]] believe that they were [[Predation|predatory]] [[carnivore]]s, the many small, coarsely serrated teeth, large denticle size, and U-shaped jaws of some species (particularly ''[[Troodon]]'') suggest that some species may have been [[Omnivore|omnivorous]] or [[Herbivore|herbivorous]]. Some suggest that the large denticle size is reminiscent of the teeth of extant [[iguana|iguanine]] lizards.<ref name="Mackovicky2004" /><ref name="holtzetal1998">{{cite journal | last1 = Holtz | first1 = T.R. Jr. | last2 = Brinkman | first2 = D.L. | last3 = Chandler | first3 = C.L. | year = 1998 | title = Denticle morphometrics and a possibly omnivorous feeding habit for the theropod dinosaur ''Troodon'' | url = http://www.geol.umd.edu/~tholtz/gaiatroo.pdf | journal = Gaia | volume = 15 | issue = | pages = 159–166 }}</ref> In contrast, a few species, such as ''[[Byronosaurus]]'', had large numbers of needle-like teeth, which seem best-suited for picking up small prey, such as birds, [[lizard]]s and small [[mammal]]s. Other morphological characteristics of the teeth, such as the detailed form of the denticles and the presence of blood grooves, also seem to indicate carnivory.<ref>{{cite journal | last1 = Currie | first1 = PJ | last2 = Dong | first2 = Z | year = 2001 | title = New information on Cretaceous troodontids (Dinosauria, Theropoda) from the People's Republic of China | url = | journal = Canadian Journal of Earth Sciences | volume = 38 | issue = | pages = 1753–1766 | doi=10.1139/e01-065}}</ref> Though little is known directly about the predatory behavior of troodontids, Fowler and colleagues theorize that the longer legs and smaller sickle claws (as compared to dromaeosaurids) indicates a more [[cursorial]] lifestyle, though the study indicates that troodontids were still likely to have used the [[ungual]]s for prey manipulation. The proportions of the metatarsals, tarsals and unguals of troodontids appear indicative of their having nimbler, but weaker feet, perhaps better adapted for capturing and subduing smaller prey. This suggests an ecological separation from the slower but more powerful Dromaeosauridae.<ref name=fowleretal2011>{{cite journal | last1 = Fowler | first1 = D.W. | last2 = Freedman | first2 = E.A. | last3 = Scannella | first3 = J.B. | last4 = Kambic | first4 = R.E. | year = 2011 | title = The Predatory Ecology of ''Deinonychus'' and the Origin of Flapping in Birds | url = http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0028964 | journal = PLoS ONE | volume = 6 | issue = 12| page = e28964 | doi = 10.1371/journal.pone.0028964 | pmid = 22194962 | pmc=3237572}}</ref>

==Paleobiology==
Many troodontid nests, including eggs that contain fossilized embryos, have been described. Hypotheses about troodontid reproduction have been developed from this evidence (see ''[[Troodon#Reproduction|Troodon]]'').

A few troodont [[fossil]]s, including specimens of ''[[Mei long|Mei]]'' and ''[[Sinornithoides]]'', demonstrate that these animals roosted like birds, with their heads tucked under their forelimbs.<ref name="Xu&Norell2004">{{cite journal | last1 = Xu | last2 = Norell | year = 2004 | title = A new troodontid dinosaur from China with avian-like sleeping posture | url = | journal = Nature | volume = 431 | issue = 7010| pages = 838–841 | doi=10.1038/nature02898 | pmid=15483610}}</ref> These fossils, as well as numerous skeletal similarities to birds and related feathered dinosaurs, support the idea that troodontids probably bore a bird-like feathered coat. The discovery of fully feathered, primitive troodontids, such as ''[[Jianianhualong]]'', lend support to this.

In 2004, Mark Norell and colleagues described two partial troodontid skulls (specimen numbers IGM 100/972 and IGM 100/974) found in a nest of [[oviraptorid]] eggs in the Djadokhta Formation of Mongolia. The nest is quite certainly that of an [[oviraptorosaur]], since an oviraptorid embryo is still preserved inside one of the eggs. The two partial troodontid skulls were first described by Norell et al. (1994) as dromaeosaurids, but reassigned to the troodontid ''[[Byronosaurus]]'' after further study.<ref name="Mackovicky2004">{{cite book |last=Mackovicky |first=Peter J. |author2=Norell, Mark A. |editor=Weishampel, David B. |editor2=Dodson, Peter |editor3=Osmólska, Halszka |title=The Dinosauria |edition=2nd |year=2004 |publisher=University of California Press |location=Berkeley |isbn=0-520-24209-2 |pages=184–195 |chapter=Troodontidae}}</ref><ref name=bever&norell2009>{{cite journal | last1 = Bever | first1 = G.S. | last2 = Norell | first2 = M.A. | year = 2009 | title = The perinate skull of ''Byronosaurus'' (Troodontidae) with observations on the cranial ontogeny of paravian theropods | url = | journal = American Museum Novitates | volume = 3657 | issue = | page = 51 }}</ref> The troodontids were either hatchlings or embryos, and fragments of eggshell are adhered to them although it seems to be oviraptorid eggshell. The presence of tiny troodontids in an oviraptorid nest is an enigma. Hypotheses explaining how they came to be there include that they were the prey of the adult oviraptorid, that they were there to prey on oviraptorid hatchlings, or that some troodontids may have been [[nest parasite]]s.<ref name="Norelletal1994">{{cite journal |last=Norell |first=Mark A. |author2=Clark, James M. |author3=Dashzeveg, Demberelyin |author4=Barsbold, Rhinchen |author5=Chiappe, Luis M. |author6=Davidson, Amy R. |author7=McKenna, Malcolm C. |author8=Perle, Altangerel |author9= Novacek, Michael J. |title=A theropod dinosaur embryo and the affinities of the Flaming Cliffs dinosaur eggs |journal=Science |volume=266 |issue=5186 |pages=779–782 |date=November 4, 1994 |doi=10.1126/science.266.5186.779 |pmid=17730398 }}</ref>

==Troodontids and bird evolution==
Troodontids are important in research into the origin of birds because they share many anatomical characters with early birds. Crucially, the substantially complete fossil identified as [[WDC DML 001]] ("Lori") is a possible troodontid from the [[Late Jurassic]] [[Morrison Formation]], close to the time of ''[[Archaeopteryx]]''. The discovery of Jurassic troodonts is positive physical evidence that derived deinonychosaurs were present before the time that avians arose. This fact strongly invalidates the "[[Temporal paradox (paleontology)|temporal paradox]]" cited by the few remaining opponents of the idea that birds are closely related to dinosaurs.<ref name="Hu et al.2009">{{cite journal | last1 = Hu | first1 = D. | last2 = Hou | first2 = L. | last3 = Zhang | first3 = L. | last4 = Xu | first4 = X. | year = 2009 | title = A pre-''Archaeopteryx'' troodontid theropod from China with long feathers on the metatarsus | url = | journal = Nature | volume = 461 | issue = | pages = 640–643 | doi = 10.1038/nature08322 | pmid = 19794491 }}</ref>

==Classification==
Troodontid fossils were among the first dinosaur remains described. Initially, [[Joseph Leidy|Leidy]] (1856) assumed they were [[lacertilia]]n (lizards), but, by 1924, they were referred to [[Dinosauria]] by [[Charles W. Gilmore|Gilmore]], who suggested that they were [[ornithischia]]ns and allied them with the [[pachycephalosauria]]n ''[[Stegoceras]]'' in a Troodontidae. It was not until 1945 that [[Charles Mortram Sternberg|C.M. Sternberg]] recognized Troodontidae as a theropod family. Since 1969, Troodontidae has typically been allied with [[Dromaeosauridae]], in a [[clade]] (natural group) known as [[Deinonychosauria]], but this was by no means a consensus. [[Thomas R. Holtz Jr.|Holtz]] (in 1994) erected the clade Bullatosauria, uniting [[Ornithomimosauria]] (the "ostrich-dinosaurs") and Troodontidae, on the basis of characteristics including, among others, an inflated braincase (''parabasisphenoid'') and a long, low opening in the upper jaw (the ''maxillary fenestra''). Features of the pelvis also suggested they were less advanced than dromaeosaurids. New discoveries of primitive troodontids from [[China]] (such as ''[[Sinovenator]]'' and ''Mei''), however, display strong similarities between Troodontidae, Dromaeosauridae and the primitive bird ''[[Archaeopteryx]]'', and most paleontologists, including Holtz, now consider troodontids to be much more closely related to birds than they are to ornithomimosaurs, causing the clade Bullatosauria to be abandoned.

One study of theropod systematics by members of the [[Theropod Working Group]] has uncovered striking similarities among the most basal dromaeosaurids, troodontids, and ''[[Archaeopteryx]]''. This clade is together called [[Paraves]] by Novas and Pol.<ref name="novas&pol2005">{{cite journal | last1 = Novas | first1 = F. E. | last2 = Pol | first2 = D. | year = 2005 | title = New evidence on deinonychosaurian dinosaurs from the Late Cretaceous of Patagonia | url = | journal = Nature | volume = 433| issue = 7028| pages = 858–861 | doi=10.1038/nature03285 | pmid=15729340}}</ref> The cladogram published in Hwang ''et al.'' found that ''Archaeopteryx'' represents a more basal branch of Paraves, and places dromaeosaurids and troodontids as more derived. This raises the possibility that aerodynamic behaviors could be ancestral to all of [[Deinonychosauria]].<ref name="hwangetal2002">{{cite journal | last1 = Hwang | first1 = S.H. | last2 = Norell | first2 = M.A. | last3 = Ji | first3 = Q. | last4 = Gao | first4 = K.-Q. | year = 2002 | title = New specimens of ''Microraptor zhaoianus'' (Theropoda: Dromaeosauridae) from Northeastern China | url = | journal = American Museum Novitates | volume = 3381 | issue = | pages = 1–44 | doi=10.1206/0003-0082(2002)381<0001:nsomzt>2.0.co;2}}</ref> The extensive cladistic analysis conducted by Turner et al. (2012) supported the monophyly of Troodontidae.<ref name=TMN2012>{{Cite journal | last1 = Turner | first1 = A. H. | last2 = Makovicky | first2 = P. J. | last3 = Norell | first3 = M. A. | title = A Review of Dromaeosaurid Systematics and Paravian Phylogeny | doi = 10.1206/748.1 | journal = Bulletin of the American Museum of Natural History | volume = 371 | pages = 1 | year = 2012 | pmid = | pmc = }}</ref>

===Relationships===
There are multiple possibilities of the genera included in Troodontidae as well as how they are related. Very primitive species, such as ''Anchiornis huxleyi'', have alternately been found to be early troodontids, early members of the closely related group Avialae, or more primitive paravians by various studies. The [[cladogram]] below follows the results of a study by Lefèvre''et al.'', 2017.<ref name=serikornis>Ulysse Lefèvre, Andrea Cau, Aude Cincotta, Dongyu Hu, Anusuya Chinsamy, François Escuillié & Pascal Godefroit (2017). A new Jurassic theropod from China documents a transitional step in the macrostructure of feathers. ''The Science of Nature'', '''104''': 74 (advance online publication). {{doi|10.1007/s00114-017-1496-y}}</ref>
{{clade| style=font-size:75%;line-height:80%
|label1=[[Eumaniraptora]]
|1={{clade
|1=[[Avialae]]
|label2=[[Deinonychosauria]]
|2={{clade
|1=[[Dromaeosauridae]]
|label2='''Troodontidae'''
|2={{clade
|1={{clade
|1=''[[Jinfengopteryx]]''
|2=''[[Mei (dinosaur)|Mei]]''}}
|2={{clade
|1=''[[Sinovenator]]''
|2={{clade
|1=''[[Sinusonasus]]''
|2={{clade
|1=''[[Sinornithoides]]''
|2={{clade
|1={{clade
|1=''[[Byronosaurus]]''
|2=''[[Gobivenator]]'' }}
|2={{clade
|1={{clade
|1=''[[Troodon]]''
|2=''[[Borogovia]]''}}
|2=''[[Saurornithoides]]''
|3=''[[Zanabazar (dinosaur)|Zanabazar]]''}} }} }} }} }} }} }} }} }}

Shen ''et al.'' (2017a) explored troodontid phylogeny using a modified version of the Tsuihiji ''et al.'' (2014) analysis.<ref name=Gobivenator>{{Cite journal | last1 = Tsuihiji | first1 = T. | last2 = Barsbold | first2 = R. | last3 = Watabe | first3 = M. | last4 = Tsogtbaatar | first4 = K. | last5 = Chinzorig | first5 = T. | last6 = Fujiyama | first6 = Y. | last7 = Suzuki | first7 = S. | doi = 10.1007/s00114-014-1143-9 | title = An exquisitely preserved troodontid theropod with new information on the palatal structure from the Upper Cretaceous of Mongolia | journal = Naturwissenschaften | year = 2014 | pmid = 24441791| pmc = | volume=101 | pages=131–142}}</ref> It was in turn based on data published by Gao ''et al.'' (2012), a slightly modified version of the Xu ''et al.'' (2011) analysis<ref name=Mei2012>{{Cite journal | last1 = Gao | first1 = C. | last2 = Morschhauser | first2 = E. M. | last3 = Varricchio | first3 = D. J. | last4 = Liu | first4 = J. | last5 = Zhao | first5 = B. | editor1-last = Farke | editor1-first = Andrew A | title = A Second Soundly Sleeping Dragon: New Anatomical Details of the Chinese Troodontid ''Mei long'' with Implications for Phylogeny and Taphonomy | doi = 10.1371/journal.pone.0045203 | journal = PLoS ONE | volume = 7 | issue = 9 | pages = e45203 | year = 2012 | pmid = 23028847| pmc = 3459897}}</ref>, focusing on advanced troodontids. A simplified version is shown below.<ref>{{Cite journal|author1=Cai-zhi Shen |author2=Bo Zhao |author3=Chun-ling Gao |author4=Jun-chang Lü |author5=Martin Kundrát |year=2017 |title=A New Troodontid Dinosaur (''Liaoningvenator curriei'' gen. et sp. nov.) from the Early Cretaceous Yixian Formation in Western Liaoning Province |journal=Acta Geoscientica Sinica |volume=38 |issue=3 |pages=359–371 |doi=10.3975/cagsb.2017.03.06 }}</ref>

{{clade| style=font-size:85%;line-height:85%
|label1=[[Deinonychosauria]] 
|1={{clade
|1=[[Dromaeosauridae]]
|label2= '''Troodontidae''' 
|2={{clade
|1=''[[Sinovenator]]''
|2={{clade
|1={{clade
|1={{clade
|1=''[[Eosinopteryx]]''
|2=''[[Liaoningvenator]]'' }}
|2={{clade
|1=''[[Anchiornis]]''
|2=''[[Xiaotingia]]'' }} }}
|2={{clade
|1=''[[Talos (dinosaur)|Talos]]''
|2={{clade
|1=''[[Mei (dinosaur)|Mei]]''
|2={{clade
|1=''[[Byronosaurus]]''
|2={{clade
|1=IGM 100/140
|2=[[SPS 100/44]]
|3=''[[Sinornithoides]]''
|4={{clade
|1=''[[Gobivenator]]''
|2={{clade
|1=''[[Linhevenator]]''
|2=''[[Philovenator]]''}}
|3={{clade
|1=''[[Troodon]]''
|2={{clade
|1=''[[Saurornithoides]]''
|2=''[[Zanabazar (dinosaur)|Zanabazar]]'' }} }} }} }} }} }} }} }} }} }} }}

In 2014, Brusatte, Lloyd, Wang and Norell published an analysis on [[Coelurosauria]], based on data from Turner ''et al.'' (2012) who named a third subfamily of troodontids, Jinfengopteryginae.<ref name=TMN2012>{{Cite journal | last1 = Turner | first1 = A. H. | last2 = Makovicky | first2 = P. J. | last3 = Norell | first3 = M. A. | title = A Review of Dromaeosaurid Systematics and Paravian Phylogeny | doi = 10.1206/748.1 | journal = Bulletin of the American Museum of Natural History | volume = 371 | pages = 1 | year = 2012 | pmid = | pmc = }}</ref> Their analysis included more basal troodontid species but failed to resolve many of their interrelationships, resulting in large "[[polytomy|polytomies]]" (sets of species where the branching order in the family tree is uncertain).<ref name="brusatteetal">{{Cite journal | doi = 10.1016/j.cub.2014.08.034| title = Gradual Assembly of Avian Body Plan Culminated in Rapid Rates of Evolution across the Dinosaur-Bird Transition| journal = Current Biology| volume = 24| issue = 20| pages = 2386| year = 2014| last1 = Brusatte | first1 = S. L. | last2 = Lloyd | first2 = G. T. | last3 = Wang | first3 = S. C. | last4 = Norell | first4 = M. A. | pmid=25264248}}</ref> An updated version of the Brusatte ''et al.'' analysis was provided by Shen ''et at.'' (2017b), who included more taxa and recovered greater resolution. Shen ''et at.'' named a fourth subfamily of troodontids, the Sinovenatorinae. A simplified version of their analysis is shown below.<ref>{{Cite journal|author1=Caizhi Shen |author2=Junchang Lü |author3=Sizhao Liu |author4=Martin Kundrát |author5=Stephen L. Brusatte |author6=Hailong Gao |year=2017 |title=A new troodontid dinosaur from the Lower Cretaceous Yixian Formation of Liaoning Province, China |journal=Acta Geologica Sinica (English Edition) |volume=91 |issue=3 |pages=763–780 |url=http://www.geojournals.cn/dzxben/ch/reader/view_abstract.aspx?file_no=2017endzxb03001&flag=1 }}</ref>

{{clade| style=font-size:85%; line-height:85%
|label1=[[Deinonychosauria]] 
|1={{clade
|1=[[Dromaeosauridae]]
|label2= '''Troodontidae''' 
|2={{clade
|1={{clade
|1=''[[Eosinopteryx]]''
|2=''[[Anchiornis]]''
|3={{clade
|1=''[[Aurornis]]''
|2=''[[Xiaotingia]]'' }} }}
|2={{clade
|1=IGM 100/44
|2=''[[Byronosaurus]]''
|3=''[[Xixiasaurus]]''
|label4= Jinfengopteryginae 
|4={{clade
|1=IGM 100/1323
|2={{clade
|1=IGM 100/1128
|2=''[[Jinfengopteryx]]'' }} }}
|label5= Sinovenatorinae 
|5={{clade
|1=''[[Mei (dinosaur)|Mei]]''
|2={{clade
|1=''[[Sinovenator]]''
|2={{clade
|1=''[[Daliansaurus]]''
|2=''[[Sinusonasus]]'' }} }} }}
|6={{clade
|1=''[[Sinornithoides]]''
|2={{clade
|1=''[[Troodon]]''
|2={{clade
|1=''[[Zanabazar]]''
|2=''[[Saurornithoides]]'' }} }} }} }} }} }} }}

==See also==
* [[Timeline of troodontid research]]

==References==
{{Reflist}}

{{Commons|Troodontidae}}
{{Portal|Dinosaurs}}

{{Troodontidae}}
[[Category:Troodontids]]
[[Category:Tithonian first appearances]]
[[Category:Tithonian taxonomic families]]
[[Category:Berriasian taxonomic families]]
[[Category:Valanginian taxonomic families]]
[[Category:Hauterivian taxonomic families]]
[[Category:Barremian taxonomic families]]
[[Category:Aptian taxonomic families]]
[[Category:Albian taxonomic families]]
[[Category:Cenomanian taxonomic families]]
[[Category:Turonian taxonomic families]]
[[Category:Coniacian taxonomic families]]
[[Category:Santonian taxonomic families]]
[[Category:Campanian taxonomic families]]
[[Category:Maastrichtian taxonomic families]]
[[Category:Maastrichtian extinctions]]

Template:Troodontidae

2017-08-27T17:30:03Z

Wikkler: We don't know if they're avialans or troodontids. Also, Xiaotingia has also been found to possibly not be in here.

{{Navbox
|name = Troodontidae
|title = [[Troodontidae]]
|state = {{{state|autocollapse}}}
|bodyclass = hlist

|above =
* Kingdom: [[Animal]]ia
* Phylum: [[Chordate|Chordata]]
* Class: [[Reptile|Sauropsida]]
* ''Clade'': [[Dinosauria]]
* Order: [[Saurischia]]
* ''Clade'': [[Maniraptora]]

|image = [[File:Byronosaurus.jpg|100px|''Byronosaurus jaffei'']]

|group1= [[Troodontidae]]
|list1 = {{Navbox|subgroup
|list1 =
* ''[[Borogovia]]''
* ''[[Byronosaurus]]''
* ''[[Geminiraptor]]''
* ''[[Jianianhualong]]''
* ''[[Sinornithoides]]''
* ''[[Talos (dinosaur)|Talos]]''
* ''[[Xixiasaurus]]''
* ''[[Polyodontosaurus]]''

|group2 = Anchiornithinae?
|list2 =
* ''[[Anchiornis]]''
* ''[[Aurornis]]''
* ''[[Eosinopteryx]]''?
* ''[[Liaoningvenator]]''?
* ''[[Xiaotingia]]''?
|group3 = [[Jinfengopteryx|Jinfengopteryginae]]
|list3 =
* ''[[Jinfengopteryx]]''
|group4 = [[Sinovenatorinae]]
|list4 =
* ''[[Daliansaurus]]''
* ''[[Mei (dinosaur)|Mei]]''
* ''[[Sinovenator]]''
* ''[[Sinusonasus]]''
|group5 = [[Troodontinae]]
|list5 =
* ''[[Albertavenator]]''
* ''[[Gobivenator]]''
* ''[[Latenivenatrix]]''
* ''[[Linhevenator]]''
* ''[[Pectinodon]]''
* ''[[Saurornithoides]]''
* ''[[Stenonychosaurus]]''
* ''[[Troodon]]''
* ''[[Zanabazar (dinosaur)|Zanabazar]]''
* ''[[Urbacodon]]''

}}
}}<noinclude>
[[Category:Archosaur navigational boxes]]
</noinclude>

Template:Troodontidae

2017-08-27T16:11:19Z

Wikkler: It redirects to Anchiornis so there's no reason to have the link right now.

{{Navbox
|name = Troodontidae
|title = [[Troodontidae]]
|state = {{{state|autocollapse}}}
|bodyclass = hlist

|above =
* Kingdom: [[Animal]]ia
* Phylum: [[Chordate|Chordata]]
* Class: [[Reptile|Sauropsida]]
* ''Clade'': [[Dinosauria]]
* Order: [[Saurischia]]
* ''Clade'': [[Maniraptora]]

|image = [[File:Byronosaurus.jpg|100px|''Byronosaurus jaffei'']]

|group1= [[Troodontidae]]
|list1 = {{Navbox|subgroup
|list1 =
* ''[[Borogovia]]''
* ''[[Byronosaurus]]''
* ''[[Geminiraptor]]''
* ''[[Jianianhualong]]''
* ''[[Sinornithoides]]''
* ''[[Talos (dinosaur)|Talos]]''
* ''[[Xixiasaurus]]''
* ''[[Polyodontosaurus]]''

|group2 = Anchiornithinae
|list2 =
* ''[[Anchiornis]]''
* ''[[Aurornis]]''
* ''[[Eosinopteryx]]''?
* ''[[Liaoningvenator]]''?
* ''[[Xiaotingia]]''
|group3 = [[Jinfengopteryx|Jinfengopteryginae]]
|list3 =
* ''[[Jinfengopteryx]]''
|group4 = [[Sinovenatorinae]]
|list4 =
* ''[[Daliansaurus]]''
* ''[[Mei (dinosaur)|Mei]]''
* ''[[Sinovenator]]''
* ''[[Sinusonasus]]''
|group5 = [[Troodontinae]]
|list5 =
* ''[[Albertavenator]]''
* ''[[Gobivenator]]''
* ''[[Latenivenatrix]]''
* ''[[Linhevenator]]''
* ''[[Pectinodon]]''
* ''[[Saurornithoides]]''
* ''[[Stenonychosaurus]]''
* ''[[Troodon]]''
* ''[[Zanabazar (dinosaur)|Zanabazar]]''
* ''[[Urbacodon]]''

}}
}}<noinclude>
[[Category:Archosaur navigational boxes]]
</noinclude>