Neornithischia

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Neornithischians
Temporal range: Middle JurassicLate Cretaceous, 170–66  Ma
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(possible Early Jurassic record)
Neornithischia Infobox Panoply.png
Six neornithischians (top left to bottom right): Psittacosaurus , Styracosaurus , Thescelosaurus , Kulindadromeus , Hypacrosaurus , Pachycephalosaurus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Ornithischia
Clade: Genasauria
Clade: Neornithischia
Cooper, 1985
Subgroups [1]

Neornithischia ("new ornithischians") is a clade of the dinosaur order Ornithischia. It is 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.

Contents

Neornithischians include a variety of basal forms historically known as "hypsilophodonts", including the Thescelosauridae. As these taxa do not all form a monophyletic clade, the term 'small-bodied early diverging ornithischian' (SBEDO) has been used to refer to these as a collective group. [2] In addition, there are derived forms classified in the groups Marginocephalia and Ornithopoda. The former includes clades Pachycephalosauria and Ceratopsia, while the latter typically 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 than to Ankylosaurus magniventris or Stegosaurus stenops ". [3] In 2021, Neornithischia was given a formal definition under the PhyloCode: "The largest clade containing Iguanodon bernissartensis and Triceratops horridus but not Ankylosaurus magniventris and Stegosaurus stenops ." [4]

A 2017 study by Matthew G. Baron, 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. [5]

Pyrodontia was named by André O. Fonseca and colleagues in 2024 to unite the members of Thescelosauridae and Cerapoda. It's name translates to "fire teeth" in reference to the rapid diversification of this clade during the Middle Jurassic. [1] The clade is formally defined in the PhyloCode as "the smallest clade containing Ceratops montanus , Iguanodon bernissartensis , Pachycephalosaurus wyomingensis , and Thescelosaurus neglectus . [1]

Cerapoda is the most diverse clade within Neornithischia. The name "Cerapoda" is a portmanteaux of "Ceratopsia" and "Ornithopoda". As the name suggests, the clade is divided into two groups: Ornithopoda ("bird-foot") and Marginocephalia ("fringed heads"). The latter group includes the Pachycephalosauria ("thick-headed lizards") and Ceratopsia ("horned faces"). The following taxonomy follows Richard J. Butler, Paul Upchurch and David B. Norman, 2008 (and Butler et al., 2011) unless otherwise noted. [3] [6]

Cerapoda was first named by Sereno in 1986 and defined by him as " Parasaurolophus walkeri Parks, 1922, Triceratops horridus Marsh, 1889, their most recent common ancestor and all descendants". [3] A similar clade Neornithopoda was tentatively proposed by David B. Norman to unite ceratopsians with advanced ornithopods in a 1984 paper. [7] In 2021, Cerapoda was given a formal definition under the PhyloCode: "The smallest clade containing Iguanodon bernissartensis Boulenger in Beneden, 1881, Pachycephalosaurus wyomingensis (Gilmore, 1931), and Triceratops horridus Marsh, 1889." [4]

The phylogenetic study of Fonseca and colleagues in 2024 recovered results similar to the previous analyses of Boyd and Herne and colleagues, [8] [9] with thescelosaurids outside Ornithopoda and heterodontosaurids outside Neornithischia, while other aspects of relationships, like the placement of Changmiania or the grouping within Ornithopoda were novel results. Their equal-weights results are below. [1]

Ornithischia

Asilisaurus

Related Research Articles

<span class="mw-page-title-main">Ornithischia</span> Extinct clade of dinosaurs

Ornithischia is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name Ornithischia, or "bird-hipped", reflects this similarity and is derived from the Greek stem ornith- (ὀρνιθ-), meaning "bird", and ischion (ἴσχιον), meaning "hip". However, birds are only distantly related to this group, as birds are theropod dinosaurs.

<span class="mw-page-title-main">Thyreophora</span> Extinct clade of dinosaurs

Thyreophora is a group of armored ornithischian dinosaurs that lived from the Early Jurassic until the end of the Cretaceous.

<span class="mw-page-title-main">Dinosaur classification</span> Various classifications of Dinosauria

Dinosaur classification began in 1842 when Sir Richard Owen placed Iguanodon, Megalosaurus, and Hylaeosaurus in "a distinct tribe or suborder of Saurian Reptiles, for which I would propose the name of Dinosauria." In 1887 and 1888 Harry Seeley divided dinosaurs into the two orders Saurischia and Ornithischia, based on their hip structure. These divisions have proved remarkably enduring, even through several seismic changes in the taxonomy of dinosaurs.

<span class="mw-page-title-main">Ornithopoda</span> Extinct suborder of dinosaurs

Ornithopoda is a clade of ornithischian dinosaurs, called ornithopods. They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively small-sized, while advanced members of the subgroup Iguanodontia became quadrupedal and developed large body size. Their major evolutionary advantage was the progressive development of a chewing apparatus that became the most sophisticated ever developed by a non-avian dinosaur, rivaling that of modern mammals such as the domestic cow. They reached their apex of diversity and ecological dominance in the hadrosaurids, before they were wiped out by the Cretaceous–Paleogene extinction event along with all other non-avian dinosaurs. Members are known worldwide.

<span class="mw-page-title-main">Ceratopsia</span> Extinct suborder of Dinosaurs

Ceratopsia or Ceratopia is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. The earliest known ceratopsian, Yinlong downsi, lived between 161.2 and 155.7 million years ago. The last ceratopsian species, Triceratops prorsus, became extinct during the Cretaceous–Paleogene extinction event, 66 million years ago.

<span class="mw-page-title-main">Pachycephalosauria</span> Extinct clade of dinosaurs

Pachycephalosauria is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachycephalosaurs lived during the Late Cretaceous Period, dating between about 85.8 and 66 million years ago. They are exclusive to the Northern Hemisphere, all of them being found in 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.

<span class="mw-page-title-main">Hypsilophodontidae</span> Extinct family of dinosaurs

Hypsilophodontidae is a traditionally used family of ornithopod dinosaurs, generally considered invalid today. It historically included many small bodied bipedal neornithischian taxa from around the world, and spanning from the Middle Jurassic until the Late Cretaceous. This inclusive status was supported by some phylogenetic analyses from the 1990s and mid 2000s, although there have also been many finding that the family is an unnatural grouping which should only include the type genus, Hypsilophodon, with the other genera being within clades like Thescelosauridae and Elasmaria. A 2014 analysis by Norman recovered a grouping of Hypsilophodon, Rhabdodontidae and Tenontosaurus, which he referred to as Hypsilophodontia. That clade is formally defined in the PhyloCode as "the smallest clade within Ornithopoda containing Hypsilophodon foxii and Tenontosaurus tilletti provided it does not include Iguanodon bernissartensis". All other analyses from around the same time have instead found these latter taxa to be within Iguanodontia. The family Hypsilophodontidae is formally defined in the PhyloCode by Daniel Madzia and colleagues in 2021 as "the largest clade containing Hypsilophodon foxii, but not Iguanodon bernissartensis and Rhabdodon priscus".

<i>Stenopelix</i> Extinct genus of dinosaurs

Stenopelix is a genus of small marginocephalian dinosaur, possibly a basal ceratopsian, from the Early Cretaceous of Germany. It lived in the late Berriasian Stage of the Cretaceous period, approximately 140 myr ago. The genus is based on a partial skeleton lacking the skull, and its classification is based on characteristics of the hips.

<span class="mw-page-title-main">Hadrosauroidea</span> Extinct clade of dinosaurs

Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or Hadrosauridae, and all dinosaurs more closely related to them than to Iguanodon. Their remains have been recovered in Asia, Europe, Africa and the Americas. Many primitive hadrosauroids, such as the Asian Probactrosaurus and Altirhinus, have traditionally been included in a paraphyletic "Iguanodontidae". With cladistic analysis, the traditional Iguanodontidae has been largely disbanded, and probably includes only Iguanodon and perhaps its closest relatives.

<span class="mw-page-title-main">Heterodontosauridae</span> Extinct family of dinosaurs

Heterodontosauridae is a family of ornithischian dinosaurs that were likely among the most basal (primitive) members of the group. Their phylogenetic placement is uncertain but they are most commonly found to be primitive, outside of the group Genasauria. Although their fossils are relatively rare and their group small in numbers, they have been found on all continents except Australia and Antarctica, with a range spanning the Early Jurassic to the Early Cretaceous.

<span class="mw-page-title-main">Chaoyangsauridae</span> Family of ceratopsian dinosaurs

Chaoyangsauridae is a family of ceratopsian dinosaurs. They are among the earliest known marginocephalian dinosaurs, with remains dating to about 160 million years ago, during the Late Jurassic period. Members of this group had sharp beaks for snipping off leaves to eat, and a very small frill.

<span class="mw-page-title-main">Dryosauridae</span> Extinct family of dinosaurs

Dryosauridae was a family of primitive iguanodonts, first proposed by Milner & Norman in 1984. They are known from Middle Jurassic to Early Cretaceous rocks of Africa, Europe, and North America.

<span class="mw-page-title-main">Thescelosauridae</span> Extinct family of dinosaurs

Thescelosauridae is a clade of neornithischians from the Cretaceous of East Asia and North America. The group was originally used as a name by Charles M. Sternberg in 1937, but was not formally defined until 2013, where it was used by Brown and colleagues as the group uniting Thescelosaurus and Orodromeus, based on their phylogenetic results. During a phylogenetic revision of neornithischians by Clint Boyd in 2015, the authorship of Thescelosauridae was given to Brown and colleagues, which meant that the similar name Parksosauridae, informally defined in 2002 by Buchholz, would have had priority over Thescelosauridae. The two clades had slightly different definitions, with Parksosauridae referring to all animals closer to Parksosaurus than Hypsilophodon, but they contained the same taxa so Boyd used Parksosauridae under the assumption it had priority. However, in formalizing the clade following the regulations of the PhyloCode, Madzia, Boyd, and colleagues identified in 2021 that Sternberg was the proper authority for Thescelosauridae, giving it priority over Parksosauridae. As well, they gave Thescelosauridae the definition of the largest clade containing Thescelosaurus neglectus but not Iguanodon bernissartensis, as long as Hypsilophodon foxii was not in the group, modifying previous definitions for Thescelosauridae in order to maintain its modern use, so that the clade was not applied if Thescelosaurus fell within Hypsilophodontidae, a family that has not been recently used but may be revived if the systematic position of Hypsilophodon was solidified at some point in the future. Madzia et al. identified the analysis of Madzia et al. in 2018 as the reference analysis for the name Thescelosauridae, an analysis based on a revised version of the 2015 Boyd analysis.

<i>Albalophosaurus</i> Extinct genus of dinosaurs

Albalophosaurus is a genus of marginocephalian ornithischian dinosaur that lived in Japan during the Early Cretaceous. The type species is Albalophosaurus yamaguchiorum.

<span class="mw-page-title-main">Ornithoscelida</span> Clade of reptiles

Ornithoscelida is a proposed clade that includes various major groupings of dinosaurs. An order Ornithoscelida was originally proposed by Thomas Henry Huxley but later abandoned in favor of Harry Govier Seeley's division of Dinosauria into Saurischia and Ornithischia. The term was revived in 2017 after a new cladistic analysis by Baron et al.

<i>Koreanosaurus</i> Extinct genus of dinosaurs

Koreanosaurus is a genus of orodromine neornithischian dinosaur. One species has been described, Koreanosaurus boseongensis.

<span class="mw-page-title-main">Elasmaria</span> Extinct clade of dinosaurs

Elasmaria is a clade of ornithopods known from Cretaceous deposits in South America, Antarctica, and Australia that contains many bipedal ornithopods that were previously considered "hypsilophodonts".

<span class="mw-page-title-main">Jeholosauridae</span> Extinct family of dinosaurs

Jeholosauridae is an extinct group of herbivorous neornithischian dinosaurs from the Cretaceous Period. The family was first proposed by Han et al. in 2012. The jeholosaurids were defined as all neornithischians more closely related to Jeholosaurus shangyuanensis than to Hypsilophodon foxii, Iguanodon bernissartensis, Protoceratops andrewsi, Pachycephalosaurus wyomingensis, or Thescelosaurus neglectus. The Jeholosauridae includes the type genus Jeholosaurus, and also possibly Yueosaurus, Changchunsaurus, and Haya.

<i>Laquintasaura</i> Extinct genus of dinosaurs

Laquintasaura is a genus of Venezuelan ornithischian dinosaur containing only the type species Laquintasaura venezuelae. It is known for being one of the and most primitive ornithischians in the fossil record, as well as the first dinosaur to have been identified from Venezuela. The name is derived from the La Quinta Formation, where it was discovered and the feminine Greek suffix for lizard, with the specific name referring to the country of Venezuela. It is known from hundreds of fossil elements, all derived from a single extensive bonebed locality. Initially discovered by French palaeontologists, numerous expeditions have been conducted to excavate from the bonebed, largely led by Marcelo R Sánchez-Villagra. Once thought to represent remains of Lesothosaurus, it was formally named in a 2014 study; much of the abundant material was not yet prepared at the time and research remains ongoing.

References

  1. 1 2 3 4 5 Fonseca, A.O.; Reid, I.J.; Venner, A.; Duncan, R.J.; Garcia, M.S.; Müller, R.T. (2024). "A comprehensive phylogenetic analysis on early ornithischian evolution". Journal of Systematic Palaeontology. 22 (1): 2346577. doi:10.1080/14772019.2024.2346577.
  2. Avrahami, Haviv M.; Makovicky, Peter J.; Tucker, Ryan T.; Zanno, Lindsay E. (2024-07-09). "A new semi-fossorial thescelosaurine dinosaur from the Cenomanian-age Mussentuchit Member of the Cedar Mountain Formation, Utah". The Anatomical Record . doi: 10.1002/ar.25505 . ISSN   1932-8486. PMID   38979930.
  3. 1 2 3 Richard J. Butler; Paul Upchurch; David B. Norman (2008). "The phylogeny of the ornithischian dinosaurs". Journal of Systematic Palaeontology. 6 (1): 1–40. doi:10.1017/S1477201907002271. S2CID   86728076.
  4. 1 2 Madzia D, Arbour VM, Boyd CA, Farke AA, Cruzado-Caballero P, Evans DC. 2021. The phylogenetic nomenclature of ornithischian dinosaurs. PeerJ9:e12362
  5. Matthew G. Baron; David B. Norman; Paul M. Barrett (2016). "Postcranial anatomy of Lesothosaurus diagnosticus (Dinosauria: Ornithischia) from the Lower Jurassic of southern Africa: implications for basal ornithischian taxonomy and systematics". Zoological Journal of the Linnean Society. doi: 10.1111/zoj.12434 .
  6. Richard J. Butler; Jin Liyong; Chen Jun; Pascal Godefroit (2011). "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". Palaeontology. 54 (3): 667–683. doi:10.1111/j.1475-4983.2011.01046.x.
  7. Norman, D.B. (1984). "A systematic reappraisal of the reptile order Ornithischia". In Reif, W.E.; Westphal, F. (eds.). Third Symposium on Mesozoic Terrestrial Ecosystems, Short Papers. ATTEMPTO Verlag. pp. 157–162.
  8. Clint A. Boyd (2015). "The systematic relationships and biogeographic history of ornithischian dinosaurs". PeerJ. 3 (e1523): e1523. doi: 10.7717/peerj.1523 . PMC   4690359 . PMID   26713260.
  9. Herne, Matthew C.; Nair, Jay P.; Evans, Alistair R.; Tait, Alan M. (2019). "New small-bodied ornithopods (Dinosauria, Neornithischia) from the Early Cretaceous Wonthaggi Formation (Strzelecki Group) of the Australian-Antarctic rift system, with revision of Qantassaurus intrepidus Rich and Vickers-Rich, 1999". Journal of Paleontology. 93 (3): 543–584. doi: 10.1017/jpa.2018.95 .