Ornithopoda

Ornithopods (/ɔːrˈnɪθəˌpɒdz, ˈɔːrnɪ-/[1][2]) or members of the clade Ornithopoda (/ˌɔːrnɪˈθɒpədə/ or /ɔːrˌnɪθəˈpoʊdə, ˌɔːrnɪ-/[3]) are a group of ornithischian dinosaurs that started out as small, bipedal running grazers, and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world, and dominated the North American landscape. 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 in the duck-bills (hadrosaurs), before they were wiped out by the Cretaceous–Paleogene extinction event along with all other non-avian dinosaurs. Members are known from all seven continents, though they are generally rare in the Southern Hemisphere.

Ornithopods
Temporal range: Middle Jurassic - Late Cretaceous, 163–66 Ma
FMNH Parasaurolophus fossil
Mounted skeleton of Parasaurolophus cyrtocristatus, Field Museum of Natural History
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Ornithischia
Clade: Cerapoda
Suborder: Ornithopoda
Marsh, 1881
Subgroups

History of research

In 1870, Thomas Henry Huxley listed Iguanodontidae (coined by Cope a year earlier[4]) as one of his three families of dinosaurs (alongside Megalosauridae and Scelidosauridae), including within it the genera Iguanodon, Hypsilophodon, and Hadrosaurus, in addition to Cetiosaurus and tentatively Stenopelix.[5] The term Ornithopoda was erected by Othniel Charles Marsh in 1881 as part of his then still ongoing investigation of the classification of Dinosauria. It was considered one of the four definite orders of dinosaurs, the others being Theropoda, Sauropoda, and Stegosauria (Hallopoda was considered a possible fifth). He subdivided the order into three families: Camptonotidae, Iguanodontidae, and Hadrosauridae; the former was a new name, whereas the latter two were carried over from the nomenclatures of Huxley and Edward Drinker Cope respectively. Within Camptonotidae he included the European Hypsilophodon and three American taxa he named himself, Camptonotus, Laosaurus, and Nanosaurus.[6] Camptonotus was in 1885 renamed to Camptosaurus, as the original name was pre-occupied by a cricket; the associated family follow suit, becoming Camptosauridae.[7] In Iguanodontidae, only found in Europe, he included Iguanodon and Vectisaurus. In Hadrosauridae, he included Hadrosaurus, Cionodon, and tentatively Agathaumas.[6]

Description

Iguanodon feet
Three-toed feet of Iguanodon

Ornithopoda means "bird feet", from the Greek ornithos ("bird") and pous ("feet"); this refers to their characteristic three-toed feet, although many early forms retained four toes. They were also characterized by having no armour, the development of a horny beak, an elongated pubis that eventually extended past the ilium, and a missing hole in the lower jaw. A variety of ornithopods and related cerapods had thin cartilaginous plates along the outside of the ribs; in some cases, these plates mineralized and were fossilized. The function of these intercostal plates is unknown. They have been found with Hypsilophodon, Othnielosaurus, Parksosaurus, Talenkauen, Thescelosaurus,[8] and Macrogryphosaurus to date.[9]

Iguanodontian Sizes
Size of a variety of numerous ornithopods

The early ornithopods were only about 1 metre (3 feet) long, but probably very fast. They had a stiff tail, like the theropods, to help them balance as they ran on their hind legs. Later ornithopods became more adapted to grazing on all fours; their spines curved, and came to resemble the spines of modern ground-feeders such as the bison. As they became more adapted to eating while bent over, they became facultative quadrupeds; still running on two legs, and comfortable reaching up into trees; but spending most of their time walking or grazing while on all fours. The taxonomy of dinosaurs previously ascribed to the Hypsilophodontidae is problematic. The group previously consisted of all non-iguanodontian bipedal ornithischians, but a phylogenetic reappraisal has shown such species to be paraphyletic. As such, the hypsilophodont family is currently represented only by Hypsilophodon.[10]

Later ornithopods became larger, but never rivalled the incredible size of the long-necked, long-tailed sauropods that they partially supplanted. The very largest, such as Shantungosaurus, were as heavy as medium-sized sauropods at up to 23 metric tons (25 short tons) but never grew much beyond 15 metres (50 feet).

Classification

Gideonmantellia
An artist's interpretation of Gideonmantellia, sometimes considered one of the most primitive ornithopods
Muttaburrasaurus NT
Restoration of Muttaburrasaurus, an early iguanodont
D. lettowvorbecki
Skeleton of Dysalotosaurus, a dryosaurid ornithopod from the Jurassic
Life restoration of Iguanacolossus
Life restoration of Iguanacolossus, an early styracosternan
Mantellisaurus atherfieldensis Steveoc
Reconstruction of Mantellisaurus, a primitive member of the Hadrosauriformes
Oxford Edmontosaurus
Mounted skeleton of Edmontosaurus, a saurolophine hadrosaur, and one of the last ornithopods

Historically, most indeterminate ornithischian bipeds were lumped in as ornithopods. Most have since been reclassified.

Taxonomy

Ornithopoda is usually given the rank of Suborder, within the order Ornithischia. While ranked taxonomy has largely fallen out of favour among dinosaur paleontologists, some researchers have continued to employ such a classification, though sources have differed on what its rank should be. Benton (2004) placed it as an infraorder within the suborder Cerapoda (originally named as an unranked clade), while others, such as Ibiricu et al. 2010, have retained it at its traditional ranking of suborder.[11]

Phylogeny

The cladogram below follows a 2017 analysis by Madzia et al.:[12]

Clypeodonta

Hypsilophodon

Cerapoda

Marginocephalia

Ornithopoda

Gideonmantellia

Elasmaria

Burianosaurus

Iguanodontia
Rhabdodontomorpha

Muttaburrasaurus

Rhabdodontidae

Tenontosaurus

Dryomorpha

Dryosauridae

Ankylopollexia

Camptosaurus

Iguanodon

Ouranosaurus

References

  1. ^ "ornithopod - definition of ornithopod in English from the Oxford dictionary". OxfordDictionaries.com. Retrieved 2016-01-20.
  2. ^ "Ornithopod". Merriam-Webster Dictionary.
  3. ^ See analogous pronunciation of "gastropoda - definition of gastropoda in English from the Oxford dictionary". OxfordDictionaries.com. Retrieved 2016-01-20.
  4. ^ Cope, Edward D. (1969). "Synopsis of the extinct batrachia, reptilia and aves of North America". American Philosophical Society. 14.
  5. ^ Huxley, T.H. (1870). "On the classification of the Dinosauria with observations on the Dinosauria of the Trias". Quarterly Journal of the Geological Society of London. 26 (1–2): 32–51. doi:10.1144/gsl.jgs.1870.026.01-02.09.
  6. ^ a b Marsh, Othniel C. (1881). "Classification of the Dinosauria". Geological Society of London, Quartly Journal. 26: 80–85.
  7. ^ Marsh, Othniel C. (1885). "Names of extinct reptiles" (PDF). American Journal of Science. 29: 169.
  8. ^ Butler, Richard J.; Galton, Peter M. (August 2008). "The 'dermal armour' of the ornithopod dinosaur Hypsilophodon from the Wealden (Early Cretaceous: Barremian) of the Isle of Wight: a reappraisal". Cretaceous Research. 29 (4): 636–642. doi:10.1016/j.cretres.2008.02.002.
  9. ^ Calvo, J. O.; Porfiri, J. D.; Novas, F. E. (2007). "Discovery of a new ornithopod dinosaur from the Portezuelo Formation (Upper Cretaceous), Neuquén, Patagonia, Argentina". Arquivos do Museu Nacional. 65 (4): 471–483.
  10. ^ Weishampel, D. B. & Heinrich, R. E. (1992). "Systematics of Hypsilophodontidae and basal Iguanodontia (Dinosauria: Ornithopoda)". Historical Biology. 6 (3): 159–184. doi:10.1080/10292389209380426.
  11. ^ Ibiricu, L. M.; Martínez, R. D.; Lamanna, M. C.; Casal, G. A.; Luna, M.; Harris, J. D. and Lacovara, K. J. (2010). "A Medium-Sized Ornithopod (Dinosauria: Ornithischia) from the Upper Cretaceous Bajo Barreal Formation of Lago Colhué Huapi, Southern Chubut Province, Argentina." Annals of Carnegie Museum, 79(1): 39-50. doi:10.2992/007.079.0103
  12. ^ Madzia, Daniel; Boyd, Clint A.; Mazuch, Martin (2017). "A basal ornithopod dinosaur from the Cenomanian of the Czech Republic". Journal of Systematic Palaeontology. 16 (11): 967–979. doi:10.1080/14772019.2017.1371258.

External links

Aralosaurini

Aralosaurini is a tribe of basal lambeosaurine hadrosaurs endemic to Eurasia. It currently contains Aralosaurus (from the Aral sea of Kazakhstan) and Canardia (from Toulouse, Southern France).

Bonapartesaurus

Bonapartesaurus is an extinct genus of herbivorous ornithopod dinosaur belonging to Hadrosauridae, which lived in the area of the modern Argentina during the Campanian and Maastrichtian stages of the Late Cretaceous.

Brachylophosaurini

Brachylophosaurini is a tribe of saurolophine hadrosaurs with known material being from N. America and potentially Asia. It contains at least four taxa; Acristavus (from Montana and Utah), Brachylophosaurus (from Montana and Alberta), Maiasaura (also from Montana), and Probrachylophosaurus (also from Montana). A hadrosaur from the Amur river, Wulagasaurus, might be a member of this tribe, but this is disputed. The group was defined by Terry A. Gates and colleagues in 2011.The clade Brachylophosaurini was defined as "Hadrosaurine ornithopods more closely related to Brachylophosaurus, Maiasaura, or Acristavus than to Gryposaurus or Saurolophus".

Cerapoda

Cerapoda ("ceratopsians and ornithopods") is a clade of the dinosaur order Ornithischia.

Choyrodon

Choyrodon is a genus of hadrosauroid dinosaur from the Early Cretaceous Albian-age Khuren Dukh Formation of Mongolia. The type and only species is Choyrodon barsboldi. The generic name is derived from the city of Choyr, and -odon, from Greek for tooth; the specific name barsboldi honours paleontologist Rinchen Barsbold. The material consists of a holotype partial skull and cervical ribs, with two other partial skulls both with associated postcranial material. It was found to be the sister taxon of Eolambia.

Delapparentia

Delapparentia is a genus of iguanodont dinosaur from the Early Cretaceous period of Galve, Teruel Province, Spain. It may be a synonym of Iguanodon bernissartensis.

Draconyx

Draconyx (meaning "dragon claw") is a genus of dinosaur from the Late Jurassic. It was an ornithopod which lived in what is now Portugal and was a herbivore. It was found in the Lourinhã Formation in 1991, and described by Octávio Mateus and Miguel Telles Antunes in 2001.

Dryosauridae

Dryosaurids were primitive iguanodonts. They are known from Middle Jurassic to Early Cretaceous rocks of Africa, Europe, and North America.

Edmontosaurini

Edmontosaurini are a tribe of saurolophine hadrosaurs that lived in the Northern Hemisphere during the Late Cretaceous period. It currently contains Edmontosaurus (from the United States and Canada), Ugrunaaluk (from Alaska, U.S.), and Shantungosaurus (from Shandong, China), though Anatosaurus might be a distinct genus. Kerberosaurus and Kundurosaurus from Russia could also be members though are more likely saurolophins.

Gideonmantellia

Gideonmantellia is an extinct genus of basal ornithopod dinosaur known from the Early Cretaceous (Barremian stage) of Galve, Province of Teruel, Spain. It contains a single species, Gideonmantellia amosanjuanae.

Iguanodontia

Iguanodontia (the iguanodonts) is a clade of herbivorous dinosaurs that lived from the Middle Jurassic to Late Cretaceous. Some members include Camptosaurus, Dryosaurus, Iguanodon, Tenontosaurus, and the hadrosaurids or "duck-billed dinosaurs". Iguanodontians were one of the first groups of dinosaurs to be found. They are among the best known of the dinosaurs, and were among the most diverse and widespread herbivorous dinosaur groups of the Cretaceous period.

Jeyawati

Jeyawati is a genus of hadrosauroid dinosaur which lived during the Turonian stage of the Late Cretaceous. The type species, J. rugoculus, was described in 2010, based on fossils recovered in the U.S. state of New Mexico.The holotype, MSM P4166, was discovered in the Moreno Hill Formation. A cladistic analysis indicates that Jeyawati was more plesiomorphic (ancestral) than Shuangmiaosaurus, Telmatosaurus, and Bactrosaurus, but more derived (less like the common ancestor) than Eolambia, Probactrosaurus, and Protohadros.

Koshisaurus

Koshisaurus is a monospecific genus of basal hadrosauroid from the Kitadani Formation in Japan. The discovery of the genus suggests that hadrosauroids had higher diversity along the eastern margin of Asia in the Early Cretaceous. "Koshi" means an old Japanese regional name including Fukui prefecture where fossils of the genus were discovered.

Lapampasaurus

Lapampasaurus is an extinct genus of hadrosaurid known from the Late Cretaceous Allen Formation (late Campanian or early Maastrichtian stage) of La Pampa Province, Argentina. It contains a single species, Lapampasaurus cholinoi.The generic name refers to the Argentine province of La Pampa. The specific name honours the late collector José Cholino. The material includes cervical, dorsal, sacral and caudal vertebrae, the forelimb girdle, and the partial hindlimb.

Neornithischia

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 historically 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 typically includes Hypsilophodon and the more derived Iguanodontia.

Notohypsilophodon

Notohypsilophodon (meaning "southern Hypsilophodon") is a genus of euornithopod dinosaur from the Late Cretaceous of Argentina. It was described as the only "hypsilophodont" known from South America, although this assessment is not universally supported, and Gasparinisaura is now believed to have been a basal euornithopod as well.

Osmakasaurus

Osmakasaurus is a genus of herbivorous iguanodontian dinosaur. It is a basal iguanodontian which lived during the lower Cretaceous period (Valanginian age) in what is now Buffalo Gap of South Dakota, United States. It is known from the Chilson Member of the Lakota Formation. This genus was named by Andrew T. McDonald in 2011 and the type species is Osmakasaurus depressus. O. depressus was previously referred to as Camptosaurus depressus, and was first described in 1909 by Charles W. Gilmore.

Tsintaosaurini

Tsintaosaurini is a tribe of basal lambeosaurine hadrosaurs native to Eurasia. It currently contains only Tsintaosaurus (from China) and Pararhabdodon (from Spain ).Koutalisaurus, also known from late Cretaceous Spain and formerly referred to Pararhabdodon

, may also be a tsintaosaurin because of its association with the latter genus; some recent work also suggests it may indeed be referrable to Pararhabdodon.

Willinakaqe

Willinakaqe is a possibly invalid genus of saurolophine hadrosaurid dinosaur described based on fossils the late Cretaceous (late Campanian-early Maastrichtian stage) of the Río Negro Province of southern Argentina.

Ornithopoda

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