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Ornithopoda () is a clade of ornithischian
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s, called ornithopods (), that started out as small,
bipedal Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
running Running is a method of terrestrial locomotion allowing humans and other animals to move rapidly on foot. Running is a type of gait characterized by an aerial phase in which all feet are above the ground (though there are exceptions). This is ...
grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
world, dominating the North American land. Their major evolutionary advantage was the progressive development of a
chewing Chewing or mastication is the process by which food is crushed and ground by teeth. It is the first step of digestion, and it increases the surface area of foods to allow a more efficient break down by enzymes. During the mastication process, th ...
apparatus that became the most sophisticated ever developed by a non-avian dinosaur, rivaling that of modern mammals such as the domestic
cow Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult ma ...
. They reached their apex of diversity and ecological dominance in the hadrosaurids (colloquially known as 'duck-bills'), before they were wiped out by the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
along with all other non- avian
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s. Members are known from all seven
continent A continent is any of several large landmasses. Generally identified by convention rather than any strict criteria, up to seven geographical regions In geography, regions, otherwise referred to as zones, lands or territories, are areas t ...
s, though they are generally rare in the Southern Hemisphere.


History of research

In 1870,
Thomas Henry Huxley Thomas Henry Huxley (4 May 1825 – 29 June 1895) was an English biologist and anthropologist specialising in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The stori ...
listed
Iguanodontidae Iguanodontidae is a family of iguanodontians belonging to Styracosterna, a derived clade within Ankylopollexia. Characterized by their elongated maxillae, they were herbivorous and typically large in size. This family exhibited locomotive dynam ...
(coined by Cope a year earlier) as one of his three families of dinosaurs (alongside
Megalosauridae Megalosauridae is a monophyletic family of carnivorous theropod dinosaurs within the group Megalosauroidea. Appearing in the Middle Jurassic, megalosaurids were among the first major radiation of large theropod dinosaurs. They were a relatively ...
and Scelidosauridae), including within it the genera ''
Iguanodon ''Iguanodon'' ( ; meaning ' iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species have been classified in the genus ''Iguanodon'', dating from the late Jurassic Period to the early Cretaceous Period of Asia, ...
'', ''
Hypsilophodon ''Hypsilophodon'' (; meaning "''Hypsilophus''-tooth") is a neornithischian dinosaur genus from the Early Cretaceous period of England. It has traditionally been considered an early member of the group Ornithopoda, but recent research has put thi ...
'', and ''
Hadrosaurus ''Hadrosaurus'' (; ) is a genus of hadrosaurid ornithopod dinosaurs that lived in North America during the Late Cretaceous Period in what is now the Woodbury Formation about 80 million to 78 million years ago. The holotype specimen was found in f ...
'', in addition to ''
Cetiosaurus ''Cetiosaurus'' () meaning 'whale lizard', from the Greek '/ meaning 'sea monster' (later, 'whale') and '/ meaning 'lizard', is a genus of herbivorous sauropod dinosaur from the Middle Jurassic Period, living about 168 million years ago in what ...
'' and tentatively ''
Stenopelix ''Stenopelix'' (meaning "narrow pelvis") 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 ag ...
''. The term Ornithopoda was erected by
Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among ...
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 Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
, and Stegosauria ( Hallopoda was considered a possible fifth). He subdivided the order into three families: Camptonotidae,
Iguanodontidae Iguanodontidae is a family of iguanodontians belonging to Styracosterna, a derived clade within Ankylopollexia. Characterized by their elongated maxillae, they were herbivorous and typically large in size. This family exhibited locomotive dynam ...
, 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 ''Hypsilophodon'' (; meaning "''Hypsilophus''-tooth") is a neornithischian dinosaur genus from the Early Cretaceous period of England. It has traditionally been considered an early member of the group Ornithopoda, but recent research has put thi ...
'' and three American taxa he named himself, '' Camptonotus'', ''
Laosaurus ''Laosaurus'' (meaning "stone or fossil lizard") is a genus of neornithischian dinosaur. The type species, ''Laosaurus celer'', was first described by O.C. Marsh in 1878 from remains from the Oxfordian-Tithonian-age Upper Jurassic Morrison Forma ...
'', and ''
Nanosaurus ''Nanosaurus'' ("small or dwarf lizard") is the name given to a genus of neornithischian dinosaur that lived about 155 to 148 million years ago, during the Late Jurassic-age. Its fossils are known from the Morrison Formation of the south-weste ...
''. ''Camptonotus'' was in 1885 renamed to ''Camptosaurus'', as the original name was pre-occupied by a cricket; the associated family followed suit, becoming Camptosauridae. In Iguanodontidae, only found in Europe, he included ''
Iguanodon ''Iguanodon'' ( ; meaning ' iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species have been classified in the genus ''Iguanodon'', dating from the late Jurassic Period to the early Cretaceous Period of Asia, ...
'' and '' Vectisaurus''. In Hadrosauridae, he included ''
Hadrosaurus ''Hadrosaurus'' (; ) is a genus of hadrosaurid ornithopod dinosaurs that lived in North America during the Late Cretaceous Period in what is now the Woodbury Formation about 80 million to 78 million years ago. The holotype specimen was found in f ...
'', '' Cionodon'', and tentatively ''
Agathaumas ''Agathaumas'' (; "great wonder") is a dubious genus of a large ceratopsid dinosaur that lived in Wyoming during the Late Cretaceous (late Maastrichtian stage, 66 million years ago). The name comes from grc, αγαν - 'much' and - 'wonder'. It ...
''.


Description

''Ornithopoda'' means "bird feet", from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''ornithos'' ("bird") and ''pous'' ("feet"); this is in reference to members’ characteristic three-
toe Toes are the digits (fingers) of the foot of a tetrapod. Animal species such as cats that walk on their toes are described as being ''digitigrade''. Humans, and other animals that walk on the soles of their feet, are described as being ''plan ...
d feet, although many early forms retained four toes. They were also characterized as lacking in body armour, not developing a horny
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for foo ...
, having an elongated pubis (that eventually extended past the ilium), and having a missing hole in the lower
jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...
(a
Mandibular fenestrae The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
). A variety of ornithopods, and related
cerapods Cerapoda ("ceratopsians" and "ornithopods") is a clade of the dinosaur order Ornithischia, that includes pachycephalosaurs, ceratopsians and ornithopods Classification Cerapoda is divided into two groups: Ornithopoda ("bird-foot") and Marginoc ...
, had thin
cartilaginous Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck a ...
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 ''Hypsilophodon'' (; meaning "''Hypsilophus''-tooth") is a neornithischian dinosaur genus from the Early Cretaceous period of England. It has traditionally been considered an early member of the group Ornithopoda, but recent research has put thi ...
'', '' Othnielosaurus'', ''
Parksosaurus ''Parksosaurus'' (meaning " William Parks's lizard") is a genus of neornithischian dinosaur from the early Maastrichtian-age Upper Cretaceous Horseshoe Canyon Formation of Alberta, Canada. It is based on most of a partially articulated skelet ...
'', ''
Talenkauen ''Talenkauen'' is a genus of basal iguanodont dinosaur from the Campanian or Maastrichtian age of the Late Cretaceous Cerro Fortaleza Formation, formerly known as the Pari Aike Formation of Patagonian Lake Viedma, in the Austral Basin of Sa ...
'', ''
Thescelosaurus ''Thescelosaurus'' ( ; ancient Greek - (''-'') meaning "godlike", "marvellous", or "wondrous" and (') "lizard") was a genus of small neornithischian dinosaur that appeared at the very end of the Late Cretaceous period in North America. It was ...
'', and '' Macrogryphosaurus'' to date. The early ornithopods were only about 1 metre (3 feet) long, but probably very fast. They had a stiff tail, like the
theropods Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally ca ...
, 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 on all fours. The taxonomy of dinosaurs previously ascribed to the ''Hypsilophodontidae'' is problematic. The group previously consisted of all non-
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-bil ...
n bipedal ornithischians, but a phylogenetic reappraisal has shown such species to be paraphyletic. As such, the hypsilophodont family is currently represented only by ''
Hypsilophodon ''Hypsilophodon'' (; meaning "''Hypsilophus''-tooth") is a neornithischian dinosaur genus from the Early Cretaceous period of England. It has traditionally been considered an early member of the group Ornithopoda, but recent research has put thi ...
''. 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 (up to 23
metric ton The tonne ( or ; symbol: t) is a unit of mass equal to 1000  kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton to distinguish it from the non-metric units of the short ton ( United State ...
s/25 short tons), but never grew much beyond 15 metres (50 feet).


Classification

Historically, most indeterminate ornithischian
biped Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
s 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 Cerapoda ("ceratopsians" and "ornithopods") is a clade of the dinosaur order Ornithischia, that includes pachycephalosaurs, ceratopsians and ornithopods Classification Cerapoda is divided into two groups: Ornithopoda ("bird-foot") and Marginoc ...
(originally named as an unranked clade), while others, such as Ibiricu ''et al.'' 2010, have retained it at its traditional ranking of suborder.


Phylogeny

In 2021, Ornithopoda was given a formal definition under the
PhyloCode The ''International Code of Phylogenetic Nomenclature'', known as the ''PhyloCode'' for short, is a formal set of rules governing phylogenetic nomenclature. Its current version is specifically designed to regulate the naming of clades, leaving the ...
: "The largest clade containing ''
Iguanodon bernissartensis ''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species have been classified in the genus ''Iguanodon'', dating from the late Jurassic Period to the early Cretaceous Period of Asia, Eu ...
'' but not ''
Pachycephalosaurus wyomingensis ''Pachycephalosaurus'' (; meaning "thick-headed lizard", from Greek ''pachys-/'' "thick", ''kephale/'' "head" and ''sauros/'' "lizard") is a genus of pachycephalosaurid dinosaurs. The type species, ''P. wyomingensis'', is the only known species, ...
'' and '' Triceratops horridus''." The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
below follows a 2017 analysis by Madzia ''et al.'':


References


External links


GEOL 104 dinosaurs: a Natural history, ornithopoda: beaks, bills & crests
by Thomas R. Holtz Jr. {{Taxonbar, from=Q467430 Bajocian first appearances Maastrichtian extinctions Taxa named by Othniel Charles Marsh