Ornithopoda
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Ornithopoda () is a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of
ornithischia Ornithischia () is an extinct order 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 s ...
n
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
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s 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 th ...
world, dominating the
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
n 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
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s 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 Avian may refer to: *Birds or Aves, winged animals *Avian (given name) (russian: Авиа́н, link=no), a male forename Aviation *Avro Avian, a series of light aircraft made by Avro in the 1920s and 1930s *Avian Limited, a hang glider manufacture ...
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 are commonly regarded as continents. Ordered from largest in area to smallest, these seven ...
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 storie ...
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 ''Scelidosaurus'' (; with the intended meaning of "limb lizard", from Greek / meaning 'rib of beef' and ''sauros''/ meaning 'lizard')Liddell & Scott (1980). Greek-English Lexicon, Abridged Edition. Oxford University Press, Oxford, UK. is a gen ...
), 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, Eu ...
'', ''
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 faunal stage, Stage of the Cretaceous period, approximatel ...
''. 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 h ...
in 1881 as part of his then still ongoing investigation of the classification of Dinosauria. It was considered one of the four definite
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
of dinosaurs, the others being
Theropoda 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 ...
,
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 Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe, ...
( 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 Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interested ...
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 ''Camptonotus'' is a genus of leaf-rolling crickets in the subfamily Gryllacridinae. It includes the following species are all found in the Americas The Americas, which are sometimes collectively called America, are a landmass compris ...
'', ''
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, Eu ...
'' 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 ''Cionodon'' (meaning 'column tooth') is a dubious genus of hadrosauridJ. R. Horner, D. B. Weishampel, and C. A. Forster. 2004. Hadrosauridae. In D. B. Weishampel, H. Osmolska, and P. Dodson (eds.), ''The Dinosauria'' (2nd edition). ''Universi ...
'', 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'. I ...
''.


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 food ...
, 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 and ...
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 Bison are large bovines in the genus ''Bison'' (Greek: "wild ox" (bison)) within the tribe Bovini. Two extant and numerous extinct species are recognised. Of the two surviving species, the American bison, ''B. bison'', found only in North Ame ...
. 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 In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
. 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
sauropod 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 ...
s that they partially supplanted. The very largest, such as ''
Shantungosaurus ''Shantungosaurus'' (meaning "''Shandong Lizard''") is a genus of very large saurolophine hadrosaurid dinosaur found in the Late Cretaceous Wangshi Group of the Shandong Peninsula in China, containing a single species, ''Shantungosaurus giganteu ...
'', 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 States c ...
s/25
short ton The short ton (symbol tn) is a measurement unit equal to . It is commonly used in the United States, where it is known simply as a ton, although the term is ambiguous, the single word being variously used for short, long, and metric ton. The vari ...
s), 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 A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
), 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 ''Triceratops'' ( ; ) is a genus of herbivorous chasmosaurine ceratopsid dinosaur that first appeared during the late Maastrichtian stage of the Late Cretaceous period, about 68 million years ago in what is now North America. It is one ...
''." 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 d ...
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