Thyreophora
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Thyreophora ("shield bearers", often known simply as "armored dinosaurs") is a group of
armored Armour (British English) or armor (American English; see spelling differences) is a covering used to protect an object, individual, or vehicle from physical injury or damage, especially direct contact weapons or projectiles during combat, or f ...
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 that lived from the Early
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
until the end of 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 ...
. Thyreophorans are characterized by the presence of body armor lined up in longitudinal rows along the body. Primitive forms had simple, low, keeled scutes or
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amp ...
s, whereas more derived forms developed more elaborate structures including spikes and plates. Most thyreophorans were
herbivorous 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 ...
and had relatively small brains for their body size. Thyreophora includes various subgroups, including the suborders
Ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
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, ...
. In both the suborders, the forelimbs were much shorter than the hindlimbs, particularly in stegosaurs. The
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, ...
has been defined as the group consisting of all species more closely related to ''
Ankylosaurus ''Ankylosaurus'' is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of the n ...
'' than to ''
Triceratops ''Triceratops'' ( ; ) is a genus of herbivore, herbivorous Chasmosaurinae, chasmosaurine Ceratopsidae, ceratopsid dinosaur that first appeared during the late Maastrichtian stage of the Late Cretaceous Period (geology), period, about 68 m ...
''. Thyreophora is the
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
of
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 ...
within
Genasauria Genasauria is a clade of extinct beaked, primarily herbivorous dinosaurs. Paleontologist Paul Sereno first named Genasauria in 1986. The name Genasauria is derived from the Latin word ''gena'' meaning ‘cheek’ and the Greek word ''saúra'' ( ...
.


Groups of thyreophorans


Basal thyreophorans

Basal thyreophorans form a
grade Grade most commonly refers to: * Grade (education), a measurement of a student's performance * Grade, the number of the year a student has reached in a given educational stage * Grade (slope), the steepness of a slope Grade or grading may also ref ...
leading to Ankylosauria and Stegosauria, or are instead sister to Ankylosauria with Stegosauria being more basal than either of them. These were small-to-medium size dinosaurs with small, primitive plates. Some of them are thought to have walked bipedally. The majority of these, such as ''
Scelidosaurus ''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 ...
'', ''
Scutellosaurus ''Scutellosaurus'' ( ) is a genus of thyreophoran ornithischian dinosaur that lived approximately 196 million years ago during the early part of the Jurassic Period in what is now Arizona, USA. It is classified in Thyreophora, the armoured di ...
'', ''
Emausaurus ''Emausaurus'' is a genus of thyreophoran or armored dinosaur from the Early Jurassic (Early Toarcian). Its fossils have been found in Mecklenburg-Vorpommern, northern Germany. ''Emausaurus'' is the only known Toarcian thyreophoran, as well as th ...
'', and ''
Yuxisaurus ''Yuxisaurus'' (meaning "Yuxi lizard") is an extinct genus of basal thyreophoran dinosaur from the Early Jurassic (Sinemurian-Toarcian) Fengjiahe Formation of southwestern China. The type species is ''Yuxisaurus kopchicki.'' Discovery and naming ...
'' are known from the Northern Hemisphere, in
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 ...
,
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
and
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
.


Ankylosauria

Among the Ankylosauria, the two main groups are the
Euankylosauria Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
(containing
ankylosaurid Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known Ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
s and
nodosaurid Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia. Description Nodosaurids, like their close relatives the ankylosaurids, we ...
s) and the
Parankylosauria Parankylosauria is a group of basal ankylosaurian dinosaurs known from the Cretaceous of South America, Antarctica, and Australia. It is thought the group split from other ankylosaurs during the mid-Jurassic period, despite this being unpreserved ...
. Ankylosaurids are one of the two families of Euankylosauria. They are noted by the presence of a large tail club composed of distended
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e that have fused into a single mass. They were heavy-set and heavily armored from head to tail in bony armor, even down to minor features such as the eyelids. Spikes and nodules, often of
horn Horn most often refers to: *Horn (acoustic), a conical or bell shaped aperture used to guide sound ** Horn (instrument), collective name for tube-shaped wind musical instruments *Horn (anatomy), a pointed, bony projection on the head of various ...
, were set into the armor. The head was flat, stocky, with little or no "neck", roughly shovel-shaped and characterized by two spikes on either side of the head approximately where the ears and cheeks were. ''
Euoplocephalus ''Euoplocephalus'' ( ) is a genus of very large, herbivorous ankylosaurid dinosaurs, living during the Late Cretaceous of Canada. It has only one named species, ''Euoplocephalus tutus''. The first fossil of ''Euoplocephalus'' was found in 1897 i ...
tutus'' is perhaps the best-known ankylosaurid. Nodosaurids, the other family in the Euankylosauria, may actually include the ancestors of the ankylosaurids. They lived during the middle
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
(approx 170 mya) on up through the late Cretaceous (66 mya) and, while armored as the ankylosaurids, did not have a tail club. Instead, the bony bumps and spikes that covered the rest of their body continued out to the tail and/or were augmented with sharp spines. Two examples of nodosaurs are ''
Sauropelta ''Sauropelta'' ( ; meaning 'lizard shield') is a genus of nodosaurid dinosaur that existed in the Early Cretaceous Period of North America. One species (''S. edwardsorum'') has been named although others may have existed. Anatomically, ''Saurop ...
'' and ''
Edmontonia ''Edmontonia'' is a genus of panoplosaurin nodosaurid dinosaur from the Late Cretaceous Period. It is part of the Nodosauridae, a family within Ankylosauria. It is named after the Edmonton Formation (now the Horseshoe Canyon Formation in Ca ...
'', the latter most notable for its formidable forward-pointing shoulder spikes. The Parankylosauria are a far more basal group of ankylosaurs recognized as a distinct group in 2021. They may have diverged from euankylosaurs during the mid-Jurassic. Unlike the euankylosaurians, these had a
Gondwanan Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stage ...
distribution, being known from southern
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, and
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
. They retain more basal traits such as longer and more slender limbs, but the most distinctive trait are their tail weapons or macuahuitls (named after the weapon of the same name), which consist of a flat array of osteoderms that form a fan-like structure on the underside of the tail. This structure is similar to but distinct from the thagomizers of stegosaurians and the tail clubs of ankylosaurids. Macahuitls are completely known from '' Stegouros'' and possibly from fragmentary remains in ''
Antarctopelta ''Antarctopelta'' ( ; meaning 'Antarctic shield') was a genus of ankylosaurian dinosaur with one known species, ''A. oliveroi'', which lived in Antarctica during the Late Cretaceous Period. It was a medium-sized ankylosaur, reaching 4 meters (13& ...
''.


Stegosauria

The suborder Stegosauria comprises
Stegosauridae Stegosauridae is a family of thyreophoran dinosaurs (armoured dinosaurs) within the suborder Stegosauria. The clade is defined as all species of dinosaurs more closely related to ''Stegosaurus'' than '' Huayangosaurus''.David B. Weishampel, Pete ...
and Huayangosauridae. These dinosaurs lived mostly from the Middle to Late
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
, although some fossils have been found 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 ...
. Stegosaurs had very small heads with simple, leaf-like teeth. Stegosaurs possessed rows of plates and/or spikes running down the dorsal midline and elongated dorsal vertebra. It has been suggested that stegosaur plates functioned in control of body temperature (
thermoregulation Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
) and/or were used as a display to identify members of a species, as well as to attract mates and intimidate rivals. Well known stegosaurs are ''
Stegosaurus ''Stegosaurus'' (; ) is a genus of herbivorous, four-legged, armored dinosaur from the Late Jurassic, characterized by the distinctive kite-shaped upright plates along their backs and spikes on their tails. Fossils of the genus have been foun ...
'' and ''
Kentrosaurus ''Kentrosaurus'' ( ; ) is a genus of stegosaurid dinosaur from the Late Jurassic in Lindi Region of Tanzania. The type species is ''K. aethiopicus'', named and described by German palaeontologist Edwin Hennig in 1915. Often thought to be a " pri ...
''.


Classification


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.


Phylogeny

Thyreophora was first named by Nopcsa in 1915. Thyreophora was defined as 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, ...
by
Paul Sereno Paul Callistus Sereno (born October 11, 1957) is a professor of paleontology at the University of Chicago and a National Geographic "explorer-in-residence" who has discovered several new dinosaur species on several continents, including at sites ...
in 1998, as "all genasaurs more closely related to ''
Ankylosaurus ''Ankylosaurus'' is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of the n ...
'' than to ''
Triceratops ''Triceratops'' ( ; ) is a genus of herbivore, herbivorous Chasmosaurinae, chasmosaurine Ceratopsidae, ceratopsid dinosaur that first appeared during the late Maastrichtian stage of the Late Cretaceous Period (geology), period, about 68 m ...
''". Thyreophoroidea was first named by Nopcsa in 1928 and defined by Sereno in 1986, as "''
Scelidosaurus ''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 ...
'', ''
Ankylosaurus ''Ankylosaurus'' is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of the n ...
'', their most recent common ancestor and all of its descendants". Eurypoda was first named by Sereno in 1986 and defined by him in 1998, as "''
Stegosaurus ''Stegosaurus'' (; ) is a genus of herbivorous, four-legged, armored dinosaur from the Late Jurassic, characterized by the distinctive kite-shaped upright plates along their backs and spikes on their tails. Fossils of the genus have been foun ...
'', ''Ankylosaurus'', their most recent common ancestor and all of their descendants". The following
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 ...
shows the
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
of Riguetti ''et al'' (2022); it incorporates the ankylosaurian taxonomy of Soto-Acuña ''et al''. (2021). In 2020, as part of his monograph on ''
Scelidosaurus ''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 ...
'', David Norman revised the relationships of early thyreophorans, finding that Stegosauria was the most basal branch, with ''
Scutellosaurus ''Scutellosaurus'' ( ) is a genus of thyreophoran ornithischian dinosaur that lived approximately 196 million years ago during the early part of the Jurassic Period in what is now Arizona, USA. It is classified in Thyreophora, the armoured di ...
'', ''
Emausaurus ''Emausaurus'' is a genus of thyreophoran or armored dinosaur from the Early Jurassic (Early Toarcian). Its fossils have been found in Mecklenburg-Vorpommern, northern Germany. ''Emausaurus'' is the only known Toarcian thyreophoran, as well as th ...
'' and ''Scelidosaurus'' being progressive stem groups to Ankylosauria, rather than to Stegosauria+Ankylosauria. A cladogram is given below:


See also

* ''
thyreophoroi The ''thyreophoroi'' or ''thureophoroi'' ( el, θυρεοφόροι; singular: ''thureophoros''/''thyreophoros'', θυρεοφόρος) were a type of infantry soldier, common in the 3rd to 1st centuries BC, who carried a large oval shield called a ...
'', Greek soldiers bearing a
thyreos A thyreos ( grc, θυρεός) was a large oval shield which was commonly used in Hellenistic armies from the 3rd century BC onwards. It was adopted from the Galatians, probably first by the Illyrians, then by the Thracians before becoming common ...
shield


References

{{Taxonbar, from=Q823931 Taxa named by Franz Nopcsa von Felső-Szilvás Hettangian first appearances Maastrichtian extinctions