Thyreophora

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Thyreophorans
Temporal range: Early Jurassic-Late Cretaceous, 200.91–66  Ma
Gastonia mount BYU 4.jpg
Skeletal mount of Gastonia burgei , BYU Museum of Paleontology
Stegosaurus (Natural History Museum, London).jpg
Skeletal mount of Stegosaurus stenops , Natural History Museum, London
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Ornithischia
Clade: Genasauria
Clade: Thyreophora
Nopcsa, 1915
Subgroups
Thyreophora distribution map.jpg
Thyreophorans distribution map

Thyreophora ("shield bearers", often known simply as "armored dinosaurs") is a group of armored ornithischian dinosaurs that lived from the Early Jurassic until the end of the Cretaceous.

Contents

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 osteoderms, whereas more derived forms developed more elaborate structures including spikes and plates. Most thyreophorans were herbivorous and had relatively small brains for their body size.

Thyreophora includes two major subgroups, Ankylosauria and Stegosauria. In both clades, the forelimbs were much shorter than the hindlimbs, particularly in stegosaurs. Thyreophora has been defined as the group consisting of all species more closely related to Ankylosaurus and Stegosaurus than to Iguanodon and Triceratops . It is the sister group of Cerapoda within Genasauria. [2]

Characteristics

Members of Thyreophora are characterised by the presence of osteoderms (bony growths within the skin), with these osteoderms having lateral keels. Characters of the skull and jaws distinctive (synapomorphic) of thyreophorans include "absence of a deep elliptic fossa along the sutural line of the nasals, presence of a wide jugal, remodeling of skull dermal bone, down-turned dentary tooth row". [3] Among primitive thyreophorans, Scutellosaurus was likely primarily bipedal, while the more quadrupedally adapted Scelidosaurus may have been bipedal for some of the time, particularly as a juvenile. Stegosaurs and ankylosaurs are thought to have been obligately quadrupedal. [4]

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. [5] Thyreophora was defined as a clade by Paul Sereno in 1998, as "all genasaurs more closely related to Ankylosaurus than to Triceratops ". Thyreophoroidea was first named by Nopcsa in 1928 and defined by Sereno in 1986, as " Scelidosaurus , Ankylosaurus , their most recent common ancestor and all of its descendants". [6] Eurypoda was first named by Sereno in 1986 and defined by him in 1998, as " Stegosaurus , Ankylosaurus, their most recent common ancestor and all of their descendants". [7]

In 2021, an international group of researchers led by Daniel Madzia registered almost all of the most commonly used ornithischian clades under the International Code of Phylogenetic Nomenclature, with the intent of standardizing their definitions. According to Madzia et al., Thyreophora defined as the largest clade containing Ankylosaurus magniventris and Stegosaurus stenops but not Iguanodon bernissartensis and Triceratops horridus. [2]

The following cladogram shows the results of the phylogenetic analysis Soto-Acuña et al. (2021). [8] In their description of Jakapil the following year, Riguetti et al modified the same matrix and found it to occupy a position as the sister taxon to the Eurypoda. [9]

Thyreophora

In 2020, as part of his monograph on Scelidosaurus , David Norman revised the relationships of early thyreophorans, finding that Stegosauria was the most basal branch, with Scutellosaurus , Emausaurus and Scelidosaurus being progressive stem groups to Ankylosauria, rather than to Stegosauria+Ankylosauria. A cladogram is given below: [10]

See also

Related Research Articles

<i>Stegosaurus</i> Thyreophoran stegosaurid dinosaur genus from Late Jurassic period

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 found in the western United States and in Portugal, where they are found in Kimmeridgian- to Tithonian-aged strata, dating to between 155 and 145 million years ago. Of the species that have been classified in the upper Morrison Formation of the western US, only three are universally recognized: S. stenops, S. ungulatus and S. sulcatus. The remains of over 80 individual animals of this genus have been found. Stegosaurus would have lived alongside dinosaurs such as Apatosaurus, Diplodocus, Camarasaurus and Allosaurus, the latter of which may have preyed on it.

<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. Ornithischians with well known anatomical adaptations include the ceratopsians or "horn-faced" dinosaurs, the pachycephalosaurs or "thick-headed" dinosaurs, the armored dinosaurs (Thyreophora) such as stegosaurs and ankylosaurs, and the ornithopods. There is strong evidence that certain groups of ornithischians lived in herds, often segregated by age group, with juveniles forming their own flocks separate from adults. Some were at least partially covered in filamentous pelts, and there is much debate over whether these filaments found in specimens of Tianyulong, Psittacosaurus, and Kulindadromeus may have been primitive feathers.

<span class="mw-page-title-main">Ankylosauria</span> Extinct order of dinosaurs

Ankylosauria is a group of herbivorous dinosaurs of the clade 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. They are known to have first appeared in the Middle Jurassic, and persisted until the end of the Cretaceous Period. The two main families of Ankylosaurs, Nodosauridae and Ankylosauridae are primarily known from the Northern Hemisphere, but the more basal Parankylosauria are known from southern Gondwana during the Cretaceous.

<i>Kentrosaurus</i> Extinct genus of dinosaurs from late Jurassic in Lindi Region, Tanzania

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 "primitive" member of the Stegosauria, several recent cladistic analyses find it as more derived than many other stegosaurs, and a close relative of Stegosaurus from the North American Morrison Formation within the Stegosauridae.

<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.

<i>Scelidosaurus</i> Genus of reptiles (fossil)

Scelidosaurus is a genus of herbivorous armoured ornithischian dinosaur from the Jurassic of England.

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

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.

<i>Paranthodon</i> Stegosaurian dinosaur genus from Early Cretaceous South Africa

Paranthodon is a genus of stegosaurian dinosaur that lived in what is now South Africa during the Early Cretaceous, between 139 and 131 million years ago. Discovered in 1845, it was one of the first stegosaurians found. Its only remains, a partial skull, isolated teeth, and fragments of vertebrae, were found in the Kirkwood Formation. British paleontologist Richard Owen initially identified the fragments as those of the pareiasaur Anthodon. After remaining untouched for years in the British Museum of Natural History, the partial skull was identified by South African paleontologist Robert Broom as belonging to a different genus; he named the specimen Palaeoscincus africanus. Several years later, Hungarian paleontologist Franz Nopcsa, unaware of Broom's new name, similarly concluded that it represented a new taxon, and named it Paranthodon owenii. Since Nopcsa's species name was assigned after Broom's, and Broom did not assign a new genus, both names are now synonyms of the current binomial, Paranthodon africanus. The genus name combines the Ancient Greek para (near) with the genus name Anthodon, to represent the initial referral of the remains.

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

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, Africa, South America and Asia. Their geographical origins are unclear; the earliest unequivocal stegosaurian, Bashanosaurus primitivus, was found in the Bathonian Shaximiao Formation of China.

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

Stegosauridae is a family of thyreophoran dinosaurs within the suborder Stegosauria. The clade is defined as all species of dinosaurs more closely related to Stegosaurus than Huayangosaurus. The name ‘Stegosauridae’ is thus a stem-based name taken from the well-represented genus – Stegosaurus. Fossil evidence of stegosaurids, dating from the Middle Jurassic through the Early Cretaceous, have been recovered from North America, Eurasia and Africa.

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

Huayangosauridae is a family of stegosaurian dinosaurs from the Jurassic of China. The group is defined as all taxa closer to the namesake genus Huayangosaurus than Stegosaurus, and was originally named as the family Huayangosaurinae by Dong Zhiming and colleagues in the description of Huayangosaurus. Huayangosaurinae was originally differentiated by the remaining taxa within Stegosauridae by the presence of teeth in the premaxilla, an antorbital fenestra, and a mandibular fenestra. Huayangosaurinae, known from the Middle Jurassic of the Shaximiao Formation, was proposed to be intermediate between Scelidosaurinae and Stegosaurinae, suggesting that the origins of stegosaurs lay in Asia. Following phylogenetic analyses, Huayangosauridae was expanded to also include the taxon Chungkingosaurus, known from specimens from younger Late Jurassic deposits of the Shaximiao Formation. Huayangosauridae is either the sister taxon to all other stegosaurs, or close to the origin of the clade, with taxa like Gigantspinosaurus or Isaberrysaura outside the Stegosauridae-Huayangosauridae split. Huayangosauridae was formally defined in 2021 by Daniel Madzia and colleagues, who used the previous definitions of all taxa closer to Huayangosaurus taibaii than Stegosaurus stenops, and chose the 2020 phylogeny of Susannah Maidment et al. to illustrate the relationships of the clade:

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

Neornithischia 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. 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.

Dracopelta is a monospecific genus of ankylosaur dinosaur from Portugal that lived during the Late Jurassic in what is now the Lourinhã Formation. The type and only species is Dracopelta zbyszewskii, which is represented by a partial skeleton including unpublished material.

<i>Emausaurus</i> Extinct genus of thyreophoran dinosaurs

Emausaurus is a genus of thyreophoran or armored dinosaur from the Early Jurassic. Its fossils have been found in Mecklenburg-Vorpommern, northern Germany. Emausaurus is the only known Toarcian thyreophoran, as well as the only dinosaur from the zone of the same age with a formal name.

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

Laquintasaura is a genus of Venezuelan ornithischian dinosaur containing only the type species Laquintasaura venezuelae. The species was the first dinosaur to have been identified from Venezuela. It is known from extensive remains, all from a singular bonebed locality which has been sampled for specimen blocks over the course of several expeditions, largely led by Marcelo R Sánchez-Villagra. A small, very primitive animal, it is known for its distinct dental anatomy and for being one of the earliest and most primitive ornithischians in the fossil record. Taxonomic uncertainty has led to conflicting theories that it is either as the base of Ornithischia or at the base of the subgroup Thyreophora. In either model, its dating to around 200 million years ago, at the start of the Jurassic, existence in equatorial latitudes, and primitive nature make it a key view into early ornithischian evolution. It's thought that Laquintasaura would've lived in groups and had a possible omnivorous diet, living on a seasonal alluvial plain and being preyed about by the contemporary Tachiraptor.

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

Spicomellus is an extinct genus of early ankylosaurian dinosaur from the El Mers III Formation (Bathonian-Callovian) of Morocco. The genus contains a single species, S. afer, known from a single rib with fused osteoderms. Spicomellus represents the oldest named ankylosaur.

<span class="mw-page-title-main">Struthiosaurini</span>

Struthiosaurini is a clade of nodosaurid ankylosaurs from the Cretaceous of Europe and North America. The group is defined as the largest clade containing Struthiosaurus austriacus, but not Nodosaurus textilis or Panoplosaurus mirus, and was named in 2021 by Madzia and colleagues for the relatively stable group found in many previous analyses. Struthiosaurini includes not only the Late Cretaceous European Struthiosaurus, but also the Early Cretaceous European Europelta, the Late Cretaceous European Hungarosaurus, and Stegopelta and Pawpawsaurus from the mid Cretaceous of North America. The approximately equivalent clade Struthiosaurinae, named in 1923 by Franz Nopcsa was previously used to include European nodosaurids, but was never formally named following the PhyloCode, so Madzia et al. named Struthiosaurini instead, as the group of taxa fell within the clade Nodosaurinae, and having the same -inae suffix on both parent and child taxon could be confusing in future. The 2018 phylogenetic analysis of Rivera-Sylva and colleagues was used as the primary reference for Struthiosaurini by Madzia et al., in addition to the supplemental analyses of Arbour et al. (2016), Brown et al. (2017), and Zheng et al. (2018).

<i>Bashanosaurus</i> Extinct genus of stegosaurian dinosaurs

Bashanosaurus is an extinct genus of stegosaurian dinosaur from the Middle Jurassic Shaximiao Formation of Yunyang County, China. The genus contains a single species, Bashanosaurus primitivus, known from incomplete skeletons belonging to three individuals. It is one of the basalmost stegosaurs, as well as one of the oldest known stegosaurs, along with Adratiklit, Isaberrysaura, and Thyreosaurus.

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

Jakapil is a genus of basal thyreophoran dinosaur from the Candeleros Formation of Argentina. The type species is Jakapil kaniukura.

References

  1. Raven, T. J.; Barrett, P. M.; Joyce, C. B.; Maidment, S. C. R. (2023). "The phylogenetic relationships and evolutionary history of the armoured dinosaurs (Ornithischia: Thyreophora)". Journal of Systematic Palaeontology. 21 (1). 2205433. doi:10.1080/14772019.2023.2205433.
  2. 1 2 Madzia, D.; Arbour, V.M.; Boyd, C.A.; Farke, A.A.; Cruzado-Caballero, P.; Evans, D.C. (2021). "The phylogenetic nomenclature of ornithischian dinosaurs". PeerJ. 9: e12362. doi: 10.7717/peerj.12362 . PMC   8667728 . PMID   34966571.
  3. Breeden, Benjamin T.; Rowe, Timothy B. (2020-07-03). "New Specimens of Scutellosaurus Lawleri Colbert, 1981, from the Lower Jurassic Kayenta Formation in Arizona Elucidate the Early Evolution of Thyreophoran Dinosaurs". Journal of Vertebrate Paleontology. 40 (4): e1791894. doi:10.1080/02724634.2020.1791894. ISSN   0272-4634.
  4. Anderson, Lilian; Brassey, Charlotte; Pond, Stuart; Bates, Karl; Sellers, William Irvin (October 2023). "Investigating the quadrupedal abilities of Scutellosaurus lawleri and its implications for locomotor behavior evolution among dinosaurs". The Anatomical Record. 306 (10): 2514–2536. doi:10.1002/ar.25189. ISSN   1932-8486. PMID   36896818.
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  9. Riguetti, Facundo J.; Apesteguía, Sebastián; Pereda-Suberbiola, Xabier (2022-08-11). "A new Cretaceous thyreophoran from Patagonia supports a South American lineage of armoured dinosaurs". Scientific Reports. 12 (1): 11621. doi:10.1038/s41598-022-15535-6. ISSN   2045-2322. PMC   9372066 . PMID   35953515.
  10. Norman, David B (2021-01-01). "Scelidosaurus harrisonii (Dinosauria: Ornithischia) from the Early Jurassic of Dorset, England: biology and phylogenetic relationships". Zoological Journal of the Linnean Society. 191 (1): 1–86. doi:10.1093/zoolinnean/zlaa061. ISSN   0024-4082.