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Thyreophorans Temporal range: Early Jurassic-Late Cretaceous, | |
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Skeletal mount of Gastonia burgei , BYU Museum of Paleontology | |
Skeletal mount of Stegosaurus stenops , Natural History Museum, London | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | † Ornithischia |
Clade: | † Genasauria |
Clade: | † Thyreophora Nopcsa, 1915 |
Subgroups | |
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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.
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]
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]
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.
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]
Thyreophora | |
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.
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.
Nodosaurus is a genus of herbivorous nodosaurid ankylosaurian dinosaur from the Late Cretaceous, the fossils of which are found exclusively in the Frontier Formation in Wyoming.
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.
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.
Scelidosaurus is a genus of herbivorous armoured ornithischian dinosaur from the Jurassic of England.
Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as a monophyletic clade as the sister taxon to the Ankylosauridae, some analyses recover it as a paraphyletic grade leading to the ankylosaurids.
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.
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.
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.
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:
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.
Panoplosaurus is a genus of armoured dinosaur from the Late Cretaceous of Alberta, Canada. Few specimens of the genus are known, all from the middle Campanian of the Dinosaur Park Formation, roughly 76 to 75 million years ago. It was first discovered in 1917, and named in 1919 by Lawrence Lambe, named for its extensive armour, meaning "well-armoured lizard". Panoplosaurus has at times been considered the proper name for material otherwise referred to as Edmontonia, complicating its phylogenetic and ecological interpretations, at one point being considered to have existed across Alberta, New Mexico and Texas, with specimens in institutions from Canada and the United States. The skull and skeleton of Panoplosaurus are similar to its relatives, but have a few significant differences, such as the lumpy form of the skull osteoderms, a completely fused shoulder blade, and regularly shaped plates on its neck and body lacking prominent spines. It was a quadrupedal animal, roughly 5 m (16 ft) long and 1,600 kg (3,500 lb) in weight. The skull has a short snout, with a very domed surface, and bony plates directly covering the cheek. The neck had circular groups of plates arranged around the top surface, both the forelimb and hindlimb were about the same length, and the hand may have only included three fingers. Almost the entire surface of the body was covered in plates, osteoderms and scutes of varying sizes, ranging from large elements along the skull and neck, to smaller, round bones underneath the chin and body, to small ossicles that filled in the spaces between other, larger osteoderms.
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.
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.
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.
Susannah "Susie" Catherine Rose Maidment is a British palaeontologist at the Natural History Museum, London. She is internationally recognised for her research on ornithischian dinosaur evolution, and was awarded the 2016 Hodson Award of the Palaeontological Association and the 2017 Lyell Fund of the Geological Society of London. She was featured as a 2019 National Geographic Women of Impact.
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.
Jakapil is a genus of basal thyreophoran dinosaur from the Candeleros Formation of Argentina. The type species is Jakapil kaniukura.