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{{featured article}} {{featured article}}
{{speciesbox {{speciesbox
| fossil_range = ], {{fossilrange|145.5|136.4}} | fossil_range = <br>], {{fossilrange|earliest=151|145.5|136.4|latest=132}}<small>Possible ] record</small>
| image = Paranthodon.png | image = Paranthodon.png
| image_alt = Skull of Paranthodon | image_alt = Skull of Paranthodon
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In identifying the remains as those of ''Palaeoscincus'', Broom initially classified ''Paranthodon'' as an ], a statement backed by the research of Coombs in the 1970s. However, in 1929, Nopcsa identified the taxon as a ], with which most modern studies agree. In 1981, the genus was reviewed with modern taxonomy, and found to be a valid genus of stegosaurid. However, a more recent 2008 review of Stegosauria could only identify one distinguishing feature, and while that study still referred it to Stegosauridae based on similarity, no diagnostic features of the family could be identified in ''Paranthodon''. In identifying the remains as those of ''Palaeoscincus'', Broom initially classified ''Paranthodon'' as an ], a statement backed by the research of Coombs in the 1970s. However, in 1929, Nopcsa identified the taxon as a ], with which most modern studies agree. In 1981, the genus was reviewed with modern taxonomy, and found to be a valid genus of stegosaurid. However, a more recent 2008 review of Stegosauria could only identify one distinguishing feature, and while that study still referred it to Stegosauridae based on similarity, no diagnostic features of the family could be identified in ''Paranthodon''.


==History of discovery==
==Description==
]
''Paranthodon'' was a small relative of larger stegosaurids such as ''Stegosaurus''. ] estimated that the animal was {{convert|16.4|ft|abbr=on|disp=flip}} long and weighed between {{convert|454|and|907|kg|abbr=on}}.<ref name="holtz2007"/><ref name="holtzsuplementary"/> The snout is elongated, though not extremely so, and ] on top. The back of the premaxilla is long and broad, and the external ] are large. The teeth have a prominent primary ridge. The fossilized nasal and maxillary bones are relatively complete, and an incomplete premaxilla is also preserved. The partial snout resembles ''Stegosaurus'' in its large ] premaxillary process and the extension of the ]. However, ''Stegosaurus'' is the only stegosaurid known from adequate cranial material to compare with ''Paranthodon'', and even though their resemblance is great, tooth ] is very distinguishing among the stegosaurians. For example, cranial material is only known from ''Stegosaurus'', ''Paranthodon'', ''Kentrosaurus'', and ''Tuojiangosaurus'', and the tooth morphology differs in all of them.<ref name="galton&coombs1981"/>

The maxilla of ''Paranthodon'' preserves the ], and shows that there is little to no overhang. This differs from ankylosaurians, where there is a large amount of overhang of the maxilla. The teeth also have a middle ridge, with five fewer prominent ridges on either side. This is similar to the size ridges seen on ''Kentrosaurus''.<ref name="PMGCratero"/> Like all stegosaurians, the ] on the teeth are rounded at the tips, in contrast to ankylosaurians. Also, like ''Huayangosaurus'', but unlike ''Kentrosaurus'' and ''Stegosaurus'', ''Paranthodon'' possesses a prominent buccal margination (a ridge beside the tooth row). ''Paranthodon'' teeth preserve wear, but wear is absent on most teeth, similar to '']'', meaning it is likely that ''Paranthodon'' lacked ] between teeth.<ref name="pmb01"/> ''Paranthodon'' is distinguished from other stegosaurs by a long, wide, posterior process of the premaxilla, teeth in the maxilla with a very large ], and large ridges on the ]s.<ref name="glut97"/> Not all of these features were considered valid in a 2008 review of Stegosauria, with the only ] found being the possession of a partial second bony ] on the maxilla.<ref name="maidment08"/>

==Discovery and naming==
] ]
In ], amateur geologists ] and ] discovered several fossils near ], ], in the ] Valley.<ref name="atherstone" /> This was the first dinosaur find in all of the Southern Hemisphere and Africa.<ref name="durand"/> In 1849 and 1853, Bain sent some of the fossils to ] for identification. Among them was an upper jaw Bain referred to as the "Cape '']''", so the site was named "Iguanodonhoek". Atherstone published a short paper about the discovery in 1857,<ref name="atherstone"/> but lamented in 1871 that it had thus far received no attention in London. Only in ] were a series of specimens from the collection named by Owen as ''Anthodon serrarius'', basing the generic name on the resemblance of the teeth to a flower.<ref name="owen1876"/> The partial ] skull BMNH 47337, the left jaw BMNH 47338, the ] BMNH 47338 including bone fragments and impressions of the ] skull, and the ] BMNH 47337a were all assigned to ''Anthodon.''<ref name="galton&coombs1981"/> In 1882, ] assigned ''Anthodon'' to ] based on BMNH 47338, and in 1890, ] found that although ''Anthodon'' was a ], its teeth were similar to those of the Stegosauridae.<ref name="galton&coombs1981"/> In ], amateur geologists ] and ] discovered several fossils near ], ], in the ] Valley.<ref name="atherstone" /> This was the first dinosaur find in all of the Southern Hemisphere and Africa.<ref name="durand"/> In 1849 and 1853, Bain sent some of the fossils to ] for identification. Among them was an upper jaw Bain referred to as the "Cape '']''", so the site was named "Iguanodonhoek". Atherstone published a short paper about the discovery in 1857,<ref name="atherstone"/> but lamented in 1871 that it had thus far received no attention in London. Only in ] were a series of specimens from the collection named by Owen as ''Anthodon serrarius'', basing the generic name on the resemblance of the teeth to a flower.<ref name="owen1876"/> The partial ] skull BMNH 47337, the left jaw BMNH 47338, the ] BMNH 47338 including bone fragments and impressions of the ] skull, and the ] BMNH 47337a were all assigned to ''Anthodon.''<ref name="galton&coombs1981"/> In 1882, ] assigned ''Anthodon'' to ] based on BMNH 47338, and in 1890, ] found that although ''Anthodon'' was a ], its teeth were similar to those of the Stegosauridae.<ref name="galton&coombs1981"/>


In ], ] visited the collection of the ]. He concluded that Owen had mixed the partial distorted skull, teeth, and a ] of a pareiasaur and a partial upper jaw of a dinosaur, BMNH 47338, which were actually from two different species.<ref name="broom1912"/><ref name="owen1876"/> Broom kept the name ''Anthodon'' for the pareiasaur, but identified the other fossil as a member of the genus '']'', naming the new species ''Paleoscincus africanus'' in ]. He found that the anatomy of the teeth were quite different, even though they resembled each other, as well as those of '']''.<ref name="broom1912"/> In ], Franz Nopcsa, unaware of Broom's previous publication, provided a new name for Broom's ''P. africanus'', as ] believed that the jaw should be differentiated from ''Anthodon''. Nopcsa named the species ''Paranthodon Owenii'', with the generic name derived from the ] ''para'', meaning "similar", "near", or "beside", and ''Anthodon'', and specific name honoring Owen.<ref name="nopcsa1929"/><ref name="holtz2007" /><ref name="glut97"/> Due to present conventions, the specific name was later emended to ''owenii''.<ref name="galton&coombs1981"/> Both names were brought into the current nomenclature by ] in his 1971 dissertation as the new combination ''Paranthodon africanus'', as the name ''Paranthodon'' was the first new generic name for the fossils and ''africanus'' was the first new specific name.<ref name="olshevsky1978"/> This makes ''Paranthodon africanus'' the proper name for the taxon previously known as ''Palaeoscincus africanus'' and ''Paranthodon owenii''.<ref name="galton&coombs1981"/> In ], ] visited the collection of the ]. He concluded that Owen had mixed the partial distorted skull, teeth, and a ] of a pareiasaur and a partial upper jaw of a dinosaur, BMNH 47338, which were actually from two different species.<ref name="broom1912"/><ref name="owen1876"/> Broom kept the name ''Anthodon'' for the pareiasaur, but identified the other fossil as a member of the genus '']'', naming the new species ''Paleoscincus africanus'' in ]. He found that the anatomy of the teeth were quite different, even though they resembled each other, as well as those of '']''.<ref name="broom1912"/> In ], Franz Nopcsa, unaware of Broom's previous publication, provided a new name for Broom's ''P. africanus'', as ] believed that the jaw should be differentiated from ''Anthodon''. Nopcsa named the species ''Paranthodon Owenii'', with the generic name derived from the ] ''para'', meaning "similar", "near", or "beside", and ''Anthodon'', and specific name honoring Owen.<ref name="nopcsa1929"/><ref name="holtz2007" /><ref name="glut97"/> Due to present conventions, the specific name was later emended to ''owenii''.<ref name="galton&coombs1981"/> Both names were brought into the current nomenclature by Walter P. Coombs in his 1971 dissertation as the new combination ''Paranthodon africanus'', as the name ''Paranthodon'' was the first new generic name for the fossils and ''africanus'' was the first new specific name.<ref name="olshevsky1978"/> This makes ''Paranthodon africanus'' the proper name for the taxon previously known as ''Palaeoscincus africanus'' and ''Paranthodon owenii''.<ref name="galton&coombs1981"/>


===Material===
The holotype of ''Paranthodon'', BMNH 47338, was found in a layer of the ] that has been dated between the ] and early ] ages. It consists of the back of the snout, containing the ] with teeth, the posterior caudodorsal ramus of the ], part of the ], and some isolated teeth probably from the lower jaw. One additional specimen can be assigned to it based on the ], BMNH 47992, including only isolated teeth sharing the same morphology as those from the holotype.<ref name="galton&coombs1981"/> However, the teeth do not bear any autapomorphies of ''Paranthodon'', and were referred to an indeterminate stegosaurid in 2008.<ref name="maidment08"/> The holotype of ''Paranthodon'', BMNH 47338, was found in a layer of the ] that has been dated between the ] and early ] ages. It consists of the back of the snout, containing the ] with teeth, the posterior caudodorsal ramus of the ], part of the ], and some isolated teeth probably from the lower jaw. One additional specimen can be assigned to it based on the ], BMNH 47992, including only isolated teeth sharing the same morphology as those from the holotype.<ref name="galton&coombs1981"/> However, the teeth do not bear any autapomorphies of ''Paranthodon'', and were referred to an indeterminate stegosaurid in 2008.<ref name="maidment08"/>

The ] of ] was screened in the 1990s by the ], and in it were discovered multiple dinosaur teeth, pertaining to many groups of taxa. The locality has been described as "the largest and most complete record of dinosaur fossils from a Late Jurassic African locality outside of ]". Two of the partial teeth discovered were referred to ''Paranthodon'' by Lee Hall and Mark Goodwin in ]. However, the reasons for the referral to ''Paranthodon'' were not discussed.<ref name=hall11/>

==Description==
]
''Paranthodon'' was a small relative of larger stegosaurids such as ''Stegosaurus''. ] estimated that the animal was {{convert|16.4|ft|abbr=on|disp=flip}} long and weighed between {{convert|454|and|907|kg|abbr=on}}.<ref name="holtz2007"/><ref name="holtzsuplementary"/> The snout is elongated, though not extremely so, and ] on top. The back of the premaxilla is long and broad, and the external ] are large. The teeth have a prominent primary ridge. The fossilized nasal and maxillary bones are relatively complete, and an incomplete premaxilla is also preserved. The partial snout resembles ''Stegosaurus'' in its large ] premaxillary process and the extension of the ]. However, ''Stegosaurus'' is the only stegosaurid known from adequate cranial material to compare with ''Paranthodon'', and even though their resemblance is great, tooth ] is very distinguishing among the stegosaurians. For example, cranial material is only known from ''Stegosaurus'', ''Paranthodon'', ''Kentrosaurus'', and ''Tuojiangosaurus'', and the tooth morphology differs in all of them.<ref name="galton&coombs1981"/>

The maxilla of ''Paranthodon'' preserves the ], and shows that there is little to no overhang. This differs from ankylosaurians, where there is a large amount of overhang of the maxilla. The teeth also have a middle ridge, with five fewer prominent ridges on either side. This is similar to the size ridges seen on ''Kentrosaurus''.<ref name="PMGCratero"/> Like all stegosaurians, the ] on the teeth are rounded at the tips, in contrast to ankylosaurians. Also, like ''Huayangosaurus'', but unlike ''Kentrosaurus'' and ''Stegosaurus'', ''Paranthodon'' possesses a prominent buccal margination (a ridge beside the tooth row). ''Paranthodon'' teeth preserve wear, but wear is absent on most teeth, similar to '']'', meaning it is likely that ''Paranthodon'' lacked ] between teeth.<ref name="pmb01"/> ''Paranthodon'' is distinguished from other stegosaurs by a long, wide, posterior process of the premaxilla, teeth in the maxilla with a very large ], and large ridges on the ]s.<ref name="glut97"/> Not all of these features were considered valid in a 2008 review of Stegosauria, with the only ] found being the possession of a partial second bony ] on the maxilla.<ref name="maidment08"/>


==Classification== ==Classification==
Line 69: Line 72:
The Kirkwood Formation is in South Africa, and many fossils of different species and genera have been discovered in it, with ''Paranthodon'' being the first uncovered.<ref name="forsteretal2009"/> The formation is of a ] to ] age, with the oldest deposits from the ], about 145.5 million years ago, and the youngest rocks being from the ], about 136.4 million years ago.<ref name="holtz2007"/><ref name="glut97"/><ref name="psetal03"/> A large variety of different ] groups have been found in the formation, including dinosaurs, at least two different ]n lizards, multiple ] fishes, a few ]s, some ] specimens, and also ]. However, a large amount of the material of the Kirkwood formation only includes isolated teeth or partial and fragmentary pieces of bone. Dinosaurs of the formation include a basal ], the primitive ]n '']'', the possible ] '']'', a potential ]n, many ]ns, ''Paranthodon'', a genus of ]n, and a "]" (the family Hypsilophodontidae is no longer considered to be a natural grouping<ref name="brown2013"/>).<ref name="forsteretal2009"/><ref name="albanymus"/> The Kirkwood Formation is in South Africa, and many fossils of different species and genera have been discovered in it, with ''Paranthodon'' being the first uncovered.<ref name="forsteretal2009"/> The formation is of a ] to ] age, with the oldest deposits from the ], about 145.5 million years ago, and the youngest rocks being from the ], about 136.4 million years ago.<ref name="holtz2007"/><ref name="glut97"/><ref name="psetal03"/> A large variety of different ] groups have been found in the formation, including dinosaurs, at least two different ]n lizards, multiple ] fishes, a few ]s, some ] specimens, and also ]. However, a large amount of the material of the Kirkwood formation only includes isolated teeth or partial and fragmentary pieces of bone. Dinosaurs of the formation include a basal ], the primitive ]n '']'', the possible ] '']'', a potential ]n, many ]ns, ''Paranthodon'', a genus of ]n, and a "]" (the family Hypsilophodontidae is no longer considered to be a natural grouping<ref name="brown2013"/>).<ref name="forsteretal2009"/><ref name="albanymus"/>


If the referral of teeth from Ethiopia to ''Paranthodon'' is correct, then the taxons geographic range is extended significantly. The Mugher locality is approimately 151 million years old, about 14 million older than has previously been suggested for ''Paranthodon'', as well as across both southern and eastern Africa. The fauna in the Mugher locality differ from elsewhere of the same time and place in Africa. While the Tendaguru has abundant stegosaurs, ]s, ]s and ]s, the Mugher Mudstone preserves the stegosaur ''Paranthodon'', a hypsilophodontid ornithopod, a probably sauropod, and theropods referred to ] and ].<ref name=hall11/>
==See also==
*]
{{Portal|Dinosaurs}}


==References== ==References==
Line 96: Line 97:
<ref name="forsteretal2009">{{cite journal|last=Forster|first=C.A.|last2=Farke|first2=A.A.|last3=McCartney|first3=J.A.|last4=de Klerk|first4=W.J.|last5=Ross|first5=C.F.|year=2009|title=A "Basal" Tetanuran from the Lower Cretaceous Kirkwood Formation of South Africa|url=|journal=Journal of Vertebrate Paleontology|volume=29|issue=1|pages=283–285|doi=10.1671/039.029.0101|jstor=20491088}}</ref> <ref name="forsteretal2009">{{cite journal|last=Forster|first=C.A.|last2=Farke|first2=A.A.|last3=McCartney|first3=J.A.|last4=de Klerk|first4=W.J.|last5=Ross|first5=C.F.|year=2009|title=A "Basal" Tetanuran from the Lower Cretaceous Kirkwood Formation of South Africa|url=|journal=Journal of Vertebrate Paleontology|volume=29|issue=1|pages=283–285|doi=10.1671/039.029.0101|jstor=20491088}}</ref>
<ref name="albanymus">{{cite book|last1=Chinsamy|first1=A.|chapter = Albany Museum, Grahamstown, South Africa|editor1-last=Currie|editor1-first=P.J.|editor2-last=Padian|editor2-first=K.|title=Encyclopedia of Dinosaurs|date=1997|publisher=Academic Press|isbn=978-0-12-226810-6|page=6}}</ref> <ref name="albanymus">{{cite book|last1=Chinsamy|first1=A.|chapter = Albany Museum, Grahamstown, South Africa|editor1-last=Currie|editor1-first=P.J.|editor2-last=Padian|editor2-first=K.|title=Encyclopedia of Dinosaurs|date=1997|publisher=Academic Press|isbn=978-0-12-226810-6|page=6}}</ref>
<ref name=hall11>{{cite journal|last=Hall|first=L.E.|last2=Goodwin|first2=M.B.|year=2011|title=A diverse dinosaur tooth assemblage from the Upper Jurassic of Ethiopia: implications for Gondwanan dinosaur biogeography|url=https://www.researchgate.net/profile/Lee_Hall4/publication/291344526_A_DIVERSE_DINOSAUR_TOOTH_ASSEMBLAGE_FROM_THE_UPPER_JURASSIC_OF_ETHIOPIA_IMPLICATIONS_FOR_GONDWANAN_DINOSAUR_BIOGEOGRAPHY/links/56a1047d08ae2afab882814a/A-DIVERSE-DINOSAUR-TOOTH-ASSEMBLAGE-FROM-THE-UPPER-JURASSIC-OF-ETHIOPIA-IMPLICATIONS-FOR-GONDWANAN-DINOSAUR-BIOGEOGRAPHY.pdf|journal=Las Vegas, Nevada, Society of Vertebrate Paleontology Annual Meeting|doi=10.13140/RG.2.1.3740.7763}}</ref>
</references> </references>

==See also==
{{Portal|Dinosaurs}}
*]


{{Thyreophora|S.}} {{Thyreophora|S.}}

Revision as of 16:10, 13 December 2017

Paranthodon
Temporal range:
Early Cretaceous, 145.5–136.4 Ma PreꞒ O S D C P T J K Pg N Possible Late Jurassic record
Skull of Paranthodon
Reconstruction of the skull; grey material is unknown.
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Ornithischia
Clade: Thyreophora
Clade: Stegosauria
Family: Stegosauridae
Genus: Paranthodon
Nopcsa, 1929
Species: P. africanus
Binomial name
Paranthodon africanus
Broom, 1912
Synonyms

Palaeoscincus africanus Broom, 1912
Paranthodon owenii Nopcsa, 1929

Paranthodon (/pəˈrænθədɒn/ pə-RAN-thə-don) is a genus of extinct stegosaurian dinosaur that lived in South Africa during the Early Cretaceous, between 145.5 and 136.4 million years ago. Discovered in 1845, it was one of the first stegosaurians found. Its only remains, a partial skull and isolated teeth, 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.

In identifying the remains as those of Palaeoscincus, Broom initially classified Paranthodon as an ankylosaurian, a statement backed by the research of Coombs in the 1970s. However, in 1929, Nopcsa identified the taxon as a stegosaurid, with which most modern studies agree. In 1981, the genus was reviewed with modern taxonomy, and found to be a valid genus of stegosaurid. However, a more recent 2008 review of Stegosauria could only identify one distinguishing feature, and while that study still referred it to Stegosauridae based on similarity, no diagnostic features of the family could be identified in Paranthodon.

History of discovery

Fossils of Anthodon, what Paranthodon was once thought to be
Skull of Anthodon serrarius, to which the skull of Paranthodon was originally assigned

In 1845, amateur geologists William Guybon Atherstone and Andrew Geddes Bain discovered several fossils near Dassieklip, Cape Province, in the Bushmans River Valley. This was the first dinosaur find in all of the Southern Hemisphere and Africa. In 1849 and 1853, Bain sent some of the fossils to Richard Owen for identification. Among them was an upper jaw Bain referred to as the "Cape Iguanodon", so the site was named "Iguanodonhoek". Atherstone published a short paper about the discovery in 1857, but lamented in 1871 that it had thus far received no attention in London. Only in 1876 were a series of specimens from the collection named by Owen as Anthodon serrarius, basing the generic name on the resemblance of the teeth to a flower. The partial holotype skull BMNH 47337, the left jaw BMNH 47338, the matrix BMNH 47338 including bone fragments and impressions of the anterior skull, and the vertebrae BMNH 47337a were all assigned to Anthodon. In 1882, Othniel Charles Marsh assigned Anthodon to Stegosauridae based on BMNH 47338, and in 1890, Richard Lydekker found that although Anthodon was a pareiasaur, its teeth were similar to those of the Stegosauridae.

In 1909, Robert Broom visited the collection of the British Museum of Natural History. He concluded that Owen had mixed the partial distorted skull, teeth, and a mandible of a pareiasaur and a partial upper jaw of a dinosaur, BMNH 47338, which were actually from two different species. Broom kept the name Anthodon for the pareiasaur, but identified the other fossil as a member of the genus Palaeoscincus, naming the new species Paleoscincus africanus in 1912. He found that the anatomy of the teeth were quite different, even though they resembled each other, as well as those of Stegosaurus. In 1929, Franz Nopcsa, unaware of Broom's previous publication, provided a new name for Broom's P. africanus, as D.M.S. Watson believed that the jaw should be differentiated from Anthodon. Nopcsa named the species Paranthodon Owenii, with the generic name derived from the Latin para, meaning "similar", "near", or "beside", and Anthodon, and specific name honoring Owen. Due to present conventions, the specific name was later emended to owenii. Both names were brought into the current nomenclature by Walter P. Coombs in his 1971 dissertation as the new combination Paranthodon africanus, as the name Paranthodon was the first new generic name for the fossils and africanus was the first new specific name. This makes Paranthodon africanus the proper name for the taxon previously known as Palaeoscincus africanus and Paranthodon owenii.

Material

The holotype of Paranthodon, BMNH 47338, was found in a layer of the Kirkwood Formation that has been dated between the Berriasian and early Valanginian ages. It consists of the back of the snout, containing the maxilla with teeth, the posterior caudodorsal ramus of the premaxilla, part of the nasals, and some isolated teeth probably from the lower jaw. One additional specimen can be assigned to it based on the dentition, BMNH 47992, including only isolated teeth sharing the same morphology as those from the holotype. However, the teeth do not bear any autapomorphies of Paranthodon, and were referred to an indeterminate stegosaurid in 2008.

The Mugher Mudstone of Ethiopia was screened in the 1990s by the University of California Museum of Paleontology, and in it were discovered multiple dinosaur teeth, pertaining to many groups of taxa. The locality has been described as "the largest and most complete record of dinosaur fossils from a Late Jurassic African locality outside of Tendaguru". Two of the partial teeth discovered were referred to Paranthodon by Lee Hall and Mark Goodwin in 2011. However, the reasons for the referral to Paranthodon were not discussed.

Description

Silhouette of Paranthodon is shown to be three times longer than the human silhouette is tall
Size comparison between Paranthodon and a human

Paranthodon was a small relative of larger stegosaurids such as Stegosaurus. Thomas R. Holtz Jr. estimated that the animal was 5.0 m (16.4 ft) long and weighed between 454 and 907 kg (1,001 and 2,000 lb). The snout is elongated, though not extremely so, and convex on top. The back of the premaxilla is long and broad, and the external nares are large. The teeth have a prominent primary ridge. The fossilized nasal and maxillary bones are relatively complete, and an incomplete premaxilla is also preserved. The partial snout resembles Stegosaurus in its large posterior premaxillary process and the extension of the palate. However, Stegosaurus is the only stegosaurid known from adequate cranial material to compare with Paranthodon, and even though their resemblance is great, tooth morphology is very distinguishing among the stegosaurians. For example, cranial material is only known from Stegosaurus, Paranthodon, Kentrosaurus, and Tuojiangosaurus, and the tooth morphology differs in all of them.

The maxilla of Paranthodon preserves the tooth row, and shows that there is little to no overhang. This differs from ankylosaurians, where there is a large amount of overhang of the maxilla. The teeth also have a middle ridge, with five fewer prominent ridges on either side. This is similar to the size ridges seen on Kentrosaurus. Like all stegosaurians, the denticles on the teeth are rounded at the tips, in contrast to ankylosaurians. Also, like Huayangosaurus, but unlike Kentrosaurus and Stegosaurus, Paranthodon possesses a prominent buccal margination (a ridge beside the tooth row). Paranthodon teeth preserve wear, but wear is absent on most teeth, similar to Huayangosaurus, meaning it is likely that Paranthodon lacked occlusion between teeth. Paranthodon is distinguished from other stegosaurs by a long, wide, posterior process of the premaxilla, teeth in the maxilla with a very large cingulum, and large ridges on the tooth crowns. Not all of these features were considered valid in a 2008 review of Stegosauria, with the only autapomorphy found being the possession of a partial second bony palate on the maxilla.

Classification

Currently, Paranthodon is classified as a stegosaur related to Stegosaurus, Tuojiangosaurus, and Loricatosaurus. Initially, when Broom assigned the name Palaeoscincus africanus to the Paranthodon fossils, he classified them as an ankylosaurian. This classification was later changed by Nopcsa, who found that Paranthodon best resembled a stegosaurid (before the group was truly defined). Coombs (1978) did not follow Nopcsa's classification, keeping Paranthodon as an ankylosaurian, like Broom, although he only classified it as Ankylosauria incertae sedis. A subsequent review by Galton and Coombs in 1981 instead confirmed Nopcsa's interpretation, redescribing Paranthodon as a stegosaurid from the Late Cretaceous. Multiple phylogenetic analyses have placed Paranthodon in Stegosauria, and often in Stegosauridae. A 2010 analysis including nearly all species of stegosaurians found that Paranthodon was outside Stegosauridae, and in a polytomy with Tuojiangosaurus, Huayangosaurus, Chungkingosaurus, Jiangjunosaurus, and Gigantspinosaurus. However, when the latter two genera were removed, Paranthodon grouped with Tuojiangosaurus just outside Stegosauridae, and Huayangosaurus grouped with Chungkingosaurus in Huayangosauridae. Below is the analysis with all included genera.

A long-necked dinosaur standing on four legs. The back has numerous bony plates extending upward.
Hypothetical restoration based on related stegosaurians
Stegosauria

Gigantspinosaurus

Jiangjunosaurus

Chungkingosaurus

Huayangosaurus

Tuojiangosaurus

Paranthodon

Stegosauridae

Kentrosaurus

Loricatosaurus

Dacentrurinae

Dacentrurus

Miragaia

Stegosaurinae

Stegosaurus

Hesperosaurus

Wuerhosaurus

Other analyses have found Paranthodon closely related to Tuojiangosaurus, Loricatosaurus, and Kentrosaurus within Stegosaurinae. Even though phylogenetic analyses recognize Paranthodon as a stegosaurid, the type material actually bears no synapomorphies of Stegosauria. However, the material is likely of stegosaurian nature, and phylogenies by many authors have found it to be within the group.

Paleoecology

Map shows the city of Port Elizabeth on the coast of South Africa, with Kirkwood to the north, and the formation just west of Kirkwood.
The Kirkwood Formation, the location of the fossil discovery

The Kirkwood Formation is in South Africa, and many fossils of different species and genera have been discovered in it, with Paranthodon being the first uncovered. The formation is of a Late Jurassic to Early Cretaceous age, with the oldest deposits from the Tithonian, about 145.5 million years ago, and the youngest rocks being from the Valanginian, about 136.4 million years ago. A large variety of different animal groups have been found in the formation, including dinosaurs, at least two different sphenodontian lizards, multiple teleost fishes, a few crocodylians, some frog specimens, and also turtles. However, a large amount of the material of the Kirkwood formation only includes isolated teeth or partial and fragmentary pieces of bone. Dinosaurs of the formation include a basal tetanuran, the primitive ornithomimosaurian Nqwebasaurus, the possible brachiosaurid Algoasaurus, a potential titanosaurian, many ornithischians, Paranthodon, a genus of iguanodontian, and a "hypsilophodontid" (the family Hypsilophodontidae is no longer considered to be a natural grouping).

If the referral of teeth from Ethiopia to Paranthodon is correct, then the taxons geographic range is extended significantly. The Mugher locality is approimately 151 million years old, about 14 million older than has previously been suggested for Paranthodon, as well as across both southern and eastern Africa. The fauna in the Mugher locality differ from elsewhere of the same time and place in Africa. While the Tendaguru has abundant stegosaurs, sauropods, ornithopods and theropods, the Mugher Mudstone preserves the stegosaur Paranthodon, a hypsilophodontid ornithopod, a probably sauropod, and theropods referred to Allosauridae and Dromaeosauridae.

References

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See also

Ankylosauria
Avemetatarsalia
Ornithischia
Ankylosauria
    • see below↓
Ankylosauria
Ankylosauria
Parankylosauria
Euankylosauria
Nodosauridae
Polacanthinae
Nodosaurinae
Panoplosaurini
Struthiosaurini
Ankylosauridae
Shamosaurinae
Ankylosaurinae
Ankylosaurini
Sauropelta edwardsorum Ankylosaurus magniventris
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