Shanxia Temporal range: Late Cretaceous, | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | † Ornithischia |
Clade: | † Thyreophora |
Clade: | † Ankylosauria |
Family: | † Ankylosauridae |
Subfamily: | † Ankylosaurinae |
Genus: | † Shanxia Barrett et al., 1998 |
Species: | †S. tianzhenensis |
Binomial name | |
†Shanxia tianzhenensis Barrett et al., 1998 | |
Shanxia (named after the Shanxi Province) is a monospecific genus of ankylosaurid dinosaur from the Shanxi Province that lived during the Late Cretaceous (Cenomanian-Campanian, ~99-71 Ma) in what is now the Huiquanpu Formation. [1] Shanxia may possibly represent a junior synonym of Tianzhenosaurus , an ankylosaurine also known from the Huiquanpu Formation of China. [2]
In 1993, Messrs. Lu and Li of the Hebei Geological Survey discovered a partial ankylosaurid skeleton at a locality in the Shanxi Province and notified staff of the Institute of Vertebrate Paleontology and Paleoanthropology, who collected the material later that year. [1] The holotype specimen, IVPP V11276, consists of a braincase, occiput, skull roof, ?quadratojugal, axis vertebra, cervical vertebrae, dorsal vertebrae, caudal vertebrae, humerus, fragment of ?ilium, femora and an osteoderm. [1] The holotype skull is partially reconstructed with plaster to stabilise parts of the skull roof and to attach the skull roof to the braincase. [3] Upchurch & Barrett, 2000 noted that the skull had not been artificially widened or lengthened by excess plaster. [3] The specimen was specifically obtained from the Huiquanpu Formation and is currently housed at the Institute of Vertebrate Paleontology and Paleoanthropology, Beijing. [1]
The generic name, Shanxia, refers to the Shanxi Province which is where the holotype was recovered from. [1] The specific name, tianzhenensis, refers to the Tian Zhen County. [1]
In 1999, Sullivans considered Shanxia as a nomen dubium and as a possible junior syonynm of Tianzhenosaurus. [2] Sullivan noted that the only diagnosable characteristic, the unique shape of the squamosal horn, of Shanxia is known to be variable within a single taxon such as Euoplocephalus . [2] Sullivan also noted that due to the presence of two ankylosaurids in the Huiquanpu Formation, they were likely to represent the same taxon. [2] However, Upchurch & Barrett, 2000 considered Shanxia as a valid taxon based on at least on autapomorphy or diagnostic trait. [3] Thompson et al., 2012 also recovered Shanxia as a valid taxon based on the caudal projection of the squamosal horns, form of cranial armour and haemal arch attachment. [4] Arbour & Currie, 2015 considered that there are no unique characteristics that distinguish Shanxia from Tianzhenosaurus, and Tianzhenosaurus from Saichania , an ankylosaurid from the Barun Goyot and Nemegt Formation. [5]
Due to the fragmentary nature of the holotype specimen, not much information is known about the exact size and appearance of Shanxia. [1]
Barrett et al., 1998 distinguished Shanxia from all other ankylosaurids based on the morphology of the squamosal horns. [1] It was noted that the squamosal horns were slender and elongate, inclined at an angle, having a narrow junction with the skull roof in occipital view, and shaped like isosceles triangles in dorsal view. [1] An isolated quadratojugal was tentatively assigned to Shanxia and shares similar features to Euoplocephalus . [1] The quadratojugal was described as sub-triangular in shape and generally similar to the quadratojugal horns of other ankylosaurids. [1] Upchurch and Barrett, 2000 established two more additional distinguishing characteristics of Shanxia that differentiate it from other ankylosaurids, including Saichania, which include the absence of a large processus medialis humeri and a pair of large domed sub-rectangular osteoderms on the skull. [3] However, similar ossifications are present in Tianzhenosaurus. [3] The holotype specimen only preserves one osteoderm, which has been described as oval in outline, sub-triangular in cross-section, excavated ventrally and has a prominent dorsal keel. [1]
Barrett et al., 1998 noted that the holotype specimen preserves a possible synapomorphy with nodosaurids, the hemispherical occipital condyle is oval to subcircular in posterior view and offset from the braincase by a distinct 'neck'. [1] However, they did state that some caution is needed in the interpretation of the synapomorphy as the shape of the occipital condyle is not entirely clear and may be primitive for Ankylosauria as the feature is also present in Tarchia , Talarurus , and Maleevus . [1]
Barrett et al. (1998) originally considered Shanxia as an ankylosaurid with unknown affiliations, but Sullivan (1999) considered Shanxia as a nomen dubium, and possibly synonymous with the ankylosaurid Tianzhenosaurus, arguing that the unique shape of the squamosal horns could be a product of individual variation. [1] [2] However, Upchurch and Barrett (2000) reaffirmed the validity of Shanxia and recognised two new distinguishing characteristics. [3] Thompson et al. (2012) recovered Shanxia as an ankylosaurine, sister taxon to “Crichtonsaurus” benxiensis and more basal than Tianzhenosaurus , while Arbour and Currie (2015) recovered Shanxia as a junior synonym of Tianzhenosaurus. [4] [5] A 2018 study by Wiersma and Irmis considered Shanxia as a valid taxon, and sister taxon to Tarchia kielanae and Minotaurasaurus ramachandrani . [6]
A phylogenetic analysis conducted by Thompson et al., 2012 is reproduced below. [4]
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The holotype specimen of Shanxia was recovered from the Huiquanpu Formation, which possibly dates to the Cenomanian or Campanian stage of the Late Cretaceous. [1] Due to the absence of index fossils, a precise dating is problematic. The proposed Cretaceous date is based in part on the discovery of pliosaur fossils. [1] The formation represents a fluvial environment, with the upper part of the formation being extensively reworked by a number of early Tertiary volcanic intrusions. [1] Shanxia would have coexisted with the sauropod Huabeisaurus , [7] the ornithopod Datonglong , [8] the tyrannosauroid Jinbeisaurus [9] and the ankylosaurid Tianzhenosaurus. [10]
Euoplocephalus is a genus of large herbivorous ankylosaurid dinosaurs, living during the Late Cretaceous of Canada. It has only one named species, Euoplocephalus tutus.
Crichtonsaurus is a genus of herbivorous ankylosaurid dinosaur that lived during the Late Cretaceous in what is now China. It was named after Michael Crichton, the author of the dinosaur novel Jurassic Park. A sister taxon was discovered, C. benxiensis, which is now identified as a separate genus.
Pinacosaurus is a genus of ankylosaurid thyreophoran dinosaur that lived in Asia during the Late Cretaceous, mainly in Mongolia and China.
Saichania is a genus of herbivorous ankylosaurid dinosaur from the Late Cretaceous period of Mongolia and China.
Anodontosaurus is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the entire span of the Late Cretaceous Horseshoe Canyon Formation of southern Alberta, Canada, and is also known from the Dinosaur Park Formation. It contains two species, A. lambei and A. inceptus.
Scolosaurus is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the lower levels of the Dinosaur Park Formation and upper levels of the Oldman Formation in the Late Cretaceous of Alberta, Canada. It contains two species, S. cutleri and S. thronus. The type species, S. cutleri, measured up to 5.6 metres (18 ft) in length and 2.2 metric tons in body mass.
Shamosaurus is an extinct genus of herbivorous basal ankylosaurid ankylosaur from Early Cretaceous deposits of Höövör, Mongolia.
Texasetes is a genus of ankylosaurian dinosaurs from the late Lower Cretaceous of North America. This poorly known genus has been recovered from the Paw Paw Formation near Haslet, Tarrant County, Texas, which has also produced the nodosaurid ankylosaur Pawpawsaurus.
Cedarpelta is an extinct genus of basal ankylosaurid dinosaur from Utah that lived during the Late Cretaceous period in what is now the Mussentuchit Member of the Cedar Mountain Formation. The type and only species, Cedarpelta bilbeyhallorum, is known from multiple specimens including partial skulls and postcranial material. It was named in 2001 by Kenneth Carpenter, James Kirkland, Don Burge, and John Bird. Cedarpelta has an estimated length of 7 metres and weight of 5 tonnes (11,023 lbs). The skull of Cedarpelta lacks extensive cranial ornamentation and is one of the only known ankylosaurs with individual skull bones that are not completely fused together.
Mymoorapelta is a nodosaurid ankylosaur from the Late Jurassic Morrison Formation of western Colorado and central Utah, USA. The animal is known from a single species, Mymoorapelta maysi, and few specimens are known. The most complete specimen is the holotype individual from the Mygatt-Moore Quarry, which includes osteoderms, a partial skull, vertebrae, and other bones. It was initially described by James Kirkland and Kenneth Carpenter in 1994. Along with Gargoyleosaurus, it is one of the earliest known nodosaurids.
Tianzhenosaurus is a monospecific genus of ankylosaurid dinosaur from the Shanxi Province that lived during the Late Cretaceous in what is now the Huiquanpu Formation. Tianzhenosaurus may represent a junior synonym of Saichania, an ankylosaurine known from the Barun Goyot and Nemegt Formation.
Tarchia is a genus of herbivorous ankylosaurid dinosaur from the late Cretaceous of Mongolia.
Talarurus is a genus of ankylosaurid dinosaur that lived in Asia during the Late Cretaceous period, about 96 million to 89 million years ago. The first remains of Talarurus were discovered in 1948 and later described by the Russian paleontologist Evgeny Maleev with the type species T. plicatospineus. It is known from multiple yet sparse specimens, making it one of the most well known ankylosaurines, along with Pinacosaurus. Elements from the specimens consists of various bones from the body; five skulls have been discovered and assigned to the genus, although the first two were very fragmented.
Nodocephalosaurus is a monospecific genus of ankylosaurid dinosaur from New Mexico that lived during the Late Cretaceous in what is now the De-na-zin member of the Kirtland Formation. The type and only species, Nodocephalosaurus kirtlandensis, is known only from a partial skull. It was named in 1999 by Robert M. Sullivan. Nodocephalosaurus has an estimated length of 4.5 metres and weight of 1.5 tonnes. It is closely related and shares similar cranial anatomy to Akainacephalus.
Heishansaurus, meaning "Heishan lizard" after the area in China where it was discovered, is the name given to a dubious genus of herbivorous ornithischian dinosaur, probably belonging to the Ankylosauridae.
Minotaurasaurus is a monospecific genus of ankylosaurid dinosaur that lived in Mongolia during the Late Cretaceous in what is now the Djadochta Formation. The type and only species, Minotaurasaurus ramachandrani, is known from two skulls, a cervical vertebra and a cervical half ring. It was named and described in 2009 by Clifford Miles and Clark Miles. The first fossils of Minotaurasaurus were illegally exported out of Mongolia.It has been suggested to be a synonym of Tarchia but more recent publications consider it as a distinct genus.
Oohkotokia is a genus of ankylosaurid dinosaur within the subfamily Ankylosaurinae. It is known from the upper levels of the Two Medicine Formation of Montana, United States. The discovery of Oohkotokia supports that Ankylosaurine dinosaurs existed and flourished continuously in Montana and/or Alberta throughout the late Campanian and early Maastrichtian stages in the Late Cretaceous period. It was a large, heavily built, quadrupedal, herbivore, that could grow up to 5 metres (16 ft) long and weigh up to 2 metric tons.
Ziapelta is an extinct genus of ankylosaurid. Its fossils have been found in the Hunter Wash and De-na-zin members of the Kirtland Formation of Upper Cretaceous (Campanian) New Mexico. It was named in 2014, in a research paper led by ankylosaur researcher Victoria Arbour. There is a single species in the genus, Ziapelta sanjuanensis. The genus is named after the Zia sun symbol, a stylized sun with four groups of rays, having religious significance to the Zia people of New Mexico, and the iconic symbol on the state flag of New Mexico, and pelta (Latin), a small shield, in reference to the osteoderms found on all ankylosaurids. The specific name is in reference to San Juan County and the San Juan basin, where the fossils were found. Multiple specimens have been described to date, though the fossils are mostly from the front part of the animal. Its closest relative appears to be either Scolosaurus or Nodocephalosaurus, depending on what cladistic model is used.
Zaraapelta is an extinct genus of herbivorous ankylosaurid thyreophoran dinosaur from the Late Cretaceous of Mongolia. The type species is Zaraapelta nomadis, named and described by Arbour et al in 2014. Zaraapelta is known from a single skull from the Barun Goyot Formation. It was found to be closest to Tarchia in the phylogenetic analysis within its description.
Akainacephalus is a monospecific genus of ankylosaurid dinosaur from southern Utah that lived during the Late Cretaceous in what is now the Horse Mountain Gryposaur Quarry of the Kaiparowits Formation. The type and only species, Akainacephalus johnsoni, is known from the most complete ankylosaur specimen ever discovered from southern Laramidia, including a complete skull, tail club, a number of osteoderms, limb elements and part of its pelvis, among other remains. It was described in 2018 by Jelle P. Wiersma and Randall B. Irmis. It is closely related and shares similar cranial anatomy to Nodocephalosaurus.