Tarchia Temporal range: Late Cretaceous, | |
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A cast of specimen PIN 3142/250, the holotype of T. teresae. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | † Ornithischia |
Clade: | † Thyreophora |
Clade: | † Ankylosauria |
Family: | † Ankylosauridae |
Subfamily: | † Ankylosaurinae |
Genus: | † Tarchia Maryanska, 1977 |
Type species | |
†Tarchia kielanae Maryanska, 1977 | |
Other species | |
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Synonyms | |
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Tarchia (meaning "brainy one") is a genus of herbivorous ankylosaurid dinosaur from the late Cretaceous of Mongolia.
In 1970, a Polish-Mongolian expedition discovered an ankylosaurian skull near Khulsan. In 1977, Teresa Maryańska named and described the type species Tarchia kielanae. The generic name is derived from Mongolian тархи (tarkhi, "brain") and Latin ~ia, in reference to a brain size presumed larger than that of the related form Saichania . The specific name honours Professor Zofia Kielan-Jaworowska, the leader of the expedition.
The holotype, ZPal MgD-I/111, was discovered in the Upper Cretaceous (possibly Campanian-Maastrichtian) Barun Goyot Formation (previously known as the 'Lower Nemegt Beds') of the Nemegt Basin of Mongolia. It consists of a skull roof, braincase and rear skull elements. [1] Maryańska referred three additional specimens: ZPAL MgDI/43, a large postcranial skeleton containing three "free" tail vertebrae, twelve tail vertebrae of the "handle" of the tail club and a scute; ZPAL MgDI/49, a right humerus; and PIN 3142/251, a skeleton with skull, that as yet remains undescribed.
Tarchia is the geologically youngest of all known Asian ankylosaurid dinosaurs. In 1977, Tatyana Tumanova named a second species: Tarchia gigantea. This was a renaming of Dyoplosaurus giganteus Maleev 1956, which had been based on specimen PIN 551/29. [2] In 1987, Tumanova concluded that both species were identical. This would make Dyoplosaurus giganteus the senior synonym of Tarchia kielanae. [3] This was generally accepted and Tarchia gigantea became the usual species name, as a combinatio nova replacing Tarchia kielanae. However, recent study by Victoria Megan Arbour indicates that D. giganteus is indistinguishable from other ankylosaurs from the late Campanian-Maastrichtian of Mongolia, and hence a nomen dubium ; the study revived the name Tarchia kielanae. [4]
A rump with tail and club, specimen ZPAL MgD I/113, once referred to Dyoplosaurus giganteus and subsequently to Tarchia gigantea, was by Arbour seen as different from the D. giganteus holotype. [5] The study by Arbour also concluded that specimen PIN 3142/250, in 1977 referred to Tarchia by Tumanova, probably belonged to Saichania instead. This would radically change the common image of Tarchia as this exemplar had been by far the best preserved and most illustrations, museum mounts and indeed scientific research had been based on it. Arbour discovered that the holotype of Tarchia shared distinguishing traits with that of Minotaurasaurus Miles & Miles 2009, concluding that the latter is a junior synonym of Tarchia. [6]
Subsequently, in 2016, a study conducted by Paul Penkalski and Tatiana Tumanova indicated that PIN 3142/250 is not referable to Saichania due to significant anatomical differences, but instead represents a new species of Tarchia, T. teresae. The study also recognized Minotaurasaurus as a distinct genus. [7] In 2021, Jin-Young Park and team named a new species of Tarchia, T. tumanovae, known from the holotype MPC-D 100/1353 which consists of a partial skeleton with associated skull. It was found in the Nemegt Formation at the Hermiin Tsav locality, making it coeval with T. teresae. [8]
Tarchia was a medium-sized ankylosaur, measuring around 5.5–6 metres (18–20 ft) long and weighing up to 2.5–3 metric tons (2.8–3.3 short tons). [9] [10] If ZPAL MgD I/113 indeed belongs to the genus, it would have belonged to an individual measuring 5.8–6.7 metres (19–22 ft) long. [11]
As an ankylosaurid, Tarchia would have had a broad, low-slung body, positioned on strong short legs. The body would have been protected by skin ossifications, named osteoderms. It probably had a bony tail club, for active defence against predators.
Tarchia had previously been distinguished from Saichania on the basis of its relatively larger basicranium, an unfused paroccipital process-quadrate contact and, based on PIN 3142/250, the fact that the premaxillary rostrum is wider than the maximum distance between the tooth rows in the maxillaries. In 2014, Arbour reported two distinguishing traits apart from those known exclusively from the holotype of Minotaurasaurus; the back of the head is visible in top view; and a deep groove runs along the front and outer side of the squamosal horn, and at the front it surrounds around an accessory osteoderm placed on the rear supraorbital, forming a deep furrow. [4]
The 2016 redescription of Tarchia notes that it differs from Saichania in having a postorbital fossa (which separates the squamosal horn from the supraorbital) and an accessory osteoderm; the occiput being visible in dorsal view; the large, deep braincase; the foramen magnum being higher than it is wide; and the nuchal osteoderms being taller laterally than medially. In addition, it differs from both Saichania and Minotaurasaurus in that it lacks postocular caputegulae (or small, polygonal bony plates behind the orbit) and has a proportionally high occiput in caudal view. [7] The study additionally found that PIN 3142/250 (i.e. T. teresae) can be distinguished from T. kielanae in that the accessory osteoderm is not fused to the roof of the skull, the quadrate and paroccipital process are not fused, the back of the skull roof is strongly sculptured, and the openings for the fourth to twelfth cranial nerves is bifurcated. [7]
Much information given about Tarchia in older work refers to PIN 3142/250 (which was briefly referred to Saichania until it was named as T. teresae in 2016). In 2001, it was stated that, in Tarchia, wear facets indicative of tooth-to-tooth occlusion are present; [12] this likely does not refer to the holotype specimen, since in the holotype no teeth are preserved.
Vickaryous et al. in 2004 stated that Tarchia was basal to two distinct clades of Late Cretaceous ankylosaurids: one comprising North American taxa ( Ankylosaurus , Euoplocephalus ) and the other comprising Asian taxa ( Pinacosaurus spp., Saichania , Tianzhenosaurus , Talarurus ). [13] However, this was again based on PIN 3142/250, the characters of which usually defined the operational taxonomic unit named Tarchia in the various cladistic analyses. Remarkably, Tarchia and Saichania nevertheless in these analyses often occupied very different positions.
The following cladogram is based on a 2015 phylogenetic analysis of the Ankylosaurinae conducted by Arbour and Currie: [14]
A limited phylogenetic analysis conducted in the 2016 redescription of Tarchia, focusing on the interrelationships between Tarchia, Saichania, and Minotaurasaurus, is reproduced below. [7]
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The rocks in which Tarchia fossils were found likely represent eolian dunes and interdune environments, with small intermittent lakes and seasonal streams. [8]
Tarchia was, like other Mongolian ankylosaurines, herbivorous and a low-level bulk feeder based on its sub-rectangular broad muzzle. [8] Instead of oral processing, ankylosaurids living in dry environments such as Tarchia may have relied more on hindgut fermentation for digestion or, alternatively, consumed succulent plants that did not require complex chewing. These ankylosaurids may have also been restricted to simple orthal pulping and might have had to deal with more grit during feeding compared to ankylosaurs that lived in tropical to subtropical climates, as indicated by the microwear pits. [15] Park et al. (2021) suggested that there was a shift from bulk feeding to selective feeding in Mongolian ankylosaurines during the Campanian and Maastrichtian stages which may have either been caused by the change in habitat, as the climate changed from semi-arid and arid to humid, or interspecific competition with saurolophine hadrosaurids that immigrated from North America to Central Asia during the Campanian stage. [8]
One skull of Tarchia shows tooth marks identified as belonging to the tyrannosaurid, Tarbosaurus, indicating the theropod hunted the ankylosaurid. [16] [17]
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 non-avian dinosaurs. It was named by Barnum Brown in 1908; it is monotypic, containing only A. magniventris. The generic name means "fused" or "bent lizard", and the specific name means "great belly". A handful of specimens have been excavated to date, but a complete skeleton has not been discovered. Though other members of Ankylosauria are represented by more extensive fossil material, Ankylosaurus is often considered the archetypal member of its group, despite having some unusual features.
Euoplocephalus is a genus of large herbivorous ankylosaurid dinosaurs, living during the Late Cretaceous of Canada. It has only one named species, Euoplocephalus tutus.
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.
Maleevus is an extinct genus of herbivorous ankylosaurid dinosaur from the late Cretaceous, around 90 million years ago, of Mongolia.
Bagaceratops is a genus of small protoceratopsid dinosaurs that lived in Asia during the Late Cretaceous, around 72 to 71 million years ago. Bagaceratops remains have been reported from the Barun Goyot Formation and Bayan Mandahu Formation. One specimen may argue the possible presence of Bagaceratops in the Djadochta Formation.
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.
Dyoplosaurus is a monospecific genus of ankylosaurid dinosaur from Alberta that lived during the Late Cretaceous in what is now the Dinosaur Park Formation. Dyoplosaurus represents a close relative of Scolosaurus and Anodontosaurus, two ankylosaurids known from the Horseshoe Canyon and Dinosaur Park Formation.
Tylocephale is a genus of pachycephalosaurid dinosaur, a group of dome-headed, herbivorous ornithischians, that lived during the Late Campanian stage of the Late Cretaceous in what is now Mongolia. It is known from a partial skull and associated mandible that were unearthed in 1971 by a Polish-Mongolian Expedition to the Barun Goyot Formation of the Gobi Desert. The specimen was described in 1974 by Polish paleontologists Teresa Maryańska and Halszka Osmólska as a new genus and species.
Shamosaurus is an extinct genus of herbivorous basal ankylosaurid ankylosaur from Early Cretaceous deposits of Höövör, Mongolia.
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.
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.
Shanxia is a monospecific genus of ankylosaurid dinosaur from the Shanxi Province that lived during the Late Cretaceous in what is now the Huiquanpu Formation. Shanxia may possibly represent a junior synonym of Tianzhenosaurus, an ankylosaurine also known from the Huiquanpu Formation of China.
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.
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.
This timeline of ankylosaur research is a chronological listing of events in the history of paleontology focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spikes and sometimes by a clubbed tail. Although formally trained scientists did not begin documenting ankylosaur fossils until the early 19th century, Native Americans had a long history of contact with these remains, which were generally interpreted through a mythological lens. The Delaware people have stories about smoking the bones of ancient monsters in a magic ritual to have wishes granted and ankylosaur fossils are among the local fossils that may have been used like this. The Native Americans of the modern southwestern United States tell stories about an armored monster named Yeitso that may have been influenced by local ankylosaur fossils. Likewise, ankylosaur remains are among the dinosaur bones found along the Red Deer River of Alberta, Canada where the Piegan people believe that the Grandfather of the Buffalo once lived.
Khulsanurus is an extinct genus of alvarezsaurid theropod dinosaur from the Late Cretaceous Barungoyot Formation of the Khulsan Locality in the Gobi Desert region of Mongolia. The type and only species is Khulsanurus magnificus.
Ondogurvel is a genus of alvarezsaurid dinosaur from the Late Cretaceous (Campanian) Barun Goyot Formation in southern Mongolia. The type and only species is O. alifanovi, known from a partial skeleton consisting of fragments of two last dorsal vertebrae, three anterior sacral vertebrae, right ilium, left and right pubis and ischium, articulated right tibia, fibula, metatarsals II and IV, and phalanges IV-1 and IV-2, right carpometacarpus, left and right manual phalanx II-1, right femur, left pedal phalanx II-1, and fragments of unidentified phalanges.
Jaculinykus is an extinct genus of alvarezsaurid theropod dinosaur from the Late Cretaceous Baruungoyot Formation of Mongolia. The genus contains a single species, J. yaruui, known from a nearly complete articulated skeleton including bones of the skull. Jaculinykus is notable for its unique hand, which has a hypertrophied first digit and greatly reduced second digit, which is intermediate between the tridactyl hand of Shuvuuia and monodactyl hand of Linhenykus.