Tarchia
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''Tarchia'' (meaning "brainy one") is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of herbivorous ankylosaurid
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
from the
late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
of
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million ...
.


Discovery and naming

In 1970, a Polish-Mongolian expedition discovered an ankylosaurian skull near Khulsan. In 1977, Teresa Maryańska named and described the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specim ...
''Tarchia kielanae''. The generic name is derived from Mongolian тархи (''tarkhi'', "brain") and
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
''~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 A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
, ZPal MgD-I/111, was discovered in the Upper
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
(possibly
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
-
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the inte ...
) 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. 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
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
n 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. In 1987, Tumanova concluded that both species were identical. This would make ''Dyoplosaurus giganteus'' the senior synonym of ''Tarchia kielanae''. This was generally accepted and ''Tarchia gigantea'' became the usual
species name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
, as a ''
combinatio nova ''Combinatio nova'', abbreviated ''comb. nov.'' (sometimes ''n. comb.''), is Latin for "new combination". It is used in taxonomic biology literature when a new name is introduced based on a pre-existing name. The term should not to be confused w ...
'' replacing ''Tarchia kielanae''. However, recent study by
Victoria Megan Arbour Victoria most commonly refers to: * Victoria (Australia), a state of the Commonwealth of Australia * Victoria, British Columbia, provincial capital of British Columbia, Canada * Victoria (mythology), Roman goddess of Victory * Victoria, Seyche ...
indicates that ''D. giganteus'' is indistinguishable from other ankylosaurs from the late Campanian-Maastrichtian of Mongolia, and hence a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
''; the study revived the name ''Tarchia kielanae''.Arbour, Victoria Megan, 2014. ''Systematics, evolution, and biogeography of the ankylosaurid dinosaurs.'' Ph.D thesis, University of Alberta. https://era.library.ualberta.ca/public/.../Arbour_Victoria_Spring2014.pdf 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. 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 The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
of ''Tarchia''. Subsequently, in 2016, a study conducted by Penkalski & 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. 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 The Nemegt Formation (also known as Nemegtskaya Svita) is a geological formation in the Gobi Desert of Mongolia, dating to the Late Cretaceous. The formation consists of river channel sediments and contains fossils of fish, turtles, crocodilians, ...
at the Hermiin Tsav locality, making it coeval with ''T. teresae''.


Description

Size estimates of ''Tarchia'' have been largely based on ''Dyoplosaurus giganteus'', the holotype of which is one of the largest ankylosaurian individuals known. This would make ''Tarchia'' the longest known Asian ankylosaur, with an estimated body length of .Holtz, Thomas R. Jr. (2011) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'
Winter 2010 Appendix.
/ref> Confusingly, the skull size often mentioned, with a length of and width of , was again based on specimen PIN 3142/250, a much smaller individual. The holotypes of ''Tarchia kielanae'' and ''Minotaurasaurus'' also indicate a medium size. In 2016, Gregory S. Paul had estimated a body length of 5.5 meters (18 ft) and a weight of 2.5 tonnes (2.75 short tons) suggesting that Minotaurasaurus was a juvenile of the species.Paul, G.S., 2016, ''The Princeton Field Guide to Dinosaurs 2nd Edition'', Princeton University Press p. 261 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 The base of skull, also known as the cranial base or the cranial floor, is the most inferior area of the skull. It is composed of the endocranium and the lower parts of the calvaria. Structure Structures found at the base of the skull are for ...
, an unfused paroccipital process- quadrate contact and, based on PIN 3142/250, the fact that the
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
ry 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. 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 The foramen magnum ( la, great hole) is a large, oval-shaped opening in the occipital bone of the skull. It is one of the several oval or circular openings (foramina) in the base of the skull. The spinal cord, an extension of the medulla oblon ...
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. 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. 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
facet Facets () are flat faces on geometric shapes. The organization of naturally occurring facets was key to early developments in crystallography, since they reflect the underlying symmetry of the crystal structure. Gemstones commonly have facets cut ...
s indicative of tooth-to-tooth occlusion are present; this likely does not refer to the holotype specimen, since in the holotype no teeth are preserved.


Phylogeny

Vickaryous ''et alii'' in 2004 stated that ''Tarchia'' was basal to two distinct
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
s of Late Cretaceous ankylosaurids: one comprising
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
n
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
(''
Ankylosaurus ''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 th ...
'', ''
Euoplocephalus ''Euoplocephalus'' ( ) is a genus of very large, herbivorous ankylosaurid dinosaurs, living during the Late Cretaceous of Canada. It has only one named species, ''Euoplocephalus tutus''. The first fossil of ''Euoplocephalus'' was found in 1 ...
'') and the other comprising Asian taxa ('' Pinacosaurus'' spp., '' Saichania'', ''
Tianzhenosaurus ''Tianzhenosaurus'' (meaning “Tianzhen lizard”) 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. ''Tianzh ...
'', ''
Talarurus ''Talarurus'' ( ; meaning "basket tail" or "wicker tail") 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 ...
''). 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 Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analyses. Remarkably, ''Tarchia'' and ''Saichania'' nevertheless in these analyses often occupied very different positions. The following
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
is based on a 2015
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
of the Ankylosaurinae conducted by Arbour and Currie: A limited phylogenetic analysis conducted in the 2016 redescription of ''Tarchia'', focusing on the interrelationships between ''Tarchia'', ''Saichania'', and ''Minotaurasaurus'', is reproduced below.


Palaeobiology

The rocks in which ''Tarchia'' fossils were found likely represent eolian dunes and interdune environments, with small intermittent lakes and seasonal streams. Hence, we know that ''Tarchia'' was a
desert A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...
animal. On the other hand, well-watered forest would have been present. ''Tarchia'' would have been preyed upon by '' Tarbosaurus''.


Paleopathology

One skull of ''Tarchia'' shows tooth marks identified as belonging to the tyrannosaurid, ''Tarbosaurus'', indicating the theropod hunted the ankylosaurid.Gallagher W.B., Tumanova T.A., Dodson P., Axel L., 1998, "CT scanning Asian ankylosaurs: paleopathology in a ''Tarchia'' skull", ''Journal of Vertebrate Paleontology'' 18: 44A-45A


See also

* Timeline of ankylosaur research * 2017 in archosaur paleontology


References


External links


''Tarchia'' in the Dino Directory
{{Taxonbar, from=Q135304 Ankylosaurids Late Cretaceous dinosaurs of Asia Fossils of Mongolia Maastrichtian life Fossil taxa described in 1977 Taxa named by Teresa Maryańska Ornithischian genera