Tongtianlong
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''Tongtianlong'' (meaning "Tongtianyan dragon") is a genus of
oviraptorid Oviraptoridae is a group of bird-like, herbivorous and omnivorous maniraptoran dinosaurs. Oviraptorids are characterized by their toothless, parrot-like beaks and, in some cases, elaborate crests. They were generally small, measuring between one ...
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s that lived in the late
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
epoch of the late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period. It contains one species, ''T. limosus''.


Description

''Tongtianlong'' was a
sheep Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to d ...
-sized member of the oviraptorids, a group of omnivorous, feathered, bird-like theropods. The describers of ''Tongtianlong'' recognized that it possessed a set of distinctive characteristics that differentiated it both from other oviraptorosaurs. In particular, unlike other oviraptorids, the crest of ''Tongtianlong'' was shaped like a dome, with its highest point just behind the eye socket; and the front edge of the toothless
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 mammals h ...
, which would have supported its beak, was very rounded. Additionally, there is a distinct ridge on the front margin of the
parietal bone The parietal bones ( ) are two bones in the skull which, when joined at a fibrous joint known as a cranial suture, form the sides and roof of the neurocranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four bord ...
, wedged between the
frontal bone In the human skull, the frontal bone or sincipital bone is an unpaired bone which consists of two portions.'' Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bo ...
s; the shaft of the
lacrimal bone The lacrimal bones are two small and fragile bones of the facial skeleton; they are roughly the size of the little fingernail and situated at the front part of the medial wall of the orbit. They each have two surfaces and four borders. Several bon ...
, which is located in front of the eye socket, is wide, flattened, and plate-like seen from the side; the
foramen magnum The foramen magnum () 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 oblongata, passes thro ...
(a hole in the back of the skull) is smaller than the
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the Atlas (anatomy), atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape ...
(the boss forming the skull-neck joint); there is no ridge on the bottom of the front lower jaws, which is also not strongly downturned; and the
xiphoid process The xiphoid process (), also referred to as the ensiform process, xiphisternum, or metasternum, constitutes a small cartilaginous process (extension) located in the inferior segment of the sternum, typically ossified in adult humans. Both the Gr ...
does not flare out from the
sternum The sternum (: sternums or sterna) or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major bl ...
behind the ribs. Other characteristics of the skull separate ''Tongtianlong'' from its contemporaries; for instance, the nostril is situated much higher than the
antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
, a trait seen otherwise only in ''
Nemegtomaia ''Nemegtomaia'' is a genus of oviraptorid dinosaur from what is now Mongolia that lived in the Late Cretaceous Period (geology), Period, about 70million years ago. The first specimen was found in 1996, and became the basis of the new genus and s ...
'' and ''
Rinchenia ''Rinchenia'' (named after Byambyn Rinchen) is a genus of oviraptorid dinosaur that lived in Asia during the Late Cretaceous epoch in what is now Mongolia, Nemegt Formation, around 70 million years ago. The type species, type and only known spec ...
''.


Discovery and naming

The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
of ''Tongtianlong'', specimen number DYM-2013-8, was discovered during the construction of a new high school near Ganxian District in
Jiangxi ; Gan: ) , translit_lang1_type2 = , translit_lang1_info2 = , translit_lang1_type3 = , translit_lang1_info3 = , image_map = Jiangxi in China (+all claims hatched).svg , mapsize = 275px , map_caption = Location ...
Province,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
. The site where it was found is part of the
Nanxiong Formation The Nanxiong Formation (also known as Yuanpu Formation) is a Late Cretaceous geologic Formation (geology), formation in Guangdong Province. Dinosaur remains are among the fossils that have been recovered from the formation. Description It cons ...
, which dates to the
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
epoch (although a more precise dating has not yet been conducted). The specimen is currently stored in the Dongyang Museum. While the exceptionally well-preserved specimen was likely originally complete, portions of the arms, right leg, and tail were destroyed by the TNT blasting which unearthed the fossil. Near the hip, a drill hole that was used to place TNT can be seen. The skeleton was preserved in an unusual pose, with a raised head and splayed arms; while it is difficult to tell exactly what led to this pose, it has been speculatively suggested that the specimen died while it was trapped in mud and trying to free itself. The genus name of ''Tongtianlong'' combines a reference to the nearby Tongtianyan ( zh, c=通天岩)
grotto A grotto or grot is a natural or artificial cave or covered recess. Naturally occurring grottoes are often small caves near water that are usually flooded or often flooded at high tide. Sometimes, artificial grottoes are used as garden fea ...
and the suffix ''-long'' ( zh, c=龍, "dragon"). "Tongtian" also is a Chinese phrase meaning "road to heaven". The specific name, ''limosus'' (the Latin word for "muddy"), refers to how the specimen was preserved in
mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from ''shale'' by its lack of fissility.Blatt, H., and R.J. Tracy, 1996, ''Petrology.'' New York, New York, ...
. It was first described and named by
Lü Junchang Lü Junchang (; 1965 – 9 October 2018) was a Chinese palaeontologist and professor at the Institute of Geology, Chinese Academy of Geological Sciences. An expert on Mesozoic reptiles, he described and named dozens of dinosaur and pterosaur taxa ...
, Chen Rongjun, Stephen L. Brusatte, ''et al''.


Classification

In 2016, ''Tongtianlong'' was found to be a member of the
Oviraptoridae Oviraptoridae is a group of bird-like, herbivorous and omnivorous maniraptoran dinosaurs. Oviraptorids are characterized by their toothless, parrot-like beaks and, in some cases, elaborate crests. They were generally small, measuring between one ...
, as a close relative of '' Banji'' and ''
Wulatelong ''Wulatelong'' is an extinct genus of basal oviraptorid dinosaur known from the Late Cretaceous Wulansuhai Formation (Campanian stage) of Bayan Mandahu, Linhe District of Inner Mongolia, northern China. It contains a single species, ''Wulatelon ...
''. The results of the
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
conducted are partially reproduced below.


Paleoecology

''Tongtianlong'' is the sixth oviraptorid described from the
Nanxiong Formation The Nanxiong Formation (also known as Yuanpu Formation) is a Late Cretaceous geologic Formation (geology), formation in Guangdong Province. Dinosaur remains are among the fossils that have been recovered from the formation. Description It cons ...
, after '' Banji'', '' Ganzhousaurus'', ''
Jiangxisaurus ''Jiangxisaurus'' is an extinct genus of oviraptorid theropod dinosaur from the Late Cretaceous Nanxiong Formation of southern China. It was similar to ''Heyuannia'', but with more strongly curved anterior claws and a thinner, frailer mandible. ...
'', ''
Nankangia ''Nankangia'' is an extinct genus of caenagnathoid oviraptorosaurian dinosaur known from the Upper Cretaceous Nanxiong Formation of Nankang, Jiangxi, Nankang County, Ganzhou City of Jiangxi Province, southeastern China. It contains a single speci ...
'', and ''
Huanansaurus ''Huanansaurus'' is an extinct genus of Oviraptoridae, oviraptorid dinosaur that lived approximately 72 million years ago, between the Campanian and Maastrichtian, during the latter part of the Cretaceous period in what is now China, in the Nanxi ...
'', in that order. Given the variation in size among these oviraptorids (for instance, ''Banji'' is much smaller than ''Tongtianlong''), it is reasonable to hypothesize that some of them might be growth stages of others; however, it is more likely that they represent genuinely distinct animals, since juvenile oviraptorid specimens show that their anatomy does not change significantly with age. The describers of ''Tongtianlong'' suggested that this diversity is indicative of an
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. A significantly large and diverse radiation within ...
of oviraptorids that occurred just prior to the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...
. However, the possibility that these species come from different points in time cannot be excluded, due to a lack of study on the stratigraphy and age of the Nanxiong Formation.


See also

*
2016 in paleontology Flora Plants Fungi Cnidarians Research * ''Yunnanoascus haikouensis'', previously thought to be a member of Ctenophora, is reinterpreted as a Crown group, crown-group medusozoan by Han ''et al.'' (2016). * A study on the fossil corals fro ...


References

{{Taxonbar, from=Q27826636 Oviraptoridae Dinosaur genera Maastrichtian dinosaurs Taxa named by Lü Junchang Taxa named by Stephen L. Brusatte Fossil taxa described in 2016 Dinosaurs of China