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''Heyuannia'' ("from
Heyuan Héyuán (, Hakka:Fò-Ngiàn) is a prefecture-level city of Guangdong province in the People's Republic of China. As of the 2020 census, its population was 2,837,686 whom 1,051,993 lived in the built-up (''or metro'') area made of Yuancheng urban ...
") 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 n ...
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 on ...
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 ...
that lived in
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 ...
during 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'', ...
epoch, in what is now China and
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 millio ...
. It was the first oviraptorid found in China; most others were found in neighbouring
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 millio ...
. Two species are known: ''H. huangi'', named by Lü Junchang in 2002 from the Dalangshan Formation; and ''H. yanshini'', originally named as a separate genus ''Ingenia'' from the
Barun Goyot Formation The Barun Goyot Formation (also known as Baruungoyot Formation or West Goyot Formation) is a geological formation dating to the Late Cretaceous Period. It is located within and is widely represented in the Gobi Desert Basin, in the Ömnögovi ...
by
Rinchen Barsbold , Rinchyengiin Barsbold, born December 21, 1935 in Ulaanbaatar) is a Mongolian paleontologist and geologist. He works with the Institute of Geology, at Ulaanbaatar, Mongolia. He is an expert in vertebrate paleontology and Mesozoic stratigraphy. Bar ...
in 1981, and later renamed to ''Ajancingenia'' in 2013 due to the preoccupation of ''Ingenia''. The latter name was eventually discarded due to various ethical issues surrounding the author.


Discovery and naming


''H. huangi''

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 specimen( ...
, ''Heyuannia huangi'', was named and described by Lü Junchang in 2002. The generic name refers to the city of Heyuan. The specific name honours Huang Dong, the director of the Heyuan Museum. 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 ...
, HYMV1-1, was discovered in
Guangdong Guangdong (, ), alternatively romanized as Canton or Kwangtung, is a coastal province in South China on the north shore of the South China Sea. The capital of the province is Guangzhou. With a population of 126.01 million (as of 2020 ...
near Huangsha in layers of the Dalangshan Formation. It consists of a partial skeleton, including the skull. Six further skeletons were assigned as
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...
s or referred to the species. Multiple other fossils have been found, including one which may retain possible reproductive organs. Thousands of eggs have also been uncovered at the site, some of them belonging to the genus and likely laid by ''Heyuannia''.


''H. yanshini''

''H. yanshini'' was first described and named by
Rinchen Barsbold , Rinchyengiin Barsbold, born December 21, 1935 in Ulaanbaatar) is a Mongolian paleontologist and geologist. He works with the Institute of Geology, at Ulaanbaatar, Mongolia. He is an expert in vertebrate paleontology and Mesozoic stratigraphy. Bar ...
in
1981 Events January * January 1 ** Greece enters the European Economic Community, predecessor of the European Union. ** Palau becomes a self-governing territory. * January 10 – Salvadoran Civil War: The FMLN launches its first major off ...
, as a new genus and species ''Ingenia yanshini''. The name "Ingenia" derives from the Ingen Khoboor Depression of
Bayankhongor Province The Bayankhongor Province or Bayanhongor Aimag, , "Rich Darling Province" is one of the 21 aimags (provinces) of Mongolia. It is located in the southwest of the country and, at 116,000 square kilometers, it is one of the largest aimags. The capit ...
(
Barun Goyot Formation The Barun Goyot Formation (also known as Baruungoyot Formation or West Goyot Formation) is a geological formation dating to the Late Cretaceous Period. It is located within and is widely represented in the Gobi Desert Basin, in the Ömnögovi ...
), Mongolia, from whence it was collected, while the specific name ''yanshini'' was chosen in honour of academician Aleksandr Leonidovich Yanshin (1911–1999), who was adviser and mentor to Rinchen Barsbold during his time at the
Paleontological Institute The Paleontological Institute, Russian Academy of Sciences (PIN; russian: Палеонтологический институт РАН) in Moscow is among the world's largest paleontological institutes. An affiliate of the Russian Academy of Scienc ...
in
St. Petersburg Saint Petersburg ( rus, links=no, Санкт-Петербург, a=Ru-Sankt Peterburg Leningrad Petrograd Piter.ogg, r=Sankt-Peterburg, p=ˈsankt pʲɪtʲɪrˈburk), formerly known as Petrograd (1914–1924) and later Leningrad (1924–1991), i ...
,
Russia Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It is the largest country in the world, with its internationally recognised territory covering , and encompassing one-eigh ...
.Barsbold, R. (1981). "Bezzubye khishchnye dinozavry Mongolii." oothless carnivorous dinosaurs of Mongolia. ''Trudy -- Sovmestnaya Sovetsko-Mongol'skaya Paleontologicheskaya Ekspeditsiya'', 15: 28-39, 124. n Russian, w/ English summary Most of the material known for this is species actually a composite of four specimens, including the holotype skull of ''
Conchoraptor ''Conchoraptor'' (meaning "conch plunderer") is a genus of oviraptorid dinosaurs that lived in Asia during the Late Cretaceous epoch, about 70 million years ago. It is known from the Barun Goyot and Nemegt formations of Mongolia. Discovery Whe ...
''. The generic name ''Ingenia'' was preoccupied by the generic name of '' Ingenia mirabilis'' (Gerlach, 1957), a tripyloidid nematode. Thus, an alternative generic name, ''Ajancingenia'', was proposed by Jesse Easter in
2013 File:2013 Events Collage V2.png, From left, clockwise: Edward Snowden becomes internationally famous for leaking classified NSA wiretapping information; Typhoon Haiyan kills over 6,000 in the Philippines and Southeast Asia; The Dhaka garment ...
. The replacement generic name is derived also from ''ajanc'' (аянч; a traveler in Mongolian), as a Western allusion to sticking one's thumb out for
hitchhiking Hitchhiking (also known as thumbing, autostop or hitching) is a means of transportation that is gained by asking individuals, usually strangers, for a ride in their car or other vehicle. The ride is usually, but not always, free. Nomads hav ...
, in reference to the first manual
ungual An ungual (from Latin ''unguis'', i.e. ''nail'') is a highly modified distal toe bone which ends in a hoof, claw, or nail. Elephants and ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammal ...
of ''Ajancingenia'' which is twice as large as the second. In 2018, Gregory Funston ''et al.'' noted that Easter's redescription had "several ethical problems", including plaigiarized text. Although this is not enough reason to invalidate ''Ajancingenia'', they reclassified it as a species of ''Heyuannia'', creating the new combination ''Heyuannia yanshini'', in order to avoid an ethical dilemma. This assignment has been accepted by many papers since.


Description

''Heyuannia'' is a medium-sized oviraptorid. Gregory S. Paul in 2010 estimated its length at 1.5 metres, the weight at twenty kilograms. Its toothless skull is relatively short with a steep snout. It had very short arms and digits, and its first digit was reduced.Lü, J. (2005). ''Oviraptorid dinosaurs from Southern China.'' Beijing: Geological Publishing House. . 200 pages + 8 plates. (In Chinese: pp 1-83, including 36 figures & 3 tables. In English: pp 85-200, including 5 geological figures.)


Classification

''Heyuannia'' was assigned by Lü to 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 on ...
in 2002. Its exact placement within this group is uncertain. Later analyses either resulted in a position in the
Oviraptorinae 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 ...
or the Ingeniinae. According to Lü the morphology of the shoulder girdle of ''Heyuannia'' supports the hypothesis that oviraptorosaurians were secondarily flightless birds. 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 ...
follows the 2017
phylogenetic analysis In biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that pro ...
by Funston and colleagues:


Paleobiology

Preservation of the pigments
biliverdin Biliverdin (latin for green bile) is a green tetrapyrrolic bile pigment, and is a product of heme catabolism.Boron W, Boulpaep E. Medical Physiology: a cellular and molecular approach, 2005. 984-986. Elsevier Saunders, United States. It is the ...
and
protoporphyrin Protoporphyrin IX is an organic compound, classified as a porphyrin, that plays an important role in living organisms as a precursor to other critical compounds like heme (hemoglobin) and chlorophyll. It is a deeply colored solid that is not solu ...
in eggshells belonging to ''Heyuannia'' indicate that they were blue-green in color. This coloration allowed for both camouflage and sexual signalling, also seen in
American robins The American robin (''Turdus migratorius'') is a migratory bird of the true thrush genus and Turdidae, the wider thrush family. It is named after the European robin because of its reddish-orange breast, though the two species are not closely ...
and ratites. The arrangement of the eggshells suggests a partially open nest arrangement for ''Heyuannia'', and also indicates that it engaged in increased parental care.


See also

*
Timeline of oviraptorosaur research This timeline of oviraptorosaur research is a chronological listing of events in the history of paleontology focused on the oviraptorosaurs, a group of beaked, bird-like theropod dinosaurs. The early history of oviraptorosaur paleontology is cha ...


References

{{Taxonbar, from=Q134594 Oviraptorids Late Cretaceous dinosaurs of Asia Maastrichtian life Fossils of China Paleontology in Guangdong Fossils of Mongolia Barun Goyot Formation Fossil taxa described in 2002 Taxa named by Lü Junchang