Heyuannia
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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 extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
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 ...
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 area ...
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'', the ...
epoch, in what is now
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
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 million, ...
. 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 million, ...
. Two species are known: ''H. huangi'', 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 ...
in 2002 from the
Dalangshan Formation The Dalangshan Formation (), also referred to as the Dalangshan Group is a geological formation in the Sanshui District of Guangdong Province, China, the strata of which date back to the Late Cretaceous period. Dinosaur remains are among the fossi ...
; 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 Pro ...
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 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 ...
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 several ...
, 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 The Dalangshan Formation (), also referred to as the Dalangshan Group is a geological formation in the Sanshui District of Guangdong Province, China, the strata of which date back to the Late Cretaceous period. Dinosaur remains are among the fossi ...
. 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). Of ...
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 offensiv ...
, 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 capi ...
(
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 Pro ...
), Mongolia, from whence it was collected, while the specific name ''yanshini'' was chosen in honour of academician
Aleksandr Leonidovich Yanshin Alexander is a male given name. The most prominent bearer of the name is Alexander the Great, the king of the Ancient Greek kingdom of Macedonia who created one of the largest empires in ancient history. Variants listed here are Aleksandar, Al ...
(1911–1999), who was adviser and mentor to Rinchen Barsbold during his time at the Paleontological Institute 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 List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
.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 Wh ...
''. The generic name ''Ingenia'' was preoccupied by the generic name of ''
Ingenia mirabilis ''Ingenia'' is a genus of marine nematode worms native to Brazil, with a single known species, ''I. mirabilis''.Gerlach, S. A. (1957). Die Nematodenfauna des Sandstrandes an der Küste von Mittelbrasilien (Brasilianische Meerse-Nematoden IV). ' ...
'' (Gerlach, 1957), a tripyloidid
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
. 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 fact ...
. 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 ungulates have ungual phalanges, as did the sauropods and horned dinosaurs. A claw is a highly modified ungual ...
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 Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
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 one ...
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 on ...
or the
Ingeniinae 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 o ...
. 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 d ...
follows the 2017
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 ...
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 pi ...
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 and
ratite A ratite () is any of a diverse group of flightless, large, long-necked, and long-legged birds of the infraclass Palaeognathae. Kiwi, the exception, are much smaller and shorter-legged and are the only nocturnal extant ratites. The systematics ...
s. 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