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''Yunganglong'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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 basal
hadrosauroid Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or hadrosaurids, and all dinosaurs more closely related to them than to ''Iguanodon''. Their remains have been recovered in Asia, Eu ...
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 ...
known from the early
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 ...
lower
Zhumapu Formation The Zhumapu Formation is an early Late Cretaceous (estimated Cenomanian)Wang et al., 2015, p.3 geologic formation in Shanxi Province, China. The hadrosauroids ''Yunganglong'' and ''Zuoyunlong''Wang et al., 2013, p.4Wang et al., 2015, p.2 and the ...
of
Zuoyun County Zuoyun County is a county in the north of Shanxi province, China, bordering Inner Mongolia to the north. It is the westernmost county-level division of the prefecture-level city of Datong Datong is a prefecture-level city in northern Shanxi ...
,
Shanxi Province Shanxi (; ; Chinese postal romanization, formerly romanised as Shansi) is a landlocked Provinces of China, province of the China, People's Republic of China and is part of the North China region. The capital and largest city of the province is ...
of northeastern
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 ...
. It contains a single species, ''Yunganglong datongensis''.


Discovery and naming

''Yunganglong'' was first described and named by Run-Fu Wang, Hai-Lu You, Shi-Chao Xu, Suo-Zhu Wang, Jian Yi, Li-Juan Xie, Lei Jia and Ya-Xian Li 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 ...
and 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 ...
is ''Y. datongensis''. The generic name honors
Yungang Grottoes The Yungang Grottoes (), formerly the Wuzhoushan Grottoes (), are ancient Chinese Buddhist temple grottoes near the city of Datong in the province of Shanxi. They are excellent examples of rock-cut architecture and one of the three most famous an ...
, a
UNESCO World Heritage site A World Heritage Site is a landmark or area with legal protection by an international convention administered by the United Nations Educational, Scientific and Cultural Organization (UNESCO). World Heritage Sites are designated by UNESCO for h ...
built in the 5th and 6th centuries about 50 km east of the fossil locality, and derived from ''long'' meaning "
dragon A dragon is a reptilian legendary creature that appears in the folklore of many cultures worldwide. Beliefs about dragons vary considerably through regions, but dragons in western cultures since the High Middle Ages have often been depicted as ...
" in
Chinese Chinese can refer to: * Something related to China * Chinese people, people of Chinese nationality, citizenship, and/or ethnicity **''Zhonghua minzu'', the supra-ethnic concept of the Chinese nation ** List of ethnic groups in China, people of va ...
. The specific name refers to the city of
Datong Datong is a prefecture-level city in northern Shanxi Province in the People's Republic of China. It is located in the Datong Basin at an elevation of and borders Inner Mongolia to the north and west and Hebei to the east. As of the 2020 cens ...
, located in northern
Shanxi province Shanxi (; ; Chinese postal romanization, formerly romanised as Shansi) is a landlocked Provinces of China, province of the China, People's Republic of China and is part of the North China region. The capital and largest city of the province is ...
, where the holotype was found. ''Yunganglong'' is known solely from 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 ...
SXMG V 00001, field number ZY007, an associated but disarticulated partial
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
housed at the Shanxi Museum of Geology. The holotype came from a single individual, and includes the caudodorsal part of the skull, ZY007-37 and ZY007-38, separated along the floor of the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
; two
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
ZY007-40 and ZY007-41; partial dorsal neural arch and neural processes ZY007-36; two
caudal vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
, including the proximal ZY007-27 and the middle ZY007-19; distal portions of both
ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from Naples. It is the largest of the Phlegrean Islands. Roughly trapezoidal in shape, it measures approximately east to west ...
(left ZY007-11 and right ZY007-12, distal end of left
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
ZY007-32, proximal portion of right
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
ZY007-1, and distal portion of left tibia with astragalus ZY007-2. The remains were collected in 2011 from Locality 7 of the
Zhumapu Formation The Zhumapu Formation is an early Late Cretaceous (estimated Cenomanian)Wang et al., 2015, p.3 geologic formation in Shanxi Province, China. The hadrosauroids ''Yunganglong'' and ''Zuoyunlong''Wang et al., 2013, p.4Wang et al., 2015, p.2 and the ...
, as a part of a project to find dinosaurs for the Shanxi Museum of Geology, initiated by the Department of Land and Resources of Shanxi Province. SXMG V 00001 was found in the vicinity of
Zuoyun County Zuoyun County is a county in the north of Shanxi province, China, bordering Inner Mongolia to the north. It is the westernmost county-level division of the prefecture-level city of Datong Datong is a prefecture-level city in northern Shanxi ...
, from the lower part of Zhumapu Formation, dating to the early
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 ...
based on
biostratigraphic Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of Bio ...
correlations, overlying the late
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
Zuoyun Formation. Other than SXMG V 00001,
ankylosaur Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
and
ceratopsia Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. ...
n remains were found from the newly discovered localities. The first dinosaur record in Shanxi Province was reported by Young (1958), from two localities in Zuoyun County. He assigned hadrosauroid material recovered from the Xinyaogou locality of the Zuoyun area to ''
Bactrosaurus johnsoni ''Bactrosaurus'' (; meaning "Club lizard," "baktron" = club + ''sauros'' = lizard) is a genus of herbivorous hadrosauroid dinosaur that lived in Asia during the Late Cretaceous, about 96-85 million years ago. The position ''Bactrosaurus'' occupie ...
'', including two isolated teeth, some vertebrae including a series of 25 caudal vertebrae, one
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
, one right
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
and several manual and pedal bones. Although no overlapping material exists, based on the more basal phylogenetic position and lower stratigraphic horizon of ''Yunganglong'' compared to ''Bactrosaurus'', Young's material more probably pertains to ''Yunganglong''.


Description

Wang ''et al.'' (2013) diagnosed ''Yunganglong datongensis'' using a unique combination of four characters. The caudal surface of the supraoccipital in ''Yunganglong'' and more advanced hadrosauroids is inclined steeply forward at approximately 45°, while nearly vertical in '' Jintasaurus'' and less derived Hadrosauriformes. The horizontal portion of the paroccipital process is caudolaterally extended and accompanied by the
squamosal The squamosal is a skull bone found in most reptiles, amphibians, and birds. In fishes, it is also called the pterotic bone. In most tetrapods, the squamosal and quadratojugal The quadratojugal is a skull bone present in many vertebrates, including ...
, but laterally extended in ''Jintasaurus'' and less derived Hadrosauriformes. As observed in ''Yunganglong'' and ''Jintasaurus'', but not in ''Bactrosaurus'' and more advanced taxa, the pendent portion of the paroccipital process does not curve cranially. Finally, ''Yunganglong'' and less derived Hadrosauriformes possess a deep, U-shaped, intercondylar extensor groove on the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
, partially enclosed by expansion of medial and lateral
condyles A condyle (;Entry "condyle"
in
Nanyangosaurus ''Nanyangosaurus'' is a genus of herbivorous ornithischian dinosaur belonging to Hadrosauroidea that lived in the Late Cretaceous of present-day Henan Province, China. In 1994 the remains of ''Nanyangosaurus'' were found near the village of Hou ...
'') it is fully enclosed. ''Yunganglong'' cannot be directly compared with three other non-hadrosaurid hadrosauroids. These include ''
Shuangmiaosaurus ''Shuangmiaosaurus'' is a genus of herbivorous ornithischian dinosaur which lived during the late Albian age of the Early Cretaceous Period, about 100 million years ago.Carpenter, K. and Ishida, Y. (2010).Early and “Middle” Cretaceous Iguano ...
'' from the early Late Cretaceous of
Liaoning Province Liaoning () is a coastal provinces of China, province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and i ...
of northeastern China, and two other early Late Cretaceous taxa from North America: the
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
''
Protohadros ''Protohadros'' (meaning "first hadrosaur") is a genus of herbivorous ornithischian dinosaur from the Late Cretaceous ( Cenomanian stage). Gary Byrd, a part-time palaeontologist, discovered some remains of this euornithopod (ribs and an ungual) ...
'' and the
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
'' Jeyawati''. Like other hadrosauroids, ''Yunganglong'' was a ground-dwelling
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
that could walk
bipedal Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
ly, and could stand on all four legs. The skull of ''Yunganglong'' is relatively wide and low.


Phylogeny

Based on its
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
Wang et al. (2013) showed that ''Yunganglong'' was more derived than the basal hadrosauriform ''
Iguanodon ''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species have been classified in the genus ''Iguanodon'', dating from the late Jurassic Period to the early Cretaceous Period of Asia, Eu ...
'' but more primitive than the
hadrosaurid Hadrosaurids (), or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which inclu ...
s, such as the well known ''
Edmontosaurus ''Edmontosaurus'' ( ) (meaning "lizard from Edmonton") is a genus of hadrosaurid (duck-billed) dinosaur. It contains two known species: ''Edmontosaurus regalis'' and ''Edmontosaurus annectens''. Fossils of ''E. regalis'' have been found in rocks ...
'' and ''
Corythosaurus ''Corythosaurus'' (; ) is a genus of hadrosaurid "duck-billed" dinosaur from the Upper Cretaceous Period (geology), Period, about 77–75.7 million years ago. It lived in what is now North America. Its name means "helmet lizard", derived fr ...
''. To further explore the
phylogenetic 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 ...
position of ''Y. datongensis'' among
hadrosauriform Ankylopollexia is an extinct clade of ornithischian dinosaurs that lived from the Late Jurassic to the Late Cretaceous. It is a derived clade of iguanodontian ornithopods and contains the subgroup Styracosterna. The name stems from the Greek word ...
s, Wang ''et al.'' (2013) used a modified version of the data matrix first presented by Sues and Averianov (2009). ''
Nanyangosaurus ''Nanyangosaurus'' is a genus of herbivorous ornithischian dinosaur belonging to Hadrosauroidea that lived in the Late Cretaceous of present-day Henan Province, China. In 1994 the remains of ''Nanyangosaurus'' were found near the village of Hou ...
'', ''
Shuangmiaosaurus ''Shuangmiaosaurus'' is a genus of herbivorous ornithischian dinosaur which lived during the late Albian age of the Early Cretaceous Period, about 100 million years ago.Carpenter, K. and Ishida, Y. (2010).Early and “Middle” Cretaceous Iguano ...
'', and ''Yunganglong'' were added to the matrix, and two character codings were modified. In the strict consensus tree, ''Yunganglong'' was recovered as more advanced than ''
Probactrosaurus ''Probactrosaurus'' (meaning "before ''Bactrosaurus''") is an early herbivorous hadrosauroid iguanodont dinosaur. It lived in China during the Late Cretaceous period. Discovery and species In 1959 and 1960 a Soviet-Chinese expedition uncovered ...
'', in an unresolved
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with '' Jintasaurus'', ''
Protohadros ''Protohadros'' (meaning "first hadrosaur") is a genus of herbivorous ornithischian dinosaur from the Late Cretaceous ( Cenomanian stage). Gary Byrd, a part-time palaeontologist, discovered some remains of this euornithopod (ribs and an ungual) ...
'', ''Nanyangosaurus'', ''Shuangmiaosaurus'', ''
Levnesovia ''Levnesovia'' is a genus of hadrosauroid dinosaur from the Late Cretaceous Bissekty Formation of Uzbekistan. It was related to ''Bactrosaurus''. The type species is ''L. transoxiana''. The genus name honours the late Russian paleontologist Lev ...
'', ''
Bactrosaurus ''Bactrosaurus'' (; meaning "Club lizard," "baktron" = club + ''sauros'' = lizard) is a genus of herbivorous hadrosauroid dinosaur that lived in Asia during the Late Cretaceous, about 96-85 million years ago. The position ''Bactrosaurus'' occupie ...
'', ''
Tanius ''Tanius'' (meaning "of Tan") is a genus of hadrosauroid dinosaur. It lived in the Late Cretaceous of China. The type species, named and described in 1929 by Carl Wiman, is ''Tanius sinensis''. The generic name honours the Chinese paleontologist ...
'', ''
Telmatosaurus ''Telmatosaurus'' (meaning "marsh lizard") is a genus of basal hadrosauromorph dinosaur from the Late Cretaceous of Romania. It was a relatively small hadrosaur, measuring approximately in length and in body mass, which has been explained as a ...
'' and the clade formed by ''Aralosaurus'' and
Hadrosauridae Hadrosaurids (), or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which includ ...
. The 50% majority rule tree resolved its phylogenetic position further, as shown in the
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 ...
below.


References

{{Taxonbar, from=Q15079382 Late Cretaceous dinosaurs of Asia Iguanodonts Fossil taxa described in 2013 Paleontology in Shanxi Ornithischian genera