Jeholosaurus Skeleton
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''Jeholosaurus'' 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
neornithischia Neornithischia ("new ornithischians") is a clade of the dinosaur order Ornithischia. It is the sister group of the Thyreophora within the clade Genasauria. Neornithischians are united by having a thicker layer of asymmetrical enamel on the inside ...
n
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 Early
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 th ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
. It is thought to have been a
herbivorous 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 ...
small
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world ...
.


History of discovery

The first two ''Jeholosaurus'' specimens were found in
2000 File:2000 Events Collage.png, From left, clockwise: Protests against Bush v. Gore after the 2000 United States presidential election; Heads of state meet for the Millennium Summit; The International Space Station in its infant form as seen from ...
at Lujiatun near
Beipiao City Beipiao () is a city in Chaoyang prefecture, Liaoning province, in Northeast China. It has a population of 202,807. The main industry in the area is coal mining. With vertical shafts of almost 1000m, these are some of the deepest coal mines in ...
,
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 ...
,
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 named and described the same year by Xu Xing, Wang Xioalin and
You Hailu In Modern English, ''you'' is the second-person pronoun. It is grammatically plural, and was historically used only for the dative case, but in most modern dialects is used for all cases and numbers. History ''You'' comes from the Proto- ...
. The type and only known species is ''Jeholosaurus shangyuanensis''. The generic name ''Jeholosaurus'' means "lizard from Jehol"; Jehol is the name of a historical province situated in western
Liaoning Liaoning () is a coastal 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 is the northernmost ...
and northern
Hebei Hebei or , (; alternately Hopeh) is a northern province of China. Hebei is China's sixth most populous province, with over 75 million people. Shijiazhuang is the capital city. The province is 96% Han Chinese, 3% Manchu, 0.8% Hui, an ...
. The specific name refers to the geographical area of Shangyuan where the fossil site is located. The type specimen of ''Jeholosaurus'', on which the genus is based, is IVPP V 12529. It was found in a layer of the early
Yixian Formation The Yixian Formation (; formerly transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its ex ...
, dating from the early
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
, about 126-125 million years old. The layers consist of fluvial sandstone interspersed with
tuff Tuff is a type of rock made of volcanic ash ejected from a vent during a volcanic eruption. Following ejection and deposition, the ash is lithified into a solid rock. Rock that contains greater than 75% ash is considered tuff, while rock cont ...
and it is thought that an enormous volcanic eruption occurred burying everything within a fifty to sixty mile radius. The holotype contains a compressed skull and a partial postcranial skeleton. The second specimen, IVPP V 12530, was referred to the species. It consists of a better preserved, perhaps smaller, skull and some neck vertebrae. Both specimens represent juvenile or at least subadult individuals.Paul, G.S., 2010, ''The Princeton Field Guide to Dinosaurs'', Princeton University Press p. 275


Description


Size

''Jeholosaurus'' was a small bipedal herbivore. Because the specimens are juvenile it is hard to ascertain the adult size. The holotype is long with a long tail, nearly half the length of the complete animal. The length of the skull is and the lower jaws of the
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
are each. The forelimbs are long and the hindlimbs are . 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 ...
is long and the
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 ...
is .Xu, Wang and You, 2000. A primitive ornithopod from the Early Cretaceous Yixian Formation of Liaoning. Vertebrata PalAsiatica 38(4)318-325.


Anatomy

Both skulls are incomplete and had to be partially restored. The referred specimens shows an extremely large orbit compared to the skull size, with a length of 40% of the total skull length. Its snout is short, also comprising 40% of the skull length. Both traits are indicative of a juvenile individual. 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 b ...
has six teeth and the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
has at least thirteen teeth. Although the back, maxillary, teeth are fan-shaped like those of a plant eater, the front premaxillary teeth are narrower and longer, more like a carnivorous dinosaur. This may mean that ''Jeholosaurus'' was
omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutri ...
, eating both plants and animals. The deeply inset ventral margin in the maxilla suggests fleshy cheeks may have been present. The nasals have large
foramina In anatomy and osteology, a foramen (;Entry "foramen"
in
dorsolaterally and a midline fossa. No
palpebrae An eyelid is a thin fold of skin that covers and protects an eye. The levator palpebrae superioris muscle retracts the eyelid, exposing the cornea to the outside, giving vision. This can be either voluntarily or involuntarily. The human eyel ...
are preserved. Though
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 ...
and
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 ...
have been preserved, their exact number is unknown. The femur is bowed and has an anterior
trochanter A trochanter is a tubercle of the femur near its joint with the hip bone. In humans and most mammals, the trochanters serve as important muscle attachment sites. Humans are known to have three trochanters, though the anatomic "normal" includes ...
slightly lower than the greater trochanter and a third as narrow as the latter. They are separated by a shallow cleft. Between the condyles of the lower femur the (anterior)
extensor In anatomy, extension is a movement of a joint that increases the angle between two bones or body surfaces at a joint. Extension usually results in straightening of the bones or body surfaces involved. For example, extension is produced by extendin ...
groove is absent. The fourth trochanter is pendant and located in a rather high position. The foot has four
metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...
s. The longest is metatarsal III with a length of ; it is placed more anteriorly than the other metatarsals. Metatarsal I is more posterior and its upper part is transversely reduced to a splint. Some distinguishing traits of ''Jeholosaurus'' include: enlarged laterodorsal nasal foramina; a quadratojugal fenestra more than 25% maximum
quadratojugal The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians. Anatomy and function In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...
length; a quadratojugal less than 30% of skull height; a
predentary Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek st ...
with almost 150% of
premaxillary 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 ...
body length; a dentary extending
posteriorly Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
almost to the posterior border of the angular; and the claw of the third toe being longer than the other third toe phalanges.


Phylogeny

''Jeholosaurus'' is an
ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek s ...
n, as is shown by its ornithischian four-pronged pelvic structure with a pubis bone pointing downward and backwards, parallel to the
ischium The ischium () form ...
, while a forward-pointing prepubic process supports the abdomen. The describers did not assign ''Jeholosaurus'' to any family, limiting themselves to a placement as
Ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek s ...
''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
''. Using the comparative method, they pointed out some similarities to basal
Euornithopoda Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world ...
: a small
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 archosauriforms, first appearing during the Triassic Period. Among extant archosaurs, birds ...
; the foramen on the quadratojugal; a large quadratic foramen; and the absence of an external fenestra in the lower jaw. However, they also noticed more derived euornithopod traits, such as the form of the greater and anterior trochanter of the thighbone, although the premaxilla not reaching the lacrimal, the high jaw joint and the premaxilla being on the same level as the maxilla, were again basal traits. The fused mandibular symphysis might indicate a relation with the
Marginocephalia Marginocephalia (/mär′jə-nō-sə-făl′ē-ən/ Latin: margin-head) is a clade of ornithischian dinosaurs that is characterized by a bony shelf or margin at the back of the skull. These fringes were likely used for display. There are two clad ...
. Very basal ornithischian traits included the presence of six teeth in the premaxilla with only a short toothless sector in front and a short hiatus with the maxillary teeth. Later
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 have recovered a basal position in the (Eu)ornithopoda. 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 ...
was based on analysis by Makovicky ''et al.'', 2011. The following cladogram was based on analysis by Zheng and colleagues in 2009.Zheng, X-.T., You, H.-L., Xu, X. and Dong, Z.-M. (2009). "An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures." ''Nature'', 458(19): 333-336. Han ''et al.''found a similar topology to that of Makovicky ''et al.'', 2011, in 2012, with a clade composed of ''Jeholosaurus'', ''Haya'', and ''Changchunsaurus''. They named this clade
Jeholosauridae Jeholosaurids were herbivorous neornithischian dinosaurs from the Cretaceous Period (Aptian - Cenomanian). The family was first proposed by Han ''et al.'' in 2012. The jeholosaurids were defined as all neornithischians more closely related to ''J ...
. The cladogram below results from analysis by Herne ''et al.'', 2019.


References

{{Taxonbar, from=Q137694 Early Cretaceous dinosaurs of Asia Fossil taxa described in 2000 Taxa named by Xu Xing Yixian fauna Ornithischian genera