Jeholosaurus
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''Jeholosaurus'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
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 insi ...
n
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 ...
from the Early
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 is thought to have been a
herbivorous A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat n ...
small
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
.


History of discovery

The first two ''Jeholosaurus'' specimens were found in
2000 2000 was designated as the International Year for the Culture of Peace and the World Mathematics, Mathematical Year. Popular culture holds the year 2000 as the first year of the 21st century and the 3rd millennium, because of a tende ...
at Lujiatun near Beipiao City,
Liaoning Province ) , image_skyline = , image_alt = , image_caption = Clockwise: Mukden Palace in Shenyang, Xinghai Square in Dalian, Dalian coast, Yalu River at Dandong , image_map = Liaoning in China (+all claims hatched).svg , ...
,
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 ...
, and named and described the same year by Xu Xing, Wang Xioalin and You Hailu. The
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...
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 ) , image_skyline = , image_alt = , image_caption = Clockwise: Mukden Palace in Shenyang, Xinghai Square in Dalian, Dalian coast, Yalu River at Dandong , image_map = Liaoning in China (+all claims hatched).svg , ...
and northern
Hebei Hebei is a Provinces of China, province in North China. It is China's List of Chinese administrative divisions by population, sixth-most populous province, with a population of over 75 million people. Shijiazhuang is the capital city. It bor ...
. The specific name refers to the geographical area of Shangyuan where the fossil site is located. The
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...
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 Romanization of Chinese, transcribed as Yihsien Formation or Yixiang Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the Barremian stage of the Early Cretaceous. I ...
, dating from the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 125.77 Ma (million years ago) and 121.4 ± 1.0 Ma (Historically, this stage was placed at 129.4 million to approximately 125 million years ago) It is a ...
, about 126-124 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 co ...
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 In 2009, 4 new well-preserved specimens were referred to ''Jeholosaurus'': IVPP V15716, a partial skull; IVPP V15717, a complete skull; IVPP V15718, a complete skull; IVPP V15719, a partial skeleton. These specimens also have features of immaturity.


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 jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
are each. The forelimbs are long and the hindlimbs are . The
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
is long and the
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
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 specimen 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 mammals h ...
has six teeth and the
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
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 regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize ...
, eating both plants and animals. The deeply inset ventral margin of the maxilla suggests fleshy cheeks may have been present. The nasals have large
foramina In anatomy and osteology, a foramen (; : foramina, or foramens ; ) is an opening or enclosed gap within the dense connective tissue (bones and deep fasciae) of extant and extinct amniote animals, typically to allow passage of nerves, arter ...
dorsolaterally and a midline fossa. No palpebrae are preserved. Though
cervical vertebrae In tetrapods, cervical vertebrae (: 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 saurop ...
and
caudal vertebrae Caudal vertebrae are the vertebrae of the tail in many vertebrates. In birds, the last few caudal vertebrae fuse into the pygostyle, and in apes, including humans, the caudal vertebrae are fused into the coccyx. In many reptiles, some of the caud ...
have been preserved, their exact number is unknown. The femur is bowed and has an anterior trochanter 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 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 (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
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: a row of small foramina on the outer surface of the nasal immediately above the premaxillary articulation and parapophyses absent from dorsal vertebrae 1 and 2. Jeholosaurus had a very rapid rate of maxillary tooth replacement at 46 days, more rapid than most basal ornithischians. As it developed ontogenetically, its premaxillary tooth replacement rate slowed from 25 days to 33 days.


Phylogeny

''Jeholosaurus'' is an
ornithischia Ornithischia () is an extinct clade 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 ...
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 (; : is ...
, 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 clade 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 ...
''
incertae sedis or is a term used for a taxonomy (biology), 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: 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 Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...
; 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. 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 ( ; from Ancient Greek 'branch') 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 ...
analyses have recovered a basal position in the Ornithopoda. The following
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
was based on analysis by Makovicky ''et al.'', 2011. 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. The cladogram below results from analysis by Herne ''et al.'', 2019.


References

{{Taxonbar, from=Q137694 Thescelosauridae Dinosaur genera Barremian dinosaurs Yixian Formation Fossil taxa described in 2000 Taxa named by Xu Xing Dinosaurs of China