Wuerhosaurus Ordosensis
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''Wuerhosaurus'' 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
stegosaurid Stegosauridae is a family of thyreophoran dinosaurs (armoured dinosaurs) within the suborder Stegosauria. The clade is defined as all species of dinosaurs more closely related to ''Stegosaurus'' than ''Huayangosaurus''.David B. Weishampel, Peter ...
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 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 ...
Period of western
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
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an autonomous region of the People's Republic of China. Its border includes most of the length of China's border with the country of Mongolia. Inner Mongolia also accounts for a ...
. As such, it was one of the last genera of
stegosauria Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe, ...
ns known to have existed, since most others lived in the late
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
.


Discovery and species

''Wuerhosaurus homheni'' is 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 ...
, described by
Dong Zhiming Dong Zhiming (Chinese: 董枝明, Pinyin: ''Dǒng Zhimíng''; born January 1937) is a Chinese vertebrate paleontologist formerly employed at the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP) in Beijing. He began working at the ...
in 1973 from the
Tugulu Group The Tugulu Group () is a geological Group in Xinjiang, China whose strata date back to the Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major ...
in
Xinjiang Xinjiang, SASM/GNC: ''Xinjang''; zh, c=, p=Xīnjiāng; formerly romanized as Sinkiang (, ), officially the Xinjiang Uygur Autonomous Region (XUAR), is an autonomous region of the People's Republic of China (PRC), located in the northwest ...
, western China. The generic name is derived from the city of Wuerho. Three separate localities in the Wuerho Valley were discovered to contain material from the new stegosaur: , 64043 and 64045. The remains consisted of 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 ...
,
Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name suggest ...
(IVPP) V.4006, a skull-less fragmentary skeleton, and the
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 ...
IVPP V.4007. Holotype material includes a mostly complete
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
and
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
lacking the
ischium The ischium () form ...
, the first
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 ...
, two
dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae and they are intermediate in size between the cervical ...
, a
scapulocoracoid The scapulocoracoid is the unit of the pectoral girdle that contains the coracoid and scapula. The coracoid itself is a beak-shaped bone that is commonly found in most vertebrates with a few exceptions. The scapula is commonly known as the ''shoulde ...
,
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
phalanx The phalanx ( grc, φάλαγξ; plural phalanxes or phalanges, , ) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar pole weapons. The term is particularly use ...
, as well as two dermal plates. Three posterior caudal vertebrae from the tail and a partial
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
of a second individual form the paratype, and Dong referred a partial
ischium The ischium () form ...
from a third locality to ''Wuerhosaurus''. A smaller stegosaur from the
Ejinhoro Formation The Ejinhoro Formation () is a geological Formation (geology), formation in Inner Mongolia, north People's Republic of China, China, whose strata date back to the Early Cretaceous period (Aptian/Albian age.Sereno, P.C. (2010). "Taxonomy, cranial mo ...
in the Ordos Basin in
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an autonomous region of the People's Republic of China. Its border includes most of the length of China's border with the country of Mongolia. Inner Mongolia also accounts for a ...
was found in 1988. When the specimen (IVPP V.6877) was described by Dong in 1993, it was named ''W. ordosensis'', as it was from a similar age and had a similar anatomy. The holotype of the species includes a nearly complete torso, consisting of three
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 ...
, all eleven dorsal vertebrae (with attached ribs), a complete
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
with a right ilium, and the first five caudal vertebrae, all articulated. An additional dorsal vertebra and dermal plate were referred to the taxon when it was named. In 2014 Ulansky named a new species of ''Wuerhosaurus'', "W. mongoliensis" for vertebrae and pelvic material, but the name is an invalid ''
nomen nudum In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...
''. It was formally described as ''
Mongolostegus ''Mongolostegus'' (meaning "Mongolian roof") is a genus of stegosaur from the Early Cretaceous (Aptian-Albian) Dzunbain Formation of Mongolia. The type and only species is ''M. exspectabilis'', known from a single specimen previously under the ...
'' in 2018.


Description

''Wuerhosaurus homheni'' was probably a broad-bodied animal, reaching in length and in body mass. Only a few scattered bones have been found, making a full restoration difficult. Its dorsal plates were at first thought to have been much rounder or flatter than other stegosaurids, but Maidment established this was an illusion caused by breakage: their actual form is unknown. ''W. homheni'' had a pelvis of which the front of the ilia strongly flared outwards indicating a very broad belly. The neural spines on the tail base were exceptionally tall. In contrast, ''W. ordosensis'' was much smaller, estimated by Paul to have been long and weigh . It had a relatively long neck with a broad pelvis, but the neural spines are shorter.


Classification


Phylogeny

''Wuerhosaurus'' is one of the most derived stegosaurians, being closely related to either ''
Dacentrurus ''Dacentrurus'' (meaning "tail full of points"), originally known as ''Omosaurus'', is a genus of stegosaurian dinosaur from the Late Jurassic to Early Cretaceous (154 - 140 mya) of Europe. Its type species, ''Omosaurus armatus'', was named in 18 ...
'' and ''Hesperosaurus'', or ''Hesperosaurus'' and ''Stegosaurus'', depending on phylogenetic analysis.
Carpenter Carpentry is a skilled trade and a craft in which the primary work performed is the cutting, shaping and installation of building materials during the construction of buildings, Shipbuilding, ships, timber bridges, concrete formwork, etc. ...
''et al.'' (2001) recovered ''Wuerhosaurus'' in the former relationship, close to ''Hesperosaurus'' and ''Dacentrurus'' as basal in
Stegosauridae Stegosauridae is a family of thyreophoran dinosaurs (armoured dinosaurs) within the suborder Stegosauria. The clade is defined as all species of dinosaurs more closely related to ''Stegosaurus'' than ''Huayangosaurus''.David B. Weishampel, Peter ...
. ''Wuerhosaurus'' was recovered in a different position by Escaso ''et al.'' (2007), still related to ''Hesperosaurus'', but basal to a clade of ''
Lexovisaurus ''Lexovisaurus'' is a genus of stegosaur from mid-to-Late Jurassic Europe, 165.7-164.7  mya. Fossils of limb bones and armor fragments have been found in middle to late Jurassic-aged strata of England. Discovery and taxonomy In the early ...
'' and ''Stegosaurus''. Maidment ''et al.'' (2008) recovered a different placement with ''Wuerhosaurus'' as being in a clade of taxa in derived
Stegosaurinae Stegosauridae is a family of thyreophoran dinosaurs (armoured dinosaurs) within the suborder Stegosauria. The clade is defined as all species of dinosaurs more closely related to ''Stegosaurus'' than ''Huayangosaurus''.David B. Weishampel, Peter ...
, most closely related to ''Hesperosaurus'' and then ''Stegosaurus''. More recently, Maidment (2017) elaborated upon her earlier analyses, and instead resolved ''Wuerhosaurus'' as closest to ''Stegosaurus'', with ''Hesperosaurus'' being more closely related to ''
Miragaia Miragaia () is a former Freguesia, civil parish in the municipality of Porto, Portugal. In 2013, the parish merged into the new parish Cedofeita, Santo Ildefonso, Sé, Miragaia, São Nicolau e Vitória. The population in 2011 was 2,067, in an area ...
''. These results are shown below.


Synonymy with ''Stegosaurus''

Maidment and colleagues proposed in
2008 File:2008 Events Collage.png, From left, clockwise: Lehman Brothers went bankrupt following the Subprime mortgage crisis; Cyclone Nargis killed more than 138,000 in Myanmar; A scene from the opening ceremony of the 2008 Summer Olympics in Beijing; ...
that ''Wuerhosaurus'' was a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
of ''Stegosaurus'', with ''W. homheni'' being renamed ''Stegosaurus homheni'', and ''W. ordosensis'' being a
dubious Doubt is a mental state in which the mind remains suspended between two or more contradictory propositions, unable to be certain of any of them. Doubt on an emotional level is indecision between belief and disbelief. It may involve uncertainty, ...
taxon. ''W. homheni'' was synonymized because the holotype that could be located was similar to ''Stegosaurus'', and because of its phylogenetic position. ''Wuerhosaurus'' placed between ''Stegosaurus'' and ''
Hesperosaurus ''Hesperosaurus'' (meaning "western lizard", from Classical Greek (') "western" and (') "lizard") is an herbivorous stegosaurian dinosaur from the Kimmeridgian epoch of the Jurassic period, approximately 156 million years ago. Fossils of ''H ...
'', the latter being considered a species of ''Stegosaurus'' because of its age, location, and anatomy. Since ''Wuerhosaurus'' placed between two possible ''Stegosaurus'' species, Maidment ''et al.'' synonymized the taxon as well. ''Wuerhosaurus ordosensis'' was considered to be a ''nomen dubium'' because the holotype could not be found in the IVPP collections. The original description did not mention any valid diagnostic traits, and no other description provided features either, so Maidment ''et al.'' considered the taxon undiagnostic. This opinion has been contested, however, by
Carpenter Carpentry is a skilled trade and a craft in which the primary work performed is the cutting, shaping and installation of building materials during the construction of buildings, Shipbuilding, ships, timber bridges, concrete formwork, etc. ...
(2010). He discussed how the diagnoses and features used by Maidment ''et al.'' were inconsistent and generalized, with ''Wuerhosaurus homheni'' bearing numerous differences. As such, Carpenter advocated for the separation of both ''Hesperosaurus'' and ''Wuerhosaurus'' from ''Stegosaurus'', and the separation of the different ''Stegosaurus'' species from ''S. armatus''.


Paleobiology

''Wuerhosaurus'' was lower to the ground than most other stegosaurids; scientists believe that this was an adaptation to let it feed on low-growing vegetation. ''Wuerhosaurus'', like other stegosaurids, perhaps had a
thagomizer A thagomizer () is the distinctive arrangement of four spikes on the tails of Stegosauridae, stegosaurine dinosaurs. These spikes are believed to have been a defensive measure against predators.Carpenter, K., Sanders, F., McWhinney, L., and Woo ...
on the end of its tail, like that of ''
Stegosaurus ''Stegosaurus'' (; ) is a genus of herbivorous, four-legged, armored dinosaur from the Late Jurassic, characterized by the distinctive kite-shaped upright plates along their backs and spikes on their tails. Fossils of the genus have been foun ...
'' which featured four bony spikes that would most likely have been used for self-defense. A single spike was found but was seen by Dong as being positioned on the shoulder.


Paleoecology

The type species, ''W. homheni'', is known from the
Tugulu Group The Tugulu Group () is a geological Group in Xinjiang, China whose strata date back to the Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major ...
, while ''W. ordosensis'' was found in the
Ejinhoro Formation The Ejinhoro Formation () is a geological Formation (geology), formation in Inner Mongolia, north People's Republic of China, China, whose strata date back to the Early Cretaceous period (Aptian/Albian age.Sereno, P.C. (2010). "Taxonomy, cranial mo ...
. The approximate age of ''Wuerhosaurus'' is 130 mya, based on the approximate dating of the Tsaganstabian fauna, and thus the stegosaur would have lived in the
Hauterivian The Hauterivian is, in the geologic timescale, an age in the Early Cretaceous Epoch or a stage in the Lower Cretaceous Series. It spans the time between 132.9 ± 2 Ma and 129.4 ± 1.5 Ma (million years ago). The Hauterivian is preceded by the Va ...
era, which is roughly coeval with the
Wealden group The Wealden Group, occasionally also referred to as the Wealden Supergroup, is a group (a sequence of rock strata) in the lithostratigraphy of southern England. The Wealden group consists of paralic to continental (freshwater) facies sedimentary ...
, from which other stegosaur material has been found. Stegosaur footprints from the Tugulu Group in Xinjiang Province, China, have been attributed as registered by ''W. homheni'' individuals. These tracks include the world's smallest stegosaur tracks, measuring 5.7 cm in length, indicative of a dinosaur that was approximately cat-sized.


See also

*
Timeline of stegosaur research This timeline of stegosaur research is a chronological listing of events in the history of paleontology focused on the stegosaurs, the iconic plate-backed, spike-tailed herbivorous eurypod dinosaurs that predominated during the Jurassic period ...


References

* {{Taxonbar, from=Q131639 Stegosaurs Taxa named by Dong Zhiming Fossil taxa described in 1973 Fossil taxa described in 1993 Early Cretaceous dinosaurs of Asia Paleontology in Xinjiang Paleontology in Inner Mongolia Ornithischian genera Hauterivian life