Wuerhosaurus

Wuerhosaurus is a genus of stegosaurid dinosaur from the Early Cretaceous Period of China and Mongolia. As such, it was one of the last genera of stegosaurians known to have existed, since most others lived in the late Jurassic.[3]

Wuerhosaurus
Temporal range:
Early Cretaceous, 132–129 Ma
Wuerhosaurus by ABelov2014
Restoration of W. homheni with low plates
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Ornithischia
Suborder: Stegosauria
Family: Stegosauridae
Subfamily: Stegosaurinae
Genus: Wuerhosaurus
Dong, 1973[1]
Type species
Wuerhosaurus homheni
Dong, 1973[1]
Species
  • W. homheni Dong, 1973[1]
  • W. ordosensis Dong, 1993[2]
Synonyms
  • Stegosaurus homheni (Dong, 1973)

Discovery and species

Wuerhosaurus-Paleozoological Museum of China
Plate of W. homheni, Paleozoological Museum of China

Wuerhosaurus homheni is the type species, described by Dong Zhiming in 1973 from the Tugulu Group in Xinjiang, 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: 64043-5, 64043 and 64045.[1] The remains consisted of the holotype, Institute of Vertebrate Paleontology and Paleoanthropology (IVPP) V.4006, a skull-less fragmentary skeleton, and the paratype IVPP V.4007.[4] Holotype material includes a mostly complete pelvis and sacrum lacking the ischium, the first caudal vertebrae, two dorsal vertebrae, a scapulocoracoid, humerus and phalanx, as well as two dermal plates. Three posterior caudal vertebrae from the tail and a partial ulna of a second individual form the paratype, and Dong referred a partial ischium from a third locality to Wuerhosaurus.[1]

A smaller stegosaur from the Ejinhoro Formation in the Ordos Basin in Inner Mongolia, 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, all eleven dorsal vertebrae (with attached ribs), a complete sacrum 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.[2] In 2014 Ulansky named a new species of Wuerhosaurus, "W. mongoliensis" for vertebrae and pelvic material, but the name is an invalid nomen nudum.[5] It was formally described as Mongolostegus in 2018.

Description

Wuerhosaurus homheni was probably a broad-bodied animal. Gregory S. Paul in 2016 estimated the length at 7 m (23 ft) and the weight at 4 tonnes (630 st).[6] Only a few scattered bones have been found, making a full restoration difficult.[3] Its dorsal plates were at first thought to have been much rounder or flatter than other stegosaurids,[7] 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.

W. ordosensis was estimated by Paul to have been 5 m (16 ft) long and weigh 1.2 tonnes (190 st). It too has a broad pelvis but the neural spines are shorter. The neck seems to have been relatively long.[6]

Classification

Phylogeny

Wuerhosaurus is one of the most derived stegosaurians, being closely related to either Dacentrurus and Hesperosaurus, or Hesperosaurus and Stegosaurus, depending on phylogenetic analysis. Carpenter et al. (2001[8]) recovered Wuerhosaurus in the former relationship, close to Hesperosaurus and Dacentrurus as basal in Stegosauridae. Wuerhosaurus was recovered in a different position by Escaso et al. (2007[9]), still related to Hesperosaurus, but basal to a clade of Lexovisaurus and Stegosaurus. Maidment et al. (2008[10]) recovered a different placement with Wuerhosaurus as being in a clade of taxa in derived Stegosaurinae, most closely related to Hesperosaurus and then Stegosaurus.[10] More recently, Maidment (2017[11]) elaborated upon her earlier analyses, and instead resolved Wuerhosaurus as closest to Stegosaurus, with Hesperosaurus being more closely related to Miragaia. These results are shown below.[11]

Wuerhosaurus
Restoration of W. homheni with tall plates
Stegosauria

Huayangosaurus taibaii

Chungkingosaurus jiangbeiensis

Tuojiangosaurus multispinus

Paranthodon africanus

Jiangjunosaurus junggarensis

Gigantspinosaurus sichuanensis

Kentrosaurus aethiopicus

Dacentrurus armatus

Loricatosaurus priscus

Hesperosaurus mjosi

Miragaia longicollum

Stegosaurus stenops

Wuerhosaurus homheni

Synonymy with Stegosaurus

Maidment and colleagues proposed in 2008 that Wuerhosaurus was a junior synonym of Stegosaurus, with W. homheni being renamed Stegosaurus homheni, and W. ordosensis being a dubious 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, 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.[10] This opinion has been contested, however, by Carpenter (2010[12]). 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.[12]

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 on the end of its tail, like that of Stegosaurus 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, while W. ordosensis was found in the Ejinhoro Formation. 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 era, which is roughly coeval with the Wealden group, from which other stegosaur material has been found.[13]

References

  1. ^ a b c d e Dong, Z. (1973). "Dinosaurs from Wuerho". Reports of Paleontological Expedition to Sinkiang (II): Pterosaurian Fauna from Wuerho, Sinkiang (in Chinese). 11. Memoirs of the Institute of Vertebrate Paleontology and Paleoanthropology, Academia Sinica. pp. 45–52.
  2. ^ a b Dong, Z. (1993). "A new species of stegosaur (Dinosauria) from the Ordos Basin, Inner Mongolia, People's Republic of China". Canadian Journal of Earth Sciences. 30 (10): 2174–2176. doi:10.1139/e93-188.
  3. ^ a b Palmer, D., ed. (1999). The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals. London: Marshall Editions. p. 156. ISBN 1-84028-152-9.
  4. ^ Dong, Z. (1990). "Stegosaurs of Asia". In Carpenter, Kenneth; Currie, Philip J. (eds.). Dinosaur Systematics: Approaches and Perspectives. Cambridge University Press. pp. 255–268. ISBN 978-0-521-43810-0.
  5. ^ Galton, P.M.; Carpenter, K. (2016). "The plated dinosaur Stegosaurus longispinus Gilmore, 1914 (Dinosauria: Ornithischia; Upper Jurassic, western USA), type species of Alcovasaurus n. gen."". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 279 (2): 185–208. doi:10.1127/njgpa/2016/0551.
  6. ^ a b Paul, G.S. (2016). The Princeton Field Guide to Dinosaurs (2 ed.). Princeton University Press. p. 248. ISBN 978-0-691-16766-4.
  7. ^ Dodson, P., ed. (1993). "Wuerhosaurus". The Age of Dinosaurs. Publications International, LTD. p. 102. ISBN 0-7853-0443-6.
  8. ^ Carpenter, K.; Miles, C.A.; Cloward, K. (2001). "New primitive stegosaur from the Morrison Formation, Wyoming". In Carpenter, K. (ed.). The Armored Dinosaurs. Indiana University Press. pp. 55–75. ISBN 978-0-253-33964-5.
  9. ^ Escaso, F.; Ortega, F.; Dantas, P.; Malafaia, E.; Pimentel, N.L.; Pereda Suberbiola, X.; Sanz, J.L.; Kullburg, J.C.; Kullburg, M.C.; Barriga, F. (2007). "New evidence of shared dinosaur across Upper Jurassic Proto-North Atlantic: Stegosaurus from Portugal". Naturwissenschaften. 94 (5): 367–374. doi:10.1007/s00114-006-0209-8. PMID 17187254.
  10. ^ a b c Maidment, Susannah C.R.; Norman, David B.; Barrett, Paul M.; Upchurch, Paul (2008). "Systematics and phylogeny of Stegosauria (Dinosauria: Ornithischia)". Journal of Systematic Palaeontology. 6 (4): 367–407. doi:10.1017/S1477201908002459.
  11. ^ a b Raven, T.j.; Maidment, S.C.R. (2017). "A new phylogeny of Stegosauria (Dinosauria, Ornithischia)" (PDF). Palaeontology. 2017 (3): 1–8. doi:10.1111/pala.12291.
  12. ^ a b Carpenter, K. (2010). "Species concept in North American stegosaurs". Swiss Journal of Geosciences. 103 (2): 155–162. doi:10.1007/s00015-010-0020-6.
  13. ^ Donovan, T. (2002). "RE: Tsagantsabian age". Dinosaur Mailing List. Cleveland Museum of Natural History. Retrieved 2018-01-08.
  • Dong Zhiming (1992). Dinosaurian Faunas of China. China Ocean Press, Beijing. ISBN 3-540-52084-8.

See also

Acantholipan

Acantholipan is a genus of herbivorous nodosaurid dinosaur from Mexico from the early Santonian age of the Late Cretaceous. It includes one species, Acantholipan gonzalezi.

Ankylosaurinae

Ankylosaurinae is a subfamily of ankylosaurid dinosaurs, existing from the Early Cretaceous about 105 million years ago until the end of the Late Cretaceous, about 66 mya. Many genera are included in the clade, such as Ankylosaurus, Pinacosaurus, Euoplocephalus, and Saichania.

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 125.0 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), approximately. The Aptian succeeds the Barremian and precedes the Albian, all part of the Lower/Early Cretaceous.The Aptian partly overlaps the upper part of the regionally used (in Western Europe) stage Urgonian.

The Selli Event, also known as OAE1a, was one of two oceanic Anoxic events in the Cretaceous period, which occurred around 120 Ma and lasted approximately 1 to 1.3 million years. The Aptian extinction was a minor extinction event hypothesized to have occurred around 116 to 117 Ma.

Aptian extinction

The Aptian extinction was an extinction event of the early Cretaceous Period. It is dated to c. 116 or 117 million years ago, in the middle of the Aptian stage of the geological time scale, and has sometimes been termed the mid-Aptian extinction event as a result.

It is classified as a minor extinction event, rather than a major event like the famous Cretaceous–Paleogene extinction event that brought about the end of the "age of dinosaurs" and the Mesozoic Era. The Aptian event is most readily detected among marine rather than terrestrial fossil deposits. Nonetheless, "From a palaeobotanical perspective, the Aptian Extinction Event is an episode of importance, deserving a higher status among other minor events."The Aptian event may have been causally connected with the Rahjamal Traps volcanism episode in the Bengal region of India, associated with the Kerguelen hotspot of volcanic activity. (At the time in question, c. 116–117 Ma, India was located in the southern Indian Ocean; plate tectonics had not yet moved the Indian landmass into its present position.)

Note that the stegosaur group of dinosaurs went extinct around this time. Wuerhosaurus, probably the last of the stegosaurs, lived during this time.

Bienosaurus

Bienosaurus (meaning "Bien's lizard") is a genus of thyreophoran dinosaur from the Lower Jurassic (probably Sinemurian) Lower Lufeng Formation in Yunnan Province in China.

Bissektipelta

Bissektipelta is a genus of herbivorous ankylosaurid dinosaur from the Upper Cretaceous of Uzbekistan. Bissektipelta is monospecific, containing only the species B. archibaldi.

Dongyangopelta

Dongyangopelta is an extinct genus of nodosaurid ankylosaurian dinosaur known from the "middle" Cretaceous Chaochuan Formation (Albian or Cenomanian stage) of Dongyang, Zhejiang Province, China. Dongyangopelta was first named by Rongjun Chen, Wenjie Zheng, Yoichi Azuma, Masateru Shibata, Tianliang Lou, Qiang Jin and Xingsheng Jin in 2013 and the type species is Dongyangopelta yangyanensis. It differs from Zhejiangosaurus, the second nodosaurid from southeast China, in the characters of presacral rod, ilium, and femur. Donyangopelta is distinguishable from Zhejiangosaurus only on the basis of the morphology of its pelvic shield.

Ejinhoro Formation

The Ejinhoro Formation (simplified Chinese: 伊金霍洛组; traditional Chinese: 伊金霍洛組; pinyin: Yījīnhuòluò Zǔ) is a geological formation in Inner Mongolia, north China, whose strata date back to the Early Cretaceous period (Aptian/Albian age.Dinosaur remains are among the fossils that have been recovered from the formation.

Hesperosaurus

Hesperosaurus (meaning "western lizard", from Classical Greek ἕσπερος/hesperos "western" and σαυρος/sauros "lizard") is a herbivorous stegosaurian dinosaur from the Kimmeridgian epoch of the Jurassic period, approximately 156 million years ago.

Fossils of Hesperosaurus have since 1985 been found in the state of Wyoming and Montana in the United States of America. The type species Hesperosaurus mjosi was named in 2001. It is from an older part of the Morrison Formation, and so a little older than other Morrison stegosaurs. Several relatively complete skeletons of Hesperosaurus are known. One specimen preserves the first known impression of the horn sheath of a stegosaurian back plate.

Hesperosaurus was six to seven metres long and two to three tonnes in weight. It was a member of the Stegosauridae, quadrupedal plant-eaters protected by vertical bony plates and spikes. It was closely related to Stegosaurus and was similar to it in having two rows of, possibly alternating, plates on its back and four spikes on its tail end. The plates on its back were perhaps not as tall, but were longer. It possibly had a deeper skull than Stegosaurus.

Lianmuqin Formation

The Lianmuqin Formation is an Early Cretaceous geologic formation composed of "interbedded red green and yellow variegated mudstones and siltstones [sic]". Dinosaur remains have been recovered from it.The formation is named after Lianmuqin Town in Shanshan County, Xinjiang.

Miragaia longicollum

Miragaia (named after Miragaia, the parish in Portugal and geologic unit where its remains were found) is a long-necked

stegosaurid dinosaur. Its fossils have been found in Lower Jurassic rocks in Portugal. Miragaia has the longest neck known for any stegosaurian, which included at least seventeen vertebrae.

Mongolostegus

Mongolostegus is a genus of stegosaur from the Early Cretaceous (Aptian-Albian) of Mongolia. The type and only species is M. exspectabilis, known from a single specimen previously under the nomen nudum Wuerhosaurus mongoliensis.

Nodosaurus

Nodosaurus (meaning "knobbed lizard") is a genus of herbivorous ankylosaurian dinosaur from the Late Cretaceous, the fossils of which are found in North America.

Silvisaurus

Silvisaurus, from the Latin silva "woodland" and Greek sauros "lizard", is a nodosaurid ankylosaur from the middle Cretaceous period.

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, Africa, South America and Asia Their geographical origins are unclear; the earliest unequivocal stegosaurian, Huayangosaurus taibaii, lived in China.

Stegosaurians were armored dinosaurs (thyreophorans). Originally, they did not differ much from more primitive members of that group, being small, low-slung, running animals protected by armored scutes. An early evolutionary innovation was the development of tail spikes, or "thagomizers", as defensive weapons. Later species, belonging to a subgroup called the Stegosauridae, became larger, and developed long hindlimbs that no longer allowed them to run. This increased the importance of active defence by the thagomizer, which could ward off even large predators because the tail was in a higher position, pointing horizontally to the rear from the broad pelvis. Stegosaurids had complex arrays of spikes and plates running along their backs, hips and tails. Their necks became longer and their small heads became narrow, able to selectively bite off the best parts of cycads with their beaks. When these plant types declined in diversity, so did the stegosaurians, which became extinct during the first half of the Cretaceous period.

The first stegosaurian finds in the early 19th century were fragmentary. Better fossil material, of the genus Dacentrurus, was discovered in 1874 in England. Soon after, in 1877, the first nearly-complete skeleton was discovered in the United States. Professor Othniel Charles Marsh that year classified such specimens in the new genus Stegosaurus, from which the group acquired its name, and which is still by far the most famous stegosaurian. During the latter half of the twentieth century, many important Chinese finds were made, representing about half of the presently known diversity of stegosaurians.

Tatisaurus

Tatisaurus is a genus of ornithischian dinosaur from the Early Jurassic from the Lower Lufeng Formation in Yunnan Province in China. Little is known as the remains are fragmentary.

Tianzhenosaurus

Tianzhenosaurus (Tianzhen + Greek sauros="lizard") is a genus of ankylosaurid dinosaurs discovered in Tianzhen County, at Kangdailiang near Zhaojiagou Village, in Shanxi Province, China, in the Late Cretaceous Huiquanpu Formation. Thus far, a virtually complete skull and postcranial skeleton have been assigned to the genus, which is monotypic (T. youngi Pang & Cheng, 1998).

This was a medium-sized ankylosaurian, the skull measuring 28 cm (11 in) in length, with a total body length around 4 m (13 ft).

Vickaryous et al. (2004) placed Tianzhenosaurus within the Ankylosauridae, nested as the sister group to Pinacosaurus. Some authors have suggested that Tianzhenosaurus is actually a junior synonym of Saichania chulsanensis.

Tugulu Group

The Tugulu Group (simplified Chinese: 吐谷鲁群; traditional Chinese: 吐谷魯群; pinyin: Tǔgǔlǔ Qún) is a geological formation in Xinjiang, China whose strata date back to the Early Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.

Valanginian

In the geologic timescale, the Valanginian is an age or stage of the Early or Lower Cretaceous. It spans between 139.8 ± 3.0 Ma and 132.9 ± 2.0 Ma (million years ago). The Valanginian stage succeeds the Berriasian stage of the Lower Cretaceous and precedes the Hauterivian stage of the Lower Cretaceous.

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