Nanshiungosaurus: Difference between revisions

Nanshiungosaurus; Temporal range: Late Cretaceous, ; ~67–66 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Skeletal restoration of IVPP V4731
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Dinosauria
Clade:	Saurischia
Clade:	Theropoda
Superfamily:	†Therizinosauroidea
Family:	†Therizinosauridae
Genus:	†Nanshiungosaurus; Dong, 1979
Type species
	†Nanshiungosaurus brevispinus; Dong, 1979

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Nanshiungosaurus (meaning "Nanxiong's lizard") is a genus of therizinosaurid dinosaurs that lived in what is know Asia during the Late Cretaceous of South China. The type species, Nanshiungosaurus brevispinus, was first discovered in 1974 and described in 1979 by Dong Zhiming. It is represented by a single specimen preserving most of the cervical and dorsal vertebrae with the pelvis. A supposed and unlikely second species, "Nanshiungosaurus" bohlini, was found in 1992 and described in 1997. It is also represented by vertebrae but this species however, differs in geological age and authentic characteristics compared to the later, making its affinity to the genus unsupported.

They were large-sized therizinosaurids that are estimated to have cover 5 m (16 ft) in length and to have weighed about 907 kg (2,000 lb). Nanshiungosaurus had a very pneumatized vertebral column with the posterior cervical vertebrae of the long neck being unsually robust and slightly more elongated than the dorsals. They also were equipped with broad torsos as seen on the bulky pelvis. As for other therizinosaurids, they had a keratinous beak used when feeding, stocky feet with four weight-bearing toes and large flattened claws.

Nanshiungosaurus are classified as therizinosaurian dinosaurs. Along with Therizinosaurus and Segnosaurus, Nanshiungosaurus was one the earliest therizinosaurs to be described and named. The unusual shape of the pelvis led Dong to interpret the remains as belonging to some dwarf sauropod, but during the 1990s the genus was recognized as a segnosaur (now therizinosaur) based the pelvic similarities with Segnosaurus. At first, the research on therizinosaurs was complicated since sparce remains were known at the time and these had features from multiple dinosaur lineages leading to their interpretation as prosauropod dinosaurs.

History of discovery

In 1974, during a geological expedition at the Nanxiong Basin led by the Institute of Vertebrate Paleontology and Paleoanthropology several fossilized remains of dinosaurs were discovered by the team. Near the village of Dapingcun at the Nanxiong Formation, Guangdong Province, a relatively large and partial skeleton was found in articulation dating back to the Late Cretaceous. The specimen was labelled under the number IVPP V4731 and consisted of 12 cervical (lacking the atlas), 10 dorsal, 5 (actually 6) sacral and the first caudal vertebrae with a nearly complete, bulky pelvis only lacking the right ilium and ischium. Later on, in 1979 the specimen was formally described by the Chinese paleontologist Dong Zhiming and used as the basis for the new genus and species Nanshiungosaurus brevispinus. The generic name, Nanshiungosaurus, refers to the site of provenance Nanxiong city and is derived from the Greek σαῦρος (sauros, meaning lizard). Lastly, the specific name, brevispinus, is derived from the Latin brevis and spina (meaning short and spine, respectively) in reference to the relatively short vertebral spines. When first described, Dong mistakenly thought the specimen to have been a dwarf, strange titanosaurine sauropod characterized by a shorter but thicker neck than other sauropods based on the pelvis structure.^[1]

In 1997 Dong Zhiming and You Hailu named and described a supposed second species: "Nanshiungosaurus" bohlini, based on a skeleton found in 1992 near Mazongshan. It consists of 11 cervical and 5 dorsal vertebrae with some ribs. The specimen is catalogued as IVPP V11116 coming from the Early Cretaceous, Upper Xinminbao Group. Also, they coined the Nanshiungosauridae to contain "both species". Dong and Yu presented no evidence or argumentation supporting the assignment of the species to Nanshiungosaurus.^[2] In 2010, the North American paleontologist Lindsay Zanno considered this referral to be highly improbable since "N". bohlini dates back from the Barremian-Aptian ages and in the view of lack of synapomorphies, she considered that the supposed second species is unrelated to Nanshiungosaurus and might warrant its own genus. In addition, she corrected the number of sacral vertebrae from 5 to 6 and noted that the holotype pelvis from the latter has suffered damage since collection and has been reconstructed with painted plaster in the affected areas.^[3] Moreover, the remains of "N". bohlini were recovered from the lower red beds of the Zhonggou Formation in the Gongpoquan Basin, a completely different geological context. As an overall consensus, this dubious specimen is no longer considered to be relatable to Nanshiungosaurus.^[4]

Description

Nanshiungosaurus were relatively large-bodied therizinosaurids, estimated at 5 m (16 ft) long weighing 907 kg (2,000 lb).^[5]^[6] This species can be differentiated by the possession of stocky posterior-most cervical vertebrae with opisthocoel (meaning that they were concave on their posterior sides) centra.^[3] Like other derived (advanced) therizinosaurids, Nanshiungosaurus were pot-bellied animals that had a strong build composed of stout hindlimbs with a functionally tretadactyl pes. Their arms endend up in large, recurved claws that were side to side flattened.^[7] Though the holotype specimen lacks cranial material, the preserved elements in therizinosaurids Erlikosaurus and Segnosaurus indicate that they had relatively small skulls with coarsely serrated teeth, and developed a prominent rhamphotheca (keratinous beak).^[8]^[9]

Vertebral column

Most cervicals have relatively short neural arches, but in the dorsals they are more elongated. The axis is well preserved and has a length of 13.5 cm (135 mm) with platycoelus (slightly concave at both ends) centra. Its odontoid process is fused with the anterior facet of centrum and has a blunt depression to the bottom area in order to articulate with the atlas centrum. The neural arch is narrow, the neural spine has a porterior orientation, and has rounded facets that connect with the atlantal posterior articular processes (bony projections). After the axis, the next cervicals gradually increase in size. The anterior cervicals have platycoelus centra and deep, pocket-like pleurocoels (small holes) develop on the lateral sides; a small ridge-like structure is also present. On the bottom surface, the centra are flat, gently flattened to the inner sides, and two projections originate from the borders of the centra. The anterior capitular process is compressed (flattened) with some degree of thickness, and extends from the bottom to lateral sides to be fused with the capitulum on the ribs. As for most cervicals, the neural arches and spines are short, the anterior articular processes are strongly developed in that direction, and the anterior lobes of the diapophyses (bony projections to the lateral sides) extend to the bottom and lateral directions to articulate with tubercles. Seventh and eighth cervicals are the longest with 18 cm (180 mm) long centra and posterior to them, the vertebral size slightly narrows. In most posterior cervicals, the centra are opisthocoelous with shallow posterior grooves.^[1]

The dorsal vertebrae centra, unlike in the cervicals, are narrow and platycoelous with an average length of 7 cm (70 mm). They are also pneumatic with shallow pneumatopores (pits leading to air pockets) on the lateral sides, but they reduce in size on posterior dorsals. Along with the centra, the neural arch are well pneumatized with most of the pneumatic areas filled with matrix (sediments). The capitular processes are undeveloped and replaced by poor rounded facets on the anterior surface of the centrum. In most dorsals, the neural spines are very elongated, the articular processes are located within the ligaments of the centrum, the neural spines are stocky, side to side thickened, and the top regions have stout tuberosities. A prominent spiral-like ridge is present on the bottom surface that becomes more notable to the posterior direction. Though most dorsals are completely preserved, eighth through tenth dorsals have damaged centra. On the posterior series the centra become platycoelous, approaching the sacral vertebrae structure. The sacrum is composed by six (originally five) sacral vertebrae and they are fused into a single structure, indicating that the individual was an adult at the time of death. They are lightly more longer than the dorsals and have stocky, side to ide broadened neural spines with the top surfaces bearing small depressions. A single caudal vertebra is present after the sacral series and represents the first caudal. Its neural spine is broken and the centrum is slightly amphicoelous (concave on both ends) with sub-equal height and length.^[1]

Pelvic girdle

The pelvis is represented by the well-preserved left side composed by the illium, pubis and both ischia; the right ilium and pubis were eroded though. As a whole, the pelvis is robustly built and some elements are deformed such as the left ilium, which due to taphonomical factors has been bent out of shape. Like other derived therizinosaurids, the pelvis has a opisthopubic condition where the pubis and ischium are fused and directed backwards. The ilium is stocky with an extremely well-developed and elongated preacetabular process (anterior expansion of the iliac blade), nevertheless, the postacetabular process (posterior expansion) is missing. Its pubic peduncle (a robust projection that connects to the pubis) is also well-developed, being straight and thick at the end. The acetabulum (hip socket in the middle) is large and relatively circular in shape. The pubic bone has lost its lower end and is well-fused to the ischial bone. The ischium is relatively thin, expanded, and the obturator process (a notably large projection at the end) is broken.^[1]

Classification

Nanshiungosaurus was in 1979 by Dong assigned to the Titanosaurinae, based on the assumption it was a sauropod genus, more specifically a titanosaurine (titanosaur).^[1] During this same year, the paleontologist Altangerel Perle described and named Segnosaurus also erecting the Segnosauridae to contain this strange taxon.^[10] Posterior to the findings of Nanshiungosaurus and Segnosaurus, more complete relatives started to be discovered, but their anatomical traits were so aberrant compared to other theropods to the point of being considered as Late Cretaceous sauropodomorph dinosaurs.^[11]^[12] In 1990, the paleontologists Rinchen Barsbold and Teresa Maryańska noted the striking similarities between the pelvises of Nanshiungosaurus and Segnosaurus, such as the ophisthopubic condition and large iliac blade. They concluded that the former was part of the Segnosauria and found this group to be a rare and aberrant group of saurischians in a position subject to change among sauropodomorphs and theropods.^[13] Dong also agreed with these similarities and placed the genus within the Segnosauridae in 1992.^[14] Nevertheles, the description of Alxasaurus in 1993 proved that the long-aberrant segnosaurs were tetanuran theropod dinosaurs and Segnosauridae was a junior synonym of the older name Therizinosauridae.^[15] Segnosauria also became synonymous with Therizinosauria.^[16] In 1997, with the description and naming of "N". bohlini, Dong and You placed the therizinosaurids Nanshiungosaurus and the former into a separate Nanshiungosauridae. They did not provide authentic traits or bases behind this new grouping.^[2] In 2010, Zanno performed one of the most complete analysis of the Therizinosauria at that point and noted that these two species do not pertain to the same genus and therefore the use of the Nanshiungosauridae was invalid and represents a synonym of Therizinosauridae.^[3]

Nanshiungosaurus features multiple therizinosaurid traits such as a ophistopubic pelvis, elongated iliac blade and an expanded obturator process. In her phylogenetic analysis, Zanno recovered this taxon as a derived therizinosaurid closer to Nothronychus and Segnosaurus.^[3] The therizinosaurid placement of Nanshiungosaurus has been widely followed and corroborated by most cladistic analyses.^[17]^[18] The extensive phylogenetic analysis conducted by Hartman and colleagues in 2019 based on Zanno's 2010 analysis, recovers Nanshiungosaurus in a more derived position than Neimongosaurus or Therizinosaurus. Below are the obtained results:^[18]

Therizinosauridae

Suzhousaurus

Neimongosaurus

	Therizinosaurus

	Erliansaurus

Nanchao embryos

Nanshiungosaurus

	Segnosaurus

	AMNH 6368

Nothronychus mckinleyi

Erlikosaurus

	"Nanshiungosaurus" bohlini

	Nothronychus graffami

Paleoecology

Nanshiungosaurus were some of the last therizinosaurs to exist. The holotype remains were unearthed from the Nanxiong Formation, which has been dated to the latest Maastrichtian stage about 66.7 ± 0.3 million years ago.^[19] This formation is mainly composed by purplish mudstones and siltstones, and was deposited in a floodplain environment, under a relatively warm, humid subtropical climate.^[20] Oviraptorid eggs are particularly common on the formation with atleast five well-preserved egg clutches.^[21] Moreover, a fossilized female oviraptorid with two eggs is known from the formation.^[22] The tracks of ornithopod dinosaurs are locally abundant at the Nanxiong region.^[23]

The therizinosaurids Nanshiungosaurus were bulky high-browsers in their ecosystem,^[6] and shared their habitat with multiple oviraptorosaur species such as Banji, Ganzhousaurus, Corythoraptor, Nankangia, Huanansaurus or Shixinggia.^[24]^[25] However, it is possible that some of these oviraptorosaurs did not actually live together given the poor stratigraphic analysis of the formation.^[24] Additional dinosaurs are represented by the fast running, long-snouted tyrannosaurids Qianzhousaurus,^[26] the sauropods Gannansaurus,^[27] and the very sparce remains of hadrosaurid dinosaurs such as Microhadrosaurus (now a nomen dubium).^[23] Other reptiles that composed the fauna were the terrestrial or semiaquatic nanhsiungchelyid turtles Nanhsiungchelys and Jiangxichelys, squamates Chianghsia and Tianyusaurus, and the crocodilians Jiangxisuchus.^[28]^[20]

References

^ ^a ^b ^c ^d ^e Dong, Z. (1979). "Cretaceous dinosaur fossils in southern China" [Cretaceous dinosaurs of the Huanan (south China)]. In Institute of Vertebrate Paleontology and Paleoanthropology; Nanjing Institute of Paleontology (eds.). Mesozoic and Cenozoic Redbeds in Southern China (in Chinese). Beijing: Science Press. pp. 342−350. Translated paper
^ ^a ^b Dong, Z.; You, H. (1997). "A new segnosaur from Mazhongshan Area, Gansu Province, China". In Dong, Z. M. (ed.). Sino-Japanese Silk Road Dinosaur Expedition. Beijing: China Ocean Press,. pp. 90−95.{{cite book}}: CS1 maint: extra punctuation (link)
^ ^a ^b ^c ^d Zanno, L. E. (2010). "A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora)". Journal of Systematic Palaeontology. 8 (4): 503−543. doi:10.1080/14772019.2010.488045.
^ You, H.; Morschhauser, E. M.; Li, D.; Dodson, P. (2018). "Introducing the Mazongshan Dinosaur Fauna". Journal of Vertebrate Paleontology. 38 (sup. 1): 1−11. doi:10.1080/02724634.2017.1396995.
^ Holtz, T. R.; Rey, L. V. (2007). Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages. Random House. ISBN 9780375824197.{{cite book}}: CS1 maint: date and year (link) Genus List for Holtz 2012 Weight Information
^ ^a ^b Paul, G. S. (2016). The Princeton Field Guide to Dinosaurs (2nd ed.). Princeton, New Jersey: Princeton University Press. pp. 162−168. ISBN 9780691167664.
^ Hedrick, B. P.; Zanno, L. E.; Wolfe, D. G.; Dodson, P. (2015). "The Slothful Claw: Osteology and Taphonomy of Nothronychus mckinleyi and N. graffami (Dinosauria: Theropoda) and Anatomical Considerations for Derived Therizinosaurids". PLOS ONE. 10 (6): e0129449. Bibcode:2015PLoSO..1029449H. doi:10.1371/journal.pone.0129449. PMC 4465624. PMID 26061728.
^ Lautenschlager, S.; Lawrence, M. W.; Perle, A.; Zanno, L. E.; Emily, J. R. (2014). "Cranial anatomy of Erlikosaurus andrewsi (Dinosauria, Therizinosauria): new insights based on digital reconstruction". Journal of Vertebrate Paleontology. 34 (6): 1263−1291. doi:10.1080/02724634.2014.874529.
^ Zanno, L. E.; Tsogtbaatar, K.; Chinzorig, T.; Gates, T. A. (2016). "Specializations of the mandibular anatomy and dentition of Segnosaurus galbinensis (Theropoda: Therizinosauria)". PeerJ. 4: e1885. doi:10.7717/peerj.1885. PMC 4824891. PMID 27069815.
^ Perle, A. (1979). "Segnosauridae — novoe semejstvo teropod iz pozdnego mela Mongolii" [Segnosauridae — a new family of theropods from the Late Cretaceous of Mongolia]. Transactions of the Joint Soviet-Mongolian Paleontological Expedition (in Russian). 8: 45−55. Translated paper
^ Gauthier, J. (1986). "Saurischian monophyly and the origin of birds". Memoirs of the California Academy of Sciences. 8: 45.
^ Sereno, P. C. (1989). "Prosauropod monophyly and basal sauropodomorph phylogeny". Abstract of Papers. Forty-Ninth Annual Meeting Society of Vertebrate Paleontology. Journal of Vertebrate Paleontology. Vol. 9, no. supp. 3. p. 39A. ISSN 0272-4634. JSTOR 4523276.
^ Barsbold, R.; Maryańska, T. (1990). "Saurischia Sedis Mutabilis: Segnosauria". In Weishampel, D. B.; Osmolska, H.; Dodson, P. (eds.). The Dinosauria (1st ed.). Berkeley: University of California Press. pp. 408−415. ISBN 9780520067271.
^ Dong, Z. (1992). Dinosaurian Faunas of China. Beijing: China Ocean Press. p. 187. ISBN 3-540-52084-8.
^ Russell, D. A.; Dong, Z. (1993). "The affinities of a new theropod from the Alxa Desert, Inner Mongolia, People's Republic of China". Canadian Journal of Earth Sciences. 30 (10): 2107−2127. Bibcode:1993CaJES..30.2107R. doi:10.1139/e93-183.
^ Russell, D. A. (1997). "Therizinosauria". In Currie, P. J.; Padian, K. (eds.). Encyclopedia of Dinosaurs. San Diego: Academic Press. pp. 729−730. ISBN 978-0-12-226810-6.
^ Pu, H.; Kobayashi, Y.; Lü, J.; Xu, L.; Wu, Y.; Chang, H.; Zhang, J.; Jia, S. (2013). "An Unusual Basal Therizinosaur Dinosaur with an Ornithischian Dental Arrangement from Northeastern China". PLOS ONE. 8 (5): e63423. doi:10.1371/journal.pone.0063423. PMC 3667168. PMID 23734177.
^ ^a ^b Hartman, S.; Mortimer, M.; Wahl, W. R.; Lomax, D. R.; Lippincott, J.; Lovelace, D. M. (2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ. 7: e7247. doi:10.7717/peerj.7247. PMC 6626525. PMID 31333906.
^ Buck, B. J.; Hanson, A. D.; Hengst, R. A.; Shu-sheng, H. (2004). "Tertiary Dinosaurs in the Nanxiong Basin, Southern China, Are Reworked from the Cretaceous". The Journal of Geology. 112 (1): 111−118. Bibcode:2004JG....112..111B. doi:10.1086/379695.
^ ^a ^b Tong, H.; Li, L. (2019). "A revision of the holotype of Nanhsiungchelys wuchingensis, Ye, 1966 (Testudines: Cryptodira: Trionychoidae: Nanhsiungchelyidae)". Cretaceous Research. 95: 151−163. doi:10.1016/j.cretres.2018.11.003. hdl:311034/9424.
^ Yang, T.-R.; Wiemann, J.; Xu, L.; Cheng, Y.-N.; Wu, X.-C.; Sander, P. M. (2019). "Reconstruction of oviraptorid clutches illuminates their unique nesting biology". Acta Palaeontologica Polonica. 466: 581−596. doi:10.4202/app.00497.2018.
^ Sato, T.; Cheng, Y.-N.; Wu, X.-C.; Zelenitsky, D. K.; Hsiao, Y.-F. (2005). "A Pair of Shelled Eggs Inside A Female Dinosaur". Science. 308 (5720): 375–375. doi:10.1126/science.1110578. PMID 15831749.
^ ^a ^b Xing, L.; Lockley, M. G.; Li, D.; Klein, H.; Ye, Y.; Scott Persons IV, W.; Ran, H. (2017). "Late Cretaceous ornithopod-dominated, theropod, and pterosaur track assemblages from the Nanxiong Basin, China: New discoveries, ichnotaxonomy, and paleoecology" (PDF). Palaeogeography, Palaeoclimatology, Palaeoecology. 466: 303−313. Bibcode:2017PPP...466..303X. doi:10.1016/j.palaeo.2016.11.035.
^ ^a ^b Lü, J.; Chen, R.; Brusatte, S. L.; Zhu, Y.; Shen, C. (2016). "A Late Cretaceous diversification of Asian oviraptorid dinosaurs: evidence from a new species preserved in an unusual posture". Scientific Reports. 6 (35780). Bibcode:2016NatSR...635780L. doi:10.1038/srep35780. PMC 5103654. PMID 27831542.
^ Lü, J.; Li, G.; Kundrát, M.; Lee, Y.-N.; Sun, Z.; Kobayashi, Y.; Shen, C.; Teng, F.; Liu, H. (2017). "High diversity of the Ganzhou Oviraptorid Fauna increased by a new cassowary-like crested species". Scientific Reports. 7 (6393). Bibcode:2017NatSR...7.6393L. doi:10.1038/s41598-017-05016-6. PMC 5532250. PMID 28751667.
^ Lü, J.; Yi, L.; Brusatte, S. L.; Yang, L.; Chen, L. (2014). "A new clade of Asian Late Cretaceous long-snouted tyrannosaurids". Nature Communications. 5 (3788). Bibcode:2014NatCo...5.3788L. doi:10.1038/ncomms4788. PMID 24807588.
^ Lü, J.; Yi, L.; Zhong, H.; Wei, X. (2013). "A New Somphospondylan Sauropod (Dinosauria, Titanosauriformes) from the Late Cretaceous of Ganzhou, Jiangxi Province of Southern China". Acta Geologica Sinica (English Edition). 87 (3): 678−685. doi:10.1111/1755-6724.12079.
^ Chun Li; Xiao-chun Wu; Scott Rufolo (2018). "A new crocodyloid (Eusuchia: Crocodylia) from the Upper Cretaceous of China". Cretaceous Research. 94: 25–39. doi:10.1016/j.cretres.2018.09.015.

[Dong1979-1] Dong, Z. (1979). "Cretaceous dinosaur fossils in southern China" [Cretaceous dinosaurs of the Huanan (south China)]. In Institute of Vertebrate Paleontology and Paleoanthropology; Nanjing Institute of Paleontology (eds.). Mesozoic and Cenozoic Redbeds in Southern China (in Chinese). Beijing: Science Press. pp. 342−350. Translated paper

[Dong1997-2] Dong, Z.; You, H. (1997). "A new segnosaur from Mazhongshan Area, Gansu Province, China". In Dong, Z. M. (ed.). Sino-Japanese Silk Road Dinosaur Expedition. Beijing: China Ocean Press,. pp. 90−95.{{cite book}}: CS1 maint: extra punctuation (link)

[Zanno2010-3] Zanno, L. E. (2010). "A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora)". Journal of Systematic Palaeontology. 8 (4): 503−543. doi:10.1080/14772019.2010.488045.

[You2018-4] You, H.; Morschhauser, E. M.; Li, D.; Dodson, P. (2018). "Introducing the Mazongshan Dinosaur Fauna". Journal of Vertebrate Paleontology. 38 (sup. 1): 1−11. doi:10.1080/02724634.2017.1396995.

[Holtz2012-5] Holtz, T. R.; Rey, L. V. (2007). Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages. Random House. ISBN 9780375824197.{{cite book}}: CS1 maint: date and year (link) Genus List for Holtz 2012 Weight Information

[Paul2016-6] Paul, G. S. (2016). The Princeton Field Guide to Dinosaurs (2nd ed.). Princeton, New Jersey: Princeton University Press. pp. 162−168. ISBN 9780691167664.

[7] Hedrick, B. P.; Zanno, L. E.; Wolfe, D. G.; Dodson, P. (2015). "The Slothful Claw: Osteology and Taphonomy of Nothronychus mckinleyi and N. graffami (Dinosauria: Theropoda) and Anatomical Considerations for Derived Therizinosaurids". PLOS ONE. 10 (6): e0129449. Bibcode:2015PLoSO..1029449H. doi:10.1371/journal.pone.0129449. PMC 4465624. PMID 26061728.

[8] Lautenschlager, S.; Lawrence, M. W.; Perle, A.; Zanno, L. E.; Emily, J. R. (2014). "Cranial anatomy of Erlikosaurus andrewsi (Dinosauria, Therizinosauria): new insights based on digital reconstruction". Journal of Vertebrate Paleontology. 34 (6): 1263−1291. doi:10.1080/02724634.2014.874529.

[9] Zanno, L. E.; Tsogtbaatar, K.; Chinzorig, T.; Gates, T. A. (2016). "Specializations of the mandibular anatomy and dentition of Segnosaurus galbinensis (Theropoda: Therizinosauria)". PeerJ. 4: e1885. doi:10.7717/peerj.1885. PMC 4824891. PMID 27069815.

[10] Perle, A. (1979). "Segnosauridae — novoe semejstvo teropod iz pozdnego mela Mongolii" [Segnosauridae — a new family of theropods from the Late Cretaceous of Mongolia]. Transactions of the Joint Soviet-Mongolian Paleontological Expedition (in Russian). 8: 45−55. Translated paper

[11] Gauthier, J. (1986). "Saurischian monophyly and the origin of birds". Memoirs of the California Academy of Sciences. 8: 45.

[12] Sereno, P. C. (1989). "Prosauropod monophyly and basal sauropodomorph phylogeny". Abstract of Papers. Forty-Ninth Annual Meeting Society of Vertebrate Paleontology. Journal of Vertebrate Paleontology. Vol. 9, no. supp. 3. p. 39A. ISSN 0272-4634. JSTOR 4523276.

[13] Barsbold, R.; Maryańska, T. (1990). "Saurischia Sedis Mutabilis: Segnosauria". In Weishampel, D. B.; Osmolska, H.; Dodson, P. (eds.). The Dinosauria (1st ed.). Berkeley: University of California Press. pp. 408−415. ISBN 9780520067271.

[14] Dong, Z. (1992). Dinosaurian Faunas of China. Beijing: China Ocean Press. p. 187. ISBN 3-540-52084-8.

[15] Russell, D. A.; Dong, Z. (1993). "The affinities of a new theropod from the Alxa Desert, Inner Mongolia, People's Republic of China". Canadian Journal of Earth Sciences. 30 (10): 2107−2127. Bibcode:1993CaJES..30.2107R. doi:10.1139/e93-183.

[16] Russell, D. A. (1997). "Therizinosauria". In Currie, P. J.; Padian, K. (eds.). Encyclopedia of Dinosaurs. San Diego: Academic Press. pp. 729−730. ISBN 978-0-12-226810-6.

[17] Pu, H.; Kobayashi, Y.; Lü, J.; Xu, L.; Wu, Y.; Chang, H.; Zhang, J.; Jia, S. (2013). "An Unusual Basal Therizinosaur Dinosaur with an Ornithischian Dental Arrangement from Northeastern China". PLOS ONE. 8 (5): e63423. doi:10.1371/journal.pone.0063423. PMC 3667168. PMID 23734177.

[Hartman2019-18] Hartman, S.; Mortimer, M.; Wahl, W. R.; Lomax, D. R.; Lippincott, J.; Lovelace, D. M. (2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ. 7: e7247. doi:10.7717/peerj.7247. PMC 6626525. PMID 31333906.

[19] Buck, B. J.; Hanson, A. D.; Hengst, R. A.; Shu-sheng, H. (2004). "Tertiary Dinosaurs in the Nanxiong Basin, Southern China, Are Reworked from the Cretaceous". The Journal of Geology. 112 (1): 111−118. Bibcode:2004JG....112..111B. doi:10.1086/379695.

[Tong2019N-20] Tong, H.; Li, L. (2019). "A revision of the holotype of Nanhsiungchelys wuchingensis, Ye, 1966 (Testudines: Cryptodira: Trionychoidae: Nanhsiungchelyidae)". Cretaceous Research. 95: 151−163. doi:10.1016/j.cretres.2018.11.003. hdl:311034/9424.

[21] Yang, T.-R.; Wiemann, J.; Xu, L.; Cheng, Y.-N.; Wu, X.-C.; Sander, P. M. (2019). "Reconstruction of oviraptorid clutches illuminates their unique nesting biology". Acta Palaeontologica Polonica. 466: 581−596. doi:10.4202/app.00497.2018.

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@@ Line 2: / Line 2: @@
 | fossil_range = [[Late Cretaceous]], <br>~{{fossilrange|67|66}}
 | image = Nanshiungosaurus.jpg
-| image_caption = Skeletal restoration
+| image_caption = Skeletal restoration of IVPP V4731
 | display_parents = 2
 | taxon = Nanshiungosaurus
 | authority = [[Dong Zhiming|Dong]], [[1979 in paleontology|1979]]
+| type_species = {{extinct}}'''''Nanshiungosaurus brevispinus'''''
-| subdivision_ranks = Species
+| type_species_authority = Dong, 1979
-| subdivision =
-*[[extinction|†]]'''''N. brevispinus''''' <small>Dong, 1979 ([[type species|type]])</small>
-*?†'''''"N". bohlini''''' <small>Dong & You, 1997</small>
 }}
-'''''Nanshiungosaurus''''' (meaning "Nanxiong's lizard") is a [[genus]] of [[therizinosaurid]] [[theropod]] [[dinosaurs]] from the [[Late Cretaceous]] of [[People's Republic of China|China]]. The [[type species]], ''Nanshiungosaurus brevispinus'', was first discovered in 1974 and described in 1979, represented by a single specimen preserving most of the cervical and dorsal vertebrae with the pelvis. A supposed, dubious second species, ''"Nanshiungosaurus" bohlini'', was found in 1992 and described in 1997. It also preserves vertebrae but this species however, have a large age difference compared to the later and issues regarding authentic characteristics, making its affinity to the genus questionable.
+'''''Nanshiungosaurus''''' (meaning "Nanxiong's lizard") is a [[genus]] of [[therizinosaurid]] [[dinosaurs]] that lived in what is know [[Asia]] during the [[Late Cretaceous]] of [[South China]]. The [[type species]], '''''Nanshiungosaurus brevispinus''''', was first discovered in 1974 and described in 1979 by [[Dong Zhiming]]. It is represented by a single specimen preserving most of the cervical and dorsal vertebrae with the pelvis. A supposed and unlikely second species, ''"Nanshiungosaurus" bohlini'', was found in 1992 and described in 1997. It is also represented by vertebrae but this species however, differs in geological age and authentic characteristics compared to the later, making its affinity to the genus unsupported.
+They were large-sized therizinosaurids that are estimated to have cover {{convert|5|m|ft|abbr=on}} in length and to have weighed about {{convert|907|kg|lb|abbr=on}}. ''Nanshiungosaurus'' had a very pneumatized [[vertebral column]] with the posterior [[cervical vertebrae]] of the long neck being unsually robust and slightly more elongated than the [[Thoracic vertebrae|dorsals]]. They also were equipped with broad torsos as seen on the bulky [[pelvis]]. As for other therizinosaurids, they had a [[Beak#Rhamphotheca|keratinous beak]] used when feeding, stocky feet with four weight-bearing [[toes]] and large flattened claws.
-==Discovery and naming==
-[[File:Therizinosauroids illium comparison.png|thumb|left|Schematic comparison of the illium of ''Nanshiungosaurus'' (in E) and other related therizinosaurs]]
-The first remains of ''Nanshiungosaurus'' were collected in 1974 by an expedition team of the [[Institute of Vertebrate Paleontology and Paleoanthropology]]. Later on, in 1979 ''Nanshiungosaurus brevispinus'' was named and described by [[Dong Zhiming]]. The generic name refers to [[Nanxiong]] and the [[specific name (zoology)|specific name]] is derived from [[Latin]] ''brevis'', "short", and ''spina'', "thorn", in reference to the short vertebral spines. The [[holotype]] specimen, '''IVPP V4731''', was near Dapingcun in [[Guangdong]] found in a layer of the [[Nanxiong Formation]], dating from the [[Late Cretaceous]] [[Campanian]]. It consists of a partial skeleton lacking the skull but including 12 [[Glossary of dinosaur anatomy#cervicals|cervical]], 10 [[Glossary of dinosaur anatomy#dorsals|dorsal]] and 6 [[Glossary of dinosaur anatomy#sacrals|sacral vertebrae]], and a bulky [[pelvis]]. It was first thought to be a small, strange [[sauropod]] characterised by a shorter but thicker neck than other sauropods.<ref name=Dong1979>{{cite book|last1=Dong|first1=Z.|date=1979|title=Mesozoic and Cenozoic Redbeds in Southern China|chapter=Cretaceous dinosaur fossils in southern China|trans-chapter=Cretaceous dinosaurs of the Huanan (south China)|editor1=Institute of Vertebrate Paleontology and Paleoanthropology|editor2=Nanjing Institute of Paleontology|publisher=Science Press|location=Beijing|pages=342–350|language=Chinese}} [https://paleoglot.org/files/Dong_79.pdf Translated paper]</ref> The Nanxiong Formation has been dated to the [[Maastrichtian]] stage about 66.7 ± 0.3 [[million years ago]].<ref>{{cite journal|last1=Buck|first1=B. J.|last2=Hanson|first2=A. D.|last3=Hengst|first3=R. A.|last4=Shu-sheng|first4=H.|date=2004|title=Tertiary Dinosaurs in the Nanxiong Basin, Southern China, Are Reworked from the Cretaceous|journal=The Journal of Geology|volume=112|issue=1|pages=111–118|doi=10.1086/379695}}</ref>
+''Nanshiungosaurus'' are classified as therizinosaurian dinosaurs. Along with ''[[Therizinosaurus]]'' and ''[[Segnosaurus]]'', ''Nanshiungosaurus'' was one the earliest therizinosaurs to be described and named. The unusual shape of the pelvis led Dong to interpret the remains as belonging to some dwarf sauropod, but during the 1990s the genus was recognized as a segnosaur (now therizinosaur) based the pelvic similarities with ''Segnosaurus''. At first, the research on therizinosaurs was complicated since sparce remains were known at the time and these had features from multiple dinosaur lineages leading to their interpretation as [[prosauropod]] dinosaurs.
-In 1997 Dong Zhiming and You Hailu named and described a second species: "''Nanshiungosaurus" bohlini'', based on a skeleton found in 1992 near [[Mazongshan]]. It consists of only 11 cervical and 5 dorsal vertebrae with some [[ribs]]. The holotype is '''IVPP V11116''', from the [[Early Cretaceous]] Upper [[Xinminbao Group]] dating from the [[Barremian]]-[[Aptian]]. The specific name honours the Swedish paleontologist [[Birger Bohlin]]. Dong and Yu presented no evidence or argumentation supporting the assignment of the species to ''Nanshiungosaurus''.<ref name=Dong1997>{{cite book|last1=Dong|first1=Z.|last2=Yu|first2=H.|date=1997|title=Sino-Japanese Silk Road Dinosaur Expedition|chapter=A new segnosaur from Mazhongshan Area, Gansu Province, China|editor-last1=Dong|editor-first1=Z. M.|publisher=China Ocean Press,|location=Beijing|pages=90–95}}</ref> In view of the lack of [[synapomorphies]], shared unique traits, and the large age difference with ''Nanshiungosaurus brevispinus'', it is generally considered that the supposed second species is unrelated to the later and might warrant its own genus.<ref name=Zanno2010/><ref name=Paul2016>{{cite book|last1=Paul|first1=G. S.|title=The Princeton Field Guide to Dinosaurs|publisher=Princeton University Press|year=2016|isbn=9780691167664|edition=2nd|location=Princeton, New Jersey|pages=162−168}}</ref><ref name=Hartman2019>{{cite journal|last1=Hartman|first1=S.|last2=Mortimer|first2=M.|last3=Wahl|first3=W. R.|last4=Lomax|first4=D. R.|last5=Lippincott|first5=J.|last6=Lovelace|first6=D. M.|title=A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight|journal=PeerJ|date=2019|volume=7|pages=e7247|doi=10.7717/peerj.7247|doi-access=free|pmid=31333906|pmc=6626525}}</ref>
+==History of discovery==
+[[File:Fossiliferous basins of China.png|thumb|left|[[Fossil]]iferous basins in China. The holotype of ''Nanshiungosaurus'' was discovered at the Nanxiong Basin]]
+In 1974, during a geological expedition at the [[Nanxiong Basin]] led by the [[Institute of Vertebrate Paleontology and Paleoanthropology]] several fossilized remains of dinosaurs were discovered by the team. Near the village of [[Dapingcun]] at the [[Nanxiong Formation]], [[Guangdong Province]], a relatively large and partial skeleton was found in articulation dating back to the [[Late Cretaceous]]. The specimen was labelled under the number '''IVPP V4731''' and consisted of 12 [[Glossary of dinosaur anatomy#cervicals|cervical]] (lacking the [[Atlas (anatomy)|atlas]]), 10 [[Glossary of dinosaur anatomy#dorsals|dorsal]], 5 (actually 6) [[Glossary of dinosaur anatomy#sacrals|sacral]] and the first [[Glossary of dinosaur anatomy#caudals|caudal vertebrae]] with a nearly complete, bulky [[pelvis]] only lacking the right [[ilium]] and [[ischium]]. Later on, in 1979 the specimen was formally described by the [[China|Chinese]] [[paleontologist]] [[Dong Zhiming]] and used as the [[Holotype|basis]] for the new genus and species ''Nanshiungosaurus brevispinus''. The [[Genus#Use|generic name]], ''Nanshiungosaurus'', refers to the site of provenance [[Nanxiong]] city and is derived from the [[Greek language|Greek]] σαῦρος (sauros, meaning lizard). Lastly, the [[specific name (zoology)|specific name]], ''brevispinus'', is derived from the [[Latin]] brevis and spina (meaning short and spine, respectively) in reference to the relatively short vertebral spines. When first described, Dong mistakenly thought the specimen to have been a dwarf, strange [[titanosaurine]] [[sauropod]] characterized by a shorter but thicker neck than other sauropods based on the pelvis structure.<ref name=Dong1979>{{cite book|last1=Dong|first1=Z.|date=1979|title=Mesozoic and Cenozoic Redbeds in Southern China|chapter=Cretaceous dinosaur fossils in southern China|trans-chapter=Cretaceous dinosaurs of the Huanan (south China)|editor1=Institute of Vertebrate Paleontology and Paleoanthropology|editor2=Nanjing Institute of Paleontology|publisher=Science Press|location=Beijing|pages=342−350|language=Chinese}} [https://paleoglot.org/files/Dong_79.pdf Translated paper]</ref>
+In 1997 Dong Zhiming and You Hailu named and described a supposed second species: "''Nanshiungosaurus" bohlini'', based on a skeleton found in 1992 near [[Mazongshan]]. It consists of 11 cervical and 5 dorsal vertebrae with some [[ribs]]. The specimen is catalogued as IVPP V11116 coming from the [[Early Cretaceous]], Upper [[Xinminbao Group]]. Also, they coined the Nanshiungosauridae to contain "both species". Dong and Yu presented no evidence or argumentation supporting the assignment of the species to ''Nanshiungosaurus''.<ref name=Dong1997>{{cite book|last1=Dong|first1=Z.|last2=You|first2=H.|date=1997|title=Sino-Japanese Silk Road Dinosaur Expedition|chapter=A new segnosaur from Mazhongshan Area, Gansu Province, China|editor-last1=Dong|editor-first1=Z. M.|publisher=China Ocean Press,|location=Beijing|pages=90−95}}</ref> In 2010, the North American paleontologist [[Lindsay Zanno]] considered this referral to be highly improbable since ''"N". bohlini'' dates back from the [[Barremian]]-[[Aptian]] ages and in the view of lack of [[synapomorphies]], she considered that the supposed second species is unrelated to ''Nanshiungosaurus'' and might warrant its own genus. In addition, she corrected the number of sacral vertebrae from 5 to 6 and noted that the holotype pelvis from the latter has suffered damage since collection and has been reconstructed with painted plaster in the affected areas.<ref name=Zanno2010>{{cite journal|first1=L. E.|last1=Zanno|date=2010|title=A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora)|journal=Journal of Systematic Palaeontology|volume=8|issue=4|pages=503−543|doi=10.1080/14772019.2010.488045|url=https://www.researchgate.net/publication/229289268}}</ref> Moreover, the remains of ''"N". bohlini'' were recovered from the lower red beds of the [[Zhonggou Formation]] in the [[Gongpoquan Basin]], a completely different geological context. As an overall consensus, this dubious specimen is no longer considered to be relatable to ''Nanshiungosaurus''.<ref name=You2018>{{cite journal|last1=You|first1=H.|last2=Morschhauser|first2=E. M.|last3=Li|first3=D.|last4=Dodson|first4=P.|date=2018|title=Introducing the Mazongshan Dinosaur Fauna|journal=Journal of Vertebrate Paleontology|volume=38|issue=sup. 1|pages=1−11|doi=10.1080/02724634.2017.1396995}}</ref>
 ==Description==
-[[File:Nanshiungosaurus scale.png|thumb|left|Size compared to a human]]
+[[File:Nanshiungosaurus scale.png|thumb|Size compared to a human]]
+''Nanshiungosaurus'' were relatively large-bodied [[therizinosaurids]], estimated at {{convert|5|m|ft|abbr=on}} long weighing {{convert|907|kg|lb|abbr=on}}.<ref name=Holtz2012>{{cite book|last1=Holtz|first1=T. R.|title=Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages|last2=Rey|first2=L. V.|date=2007|publisher=Random House|year=2007|isbn=9780375824197}} [https://www.geol.umd.edu/~tholtz/dinoappendix/HoltzappendixWinter2011.pdf Genus List for Holtz 2012]  [https://www.geol.umd.edu/~tholtz/dinoappendix/appendix.html Weight Information]</ref><ref name=Paul2016>{{cite book|last1=Paul|first1=G. S.|title=The Princeton Field Guide to Dinosaurs|publisher=Princeton University Press|year=2016|isbn=9780691167664|edition=2nd|location=Princeton, New Jersey|pages=162−168}}</ref> This species can be differentiated by the possession of stocky posterior-most cervical vertebrae with [[Vertebral column#Variations in vertebrae|opisthocoel]] (meaning that they were [[wikt:Special:Search/concave|concave]] on their posterior sides) [[Vertebra#Structure|centra]].<ref name=Zanno2010/> Like other derived (advanced) therizinosaurids, ''Nanshiungosaurus'' were pot-bellied animals that had a strong build composed of stout hindlimbs with a functionally [[Dactyly#Tetradactyly|tretadactyl]] [[Pes (anatomy)|pes]]. Their arms endend up in large, recurved [[claws]] that were side to side flattened.<ref>{{cite journal|last1=Hedrick|first1=B. P.|last2=Zanno|first2=L. E.|last3=Wolfe|first3=D. G.|last4=Dodson|first4=P.|date=2015|title=The Slothful Claw: Osteology and Taphonomy of Nothronychus mckinleyi and N. graffami (Dinosauria: Theropoda) and Anatomical Considerations for Derived Therizinosaurids|journal=PLOS ONE|volume=10|issue=6|pages=e0129449|bibcode=2015PLoSO..1029449H|doi=10.1371/journal.pone.0129449|doi-access=free|pmc=4465624|pmid=26061728}}</ref> Though the holotype specimen lacks cranial material, the preserved elements in therizinosaurids ''[[Erlikosaurus]]'' and ''[[Segnosaurus]]'' indicate that they had relatively small skulls with coarsely [[Denticle (tooth feature)|serrated]] [[teeth]], and developed a prominent [[rhamphotheca]] (keratinous beak).<ref>{{cite journal|last1=Lautenschlager|first1=S.|last2=Lawrence|first2=M. W.|last3=Perle|first3=A.|last4=Zanno|first4=L. E.|last5=Emily|first5=J. R.|date=2014|title=Cranial anatomy of Erlikosaurus andrewsi (Dinosauria, Therizinosauria): new insights based on digital reconstruction|journal=Journal of Vertebrate Paleontology|volume=34|issue=6|pages=1263−1291|doi=10.1080/02724634.2014.874529|doi-access=free}}</ref><ref>{{cite journal|last1=Zanno|first1=L. E.|last2=Tsogtbaatar|first2=K.|last3=Chinzorig|first3=T.|last4=Gates|first4=T. A.|date=2016|title=Specializations of the mandibular anatomy and dentition of Segnosaurus galbinensis (Theropoda: Therizinosauria)|journal=PeerJ|volume=4|page=e1885|doi=10.7717/peerj.1885|doi-access=free|pmc=4824891|pmid=27069815}}</ref>
-[[File:Nanshiungosaurus dinosaur.png|thumb|[[Paleoart|Life restoration]]]]
-''Nanshiungosaurus'' were relatively large-bodied therizinosaurids, estimated at {{convert|4.4|-|5|m|ft|abbr=on}} long weighing {{convert|907|kg|lb|abbr=on}}.<ref name=Holtz2012>{{cite book|last1=Holtz|first1=T. R.|title=Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages|last2=Rey|first2=L. V.|date=2007|publisher=Random House|year=2007|isbn=9780375824197}} [https://www.geol.umd.edu/~tholtz/dinoappendix/HoltzappendixWinter2011.pdf Genus List for Holtz 2012]  [https://www.geol.umd.edu/~tholtz/dinoappendix/appendix.html Weight Information]</ref><ref name=Paul2016/> ''"Nanshiungosaurus" bohlini'' were slightly larger, estimated by [[Gregory S. Paul|Paul]] at {{convert|6|m|ft|abbr=on}} and {{convert|1.3|t|kg|abbr=on}}.<ref name=Paul2016/> The number of sacral vertebrae was first determined at five,<ref name=Dong1979/> later corrected to six.<ref name=Zanno2010>{{cite journal|first1=L. E.|last1=Zanno|year=2010|title=A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora)|journal=Journal of Systematic Palaeontology|volume=8|issue=4|pages=503–543|doi=10.1080/14772019.2010.488045|url=https://www.researchgate.net/publication/229289268}}</ref> ''N. brevispinus'' distinguishes itself by the possession of twelve, stocky cervical vertebrae with [[Vertebral column#Variations in vertebrae|opisthocoel]] (meaning that they were [[wikt:Special:Search/concave|concave]] on their posterior sides) [[Vertebra#Structure|centra]].<ref name=Zanno2010/> Most cervicals have relatively short [[neural arches]], but in the dorsals they are more elongated.<ref name=Dong1979/>
+===Vertebral column===
-The pelvis is represented by a well-preserved [[sacrum]] and left side composed by the [[illium]], [[Pubis (bone)#Dinosaurs|pubis]] and [[ischium]]; the pubis has lost its distal end though. The right side only preserves the ischium. As in most therizinosaurs, the pubis and ischium are fused into a solid structure. As a whole, the pelvis is robustly built. The sacral vertebrae are fused, indicating that the individual was an adult at the time of death.<ref name=Dong1979/>
+[[File:Nanshiungosaurus dinosaur.png|thumb|left|[[Paleoart|Life restoration]]]]
+Most cervicals have relatively short [[neural arches]], but in the dorsals they are more elongated. The [[Axis (anatomy)|axis]] is well preserved and has a length of {{convert|13.5|cm|mm|abbr=on}} with [[Vertebral column#Variations in vertebrae|platycoelus]] (slightly concave at both ends) centra. Its [[Axis (anatomy)#Dens|odontoid process]] is fused with the anterior facet of centrum and has a blunt depression to the bottom area in order to articulate with the [[Atlas (anatomy)|atlas]] centrum. The neural arch is narrow, the neural spine has a porterior orientation, and has rounded facets that connect with the atlantal posterior [[articular processes]] (bony projections). After the axis, the next cervicals gradually increase in size. The anterior cervicals have platycoelus centra and deep, pocket-like [[Glossary of dinosaur anatomy#P|pleurocoels]] (small holes) develop on the lateral sides; a small ridge-like structure is also present. On the bottom surface, the centra are flat, gently flattened to the inner sides, and two projections originate from the borders of the centra. The anterior capitular process is compressed (flattened) with some degree of thickness, and extends from the bottom to lateral sides to be fused with the [[Glossary of dinosaur anatomy#R|capitulum]] on the [[ribs]]. As for most cervicals, the neural arches and spines are short, the anterior articular processes are strongly developed in that direction, and the anterior lobes of the diapophyses (bony projections to the lateral sides) extend to the bottom and lateral directions to articulate with tubercles. Seventh and eighth cervicals are the longest with {{convert|18|cm|mm|abbr=on}} long centra and posterior to them, the vertebral size slightly narrows. In most posterior cervicals, the centra are opisthocoelous with shallow posterior grooves.<ref name=Dong1979/>
+The [[dorsal vertebrae]] centra, unlike in the cervicals, are narrow and platycoelous with an average length of {{convert|7|cm|mm|abbr=on}}. They are also [[Skeletal pneumaticity|pneumatic]] with shallow [[pneumatopore]]s (pits leading to air pockets) on the lateral sides, but they reduce in size on posterior dorsals. Along with the centra, the neural arch are well pneumatized with most of the pneumatic areas filled with matrix (sediments). The capitular processes are undeveloped and replaced by poor rounded facets on the anterior surface of the centrum. In most dorsals, the [[neural spine]]s are very elongated, the articular processes are located within the ligaments of the centrum, the neural spines are stocky, side to side thickened, and the top regions have stout tuberosities. A prominent spiral-like ridge is present on the bottom surface that becomes more notable to the posterior direction. Though most dorsals are completely preserved, eighth through tenth dorsals have damaged centra. On the posterior series the centra become platycoelous, approaching the sacral vertebrae structure. The [[sacrum]] is composed by six (originally five) [[Glossary of dinosaur anatomy#sacrals|sacral vertebrae]] and they are fused into a single structure, indicating that the individual was an adult at the time of death. They are lightly more longer than the dorsals and have stocky, side to ide broadened neural spines with the top surfaces bearing small depressions. A single [[Glossary of dinosaur anatomy#caudals|caudal vertebra]] is present after the sacral series and represents the first caudal. Its neural spine is broken and the centrum is slightly [[Vertebral column#Variations in vertebrae|amphicoelous]] (concave on both ends) with sub-equal height and length.<ref name=Dong1979/>
+===Pelvic girdle===
+[[File:Therizinosauroids illium comparison.png|thumb|Schematic comparison of the illium of ''Nanshiungosaurus'' (in E) compared to other therizinosaurs]]
+The pelvis is represented by the well-preserved left side composed by the [[illium]], [[Pubis (bone)#Dinosaurs|pubis]] and both [[Ischium|ischia]]; the right ilium and pubis were eroded though. As a whole, the pelvis is robustly built and some elements are deformed such as the left ilium, which due to [[taphonomical]] factors has been bent out of shape. Like other derived therizinosaurids, the pelvis has a [[Pubis (bone)#Dinosaurs|opisthopubic]] condition where the pubis and ischium are fused and directed backwards. The ilium is stocky with an extremely well-developed and elongated preacetabular process (anterior expansion of the iliac blade), nevertheless, the postacetabular process (posterior expansion) is missing. Its pubic peduncle (a robust projection that connects to the pubis) is also well-developed, being straight and thick at the end. The [[acetabulum]] (hip socket in the middle) is large and relatively circular in shape. The [[Pubis (bone)#Dinosaurs|pubic bone]] has lost its lower end and is well-fused to the ischial bone. The [[ischium]] is relatively thin, expanded, and the obturator process (a notably large projection at the end) is broken.<ref name=Dong1979/>
 ==Classification==
+''Nanshiungosaurus'' was in 1979 by Dong assigned to the [[Titanosaurinae]], based on the assumption it was a sauropod genus, more specifically a titanosaurine ([[titanosaur]]).<ref name=Dong1979/> During this same year, the paleontologist [[Altangerel Perle]] described and named ''[[Segnosaurus]]'' also erecting the Segnosauridae to contain this strange taxon.<ref>{{cite journal|last1=Perle|first1=A.|title=Segnosauridae — novoe semejstvo teropod iz pozdnego mela Mongolii|trans-title=Segnosauridae — a new family of theropods from the Late Cretaceous of Mongolia|journal=Transactions of the Joint Soviet-Mongolian Paleontological Expedition|date=1979|volume=8|pages=45−55|language=Russian}} [http://paleoglot.org/files/Perle%2079.pdf Translated paper]</ref> Posterior to the findings of ''Nanshiungosaurus'' and ''Segnosaurus'', more complete relatives started to be discovered, but their anatomical traits were so aberrant compared to other theropods to the point of being considered as [[Late Cretaceous]] [[sauropodomorph]] dinosaurs.<ref>{{cite journal|last1=Gauthier|first1=J.|title=Saurischian monophyly and the origin of birds|journal=Memoirs of the California Academy of Sciences|date=1986|volume=8|page=45|url=https://www.biodiversitylibrary.org/page/15651737#page/331/mode/1up}}</ref><ref>{{cite conference|last1=Sereno|first1=P. C.|title=Prosauropod monophyly and basal sauropodomorph phylogeny|book-title=Abstract of Papers. Forty-Ninth Annual Meeting Society of Vertebrate Paleontology|journal=Journal of Vertebrate Paleontology|date=1989|volume=9|issue=supp. 3|page=39A|jstor=4523276|issn=0272-4634}}</ref> In 1990, the paleontologists [[Rinchen Barsbold]] and [[Teresa Maryańska]] noted the striking similarities between the pelvises of ''Nanshiungosaurus'' and ''Segnosaurus'', such as the ophisthopubic condition and large iliac blade. They concluded that the former was part of the Segnosauria and found this group to be a rare and aberrant group of saurischians in a position subject to change among sauropodomorphs and theropods.<ref>{{cite book|last1=Barsbold|first1=R.|last2=Maryańska|first2=T.|chapter=Saurischia Sedis Mutabilis: Segnosauria|editor1-last=Weishampel|editor1-first=D. B.|editor2-last=Osmolska|editor2-first=H.|editor3-last=Dodson|editor3-first=P.|year=1990|title=The Dinosauria|edition=1st|publisher=University of California Press|location=Berkeley|isbn=9780520067271|pages=408−415}}</ref> Dong also agreed with these similarities and placed the genus within the Segnosauridae in 1992.<ref>{{cite book|last1=Dong|first1=Z.|year=1992|title=Dinosaurian Faunas of China|publisher=China Ocean Press|location=Beijing|pages=187|isbn=3-540-52084-8}}</ref> Nevertheles, the description of ''[[Alxasaurus]]'' in 1993 proved that the long-aberrant segnosaurs were [[tetanuran theropod]] dinosaurs and Segnosauridae was a [[junior synonym]] of the older name [[Therizinosauridae]].<ref>{{cite journal|last1=Russell|first1=D. A.|last2=Dong|first2=Z.|date=1993|title=The affinities of a new theropod from the Alxa Desert, Inner Mongolia, People's Republic of China|journal=Canadian Journal of Earth Sciences|volume=30|issue=10|pages=2107−2127|doi=10.1139/e93-183|bibcode=1993CaJES..30.2107R}}</ref> Segnosauria also became synonymous with [[Therizinosauria]].<ref>{{cite book|last1=Russell|first=D. A.|year=1997|chapter=Therizinosauria|editor-last1=Currie|editor-first1=P. J.|editor-last2=Padian|editor-first2=K.|title=Encyclopedia of Dinosaurs|publisher=Academic Press|location=San Diego|pages=729−730|isbn=978-0-12-226810-6}}</ref> In 1997, with the description and naming of ''"N". bohlini'', Dong and You placed the therizinosaurids ''Nanshiungosaurus'' and the former into a separate Nanshiungosauridae. They did not provide authentic traits or bases behind this new grouping.<ref name=Dong1997/> In 2010, Zanno performed one of the most complete analysis of the Therizinosauria at that point and noted that these two species do not pertain to the same genus and therefore the use of the Nanshiungosauridae was invalid and represents a synonym of Therizinosauridae.<ref name=Zanno2010/>
-''Nanshiungosaurus'' was in 1979 by Dong assigned to the [[Titanosaurinae]], based on the assumption it were a sauropod.<ref name=Dong1979/> In 1992 the species was assigned to the [[Segnosauridae]], the group today called [[Therizinosauridae]].<ref>{{cite book|last1=Dong|first1=Z.|year=1992|title=Dinosaurian Faunas of China|publisher=China Ocean Press|location=Beijing|pages=187|isbn=3-540-52084-8}}</ref> In 1997 however, Dong placed ''N. brevispinus'' and ''"N". bohlini'' in a separate Nanshungosauridae (now obsolete).<ref name=Dong1997/> [[Cladistic]] analyses indicate a position for ''Nanshiungosaurus'' that is either basal in the Therizinosauridae<ref name=Zanno2010/><ref>{{cite journal|last1=Zanno|first1=L. E.|last2=Gillete|first2=D. D.|last3=Albright|first3=L. B.|last4=Titus|first4=A. L.|date=2009|title=A new North American therizinosaurid and the role of herbivory in ‘predatory’ dinosaur evolution|journal=Proceedings of the Royal Society B|volume=276|issue=1672|pages=3505–3511|doi=10.1098/rspb.2009.1029|pmc=2817200|pmid=19605396}}</ref> or just outside that clade, basal in the [[Therizinosauroidea]].<ref>{{cite journal|last1=Senter|first1=P.|date=2007|title=A new look at the phylogeny of coelurosauria (Dinosauria: Theropoda)|journal=Journal of Systematic Palaeontology|volume=5|issue=4|pages=429–463|doi=10.1017/S1477201907002143}}</ref> Under the definition of Therizinosauridae ''sensu'' [[Xu Xing (paleontologist)|Xu]] 2002, ''Nanshuingosaurus'' is by definition a therizinosaurid.<ref>{{cite journal|last1=Xu|first1=X.|last2=Zhang|first2=Z.-H.|last3=Sereno|first3=P. C.|last4=Zhao|first4=X.-J.|last5=Kuang|first5=X.-W.|last6=Han|first6=J.|last7=Tan|first7=L.|date=2002|title=A new therizinosauroid (Dinosauria, Theropoda) from the Upper Cretaceous Iren Dabasu Formation of Nei Mongol|journal=Vertebrata PalAsiatica|volume=40|pages=228–240|url=http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/200901/W020090813371861636279.pdf}}</ref> The phylogenetic analysis conducted by Hartaman et al. 2019 recovers ''Nanshiungosaurus'' as a therizinosaurid in a more derived position than ''[[Erliansaurus]]'' or ''[[Neimongosaurus]]''. Below are the obtained results:<ref name=Hartman2019/>
-{{clade|style=font-size:90%; line-height:90%;
+''Nanshiungosaurus'' features multiple therizinosaurid traits such as a ophistopubic pelvis, elongated iliac blade and an expanded obturator process. In her [[phylogenetic]] analysis, Zanno recovered this taxon as a derived therizinosaurid closer to ''[[Nothronychus]]'' and ''Segnosaurus''.<ref name=Zanno2010/> The therizinosaurid placement of ''Nanshiungosaurus'' has been widely followed and corroborated by most cladistic analyses.<ref>{{cite journal|last1=Pu|first1=H.|last2=Kobayashi|first2=Y.|last3=Lü|first3=J.|last4=Xu|first4=L.|last5=Wu|first5=Y.|last6=Chang|first6=H.|last7=Zhang|first7=J.|last8=Jia|first8=S.|date=2013|title=An Unusual Basal Therizinosaur Dinosaur with an Ornithischian Dental Arrangement from Northeastern China|journal=PLOS ONE|volume=8|issue=5|pages=e63423|doi=10.1371/journal.pone.0063423|doi-access=free|pmc=3667168|pmid=23734177}}</ref><ref name=Hartman2019>{{cite journal|last1=Hartman|first1=S.|last2=Mortimer|first2=M.|last3=Wahl|first3=W. R.|last4=Lomax|first4=D. R.|last5=Lippincott|first5=J.|last6=Lovelace|first6=D. M.|title=A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight|journal=PeerJ|date=2019|volume=7|pages=e7247|doi=10.7717/peerj.7247|doi-access=free|pmid=31333906|pmc=6626525}}</ref> The extensive phylogenetic analysis conducted by Hartman and colleagues in 2019 based on Zanno's 2010 analysis, recovers ''Nanshiungosaurus'' in a more derived position than ''[[Neimongosaurus]]'' or ''[[Therizinosaurus]]''. Below are the obtained results:<ref name=Hartman2019/>
-|label1=[[Therizinosauridae]]
+[[File:Therizinosauria hips variation.JPG|thumb|Hip variation among therizinosaur genera (''Nanshiungosaurus'' right corner)]]
+{{clade|style=font-size:100%; line-height:100%;
+|label1='''[[Therizinosauridae]]'''
 |1={{clade
 |1={{clade
@@ Line 42: / Line 54: @@
 |1=[[Nanchao Formation|Nanchao]] embryos
 |2={{clade
-|1='''''Nanshiungosaurus brevispinus'''''
+|1='''''Nanshiungosaurus'''''
 |2={{clade
 |1={{clade
@@ Line 53: / Line 65: @@
 |1=''[[Erlikosaurus]]''
 |2={{clade
-|1='''''"Nanshiungosaurus" bohlini'''''
+|1=''"Nanshiungosaurus" bohlini''
 |2=''[[Nothronychus graffami]]'' }} }} }} }} }} }} }} }} }}
+==Paleoecology==
+''Nanshiungosaurus'' were some of the last therizinosaurs to exist. The holotype remains were unearthed from the [[Nanxiong Formation]], which has been dated to the latest [[Maastrichtian]] stage about 66.7 ± 0.3 [[million years ago]].<ref>{{cite journal|last1=Buck|first1=B. J.|last2=Hanson|first2=A. D.|last3=Hengst|first3=R. A.|last4=Shu-sheng|first4=H.|date=2004|title=Tertiary Dinosaurs in the Nanxiong Basin, Southern China, Are Reworked from the Cretaceous|journal=The Journal of Geology|volume=112|issue=1|pages=111−118|bibcode=2004JG....112..111B|doi=10.1086/379695|url=https://www.academia.edu/476028}}</ref> This formation is mainly composed by purplish [[mudstones]] and [[siltstones]], and was deposited in a [[floodplain]] environment, under a relatively warm, [[humid subtropical climate]].<ref name=Tong2019N>{{cite journal|last1=Tong|first1=H.|last2=Li|first2=L.|date=2019|title=A revision of the holotype of Nanhsiungchelys wuchingensis, Ye, 1966 (Testudines: Cryptodira: Trionychoidae: Nanhsiungchelyidae)|journal=Cretaceous Research|volume=95|pages=151−163|doi=10.1016/j.cretres.2018.11.003|hdl=311034/9424|url=https://www.researchgate.net/publication/328822087}}</ref> [[Oviraptorid]] [[egg]]s are particularly common on the formation with atleast five well-preserved egg clutches.<ref>{{cite journal|last1=Yang|first1=T.-R.|last2=Wiemann|first2=J.|last3=Xu|first3=L.|last4=Cheng|first4=Y.-N.|last5=Wu|first5=X.-C.|last6=Sander|first6=P. M.|date=2019|title=Reconstruction of oviraptorid clutches illuminates their unique nesting biology|journal=Acta Palaeontologica Polonica|volume=466|pages=581−596|doi=10.4202/app.00497.2018|doi-access=free}}</ref> Moreover, a fossilized female oviraptorid with two eggs is known from the formation.<ref>{{cite journal|first1=T.|last1=Sato|first2=Y.-N.|last2=Cheng|first3=X.-C.|last3=Wu|first4=D. K.|last4=Zelenitsky|first5=Y.-F.|last5=Hsiao|date=2005|title=A Pair of Shelled Eggs Inside A Female Dinosaur|journal=Science|volume=308|issue=5720|pages=375–375|doi=10.1126/science.1110578|pmid=15831749|url=https://www.researchgate.net/publication/7904913}}</ref> The tracks of [[ornithopod]] dinosaurs are locally abundant at the Nanxiong region.<ref name=Xing2017>{{cite journal|last1=Xing|first1=L.|last2=Lockley|first2=M. G.|last3=Li|first3=D.|last4=Klein|first4=H.|last5=Ye|first5=Y.|last6=Scott Persons IV|first6=W.|last7=Ran|first7=H.|date=2017|title=Late Cretaceous ornithopod-dominated, theropod, and pterosaur track assemblages from the Nanxiong Basin, China: New discoveries, ichnotaxonomy, and paleoecology|journal=Palaeogeography, Palaeoclimatology, Palaeoecology|volume=466|pages=303−313|bibcode=2017PPP...466..303X|doi=10.1016/j.palaeo.2016.11.035|url=http://www.xinglida.net/pdf/Xing%20et%20al%202017%20Nanxiong%20tracks.pdf}}</ref>
+The therizinosaurids ''Nanshiungosaurus'' were bulky [[Browsing (herbivory)|high-browsers]] in their [[ecosystem]],<ref name=Paul2016/> and shared their habitat with multiple [[oviraptorosaur]] species such as ''[[Banji]]'', ''[[Ganzhousaurus]]'', ''[[Corythoraptor]]'', ''[[Nankangia]]'', ''[[Huanansaurus]]'' or ''[[Shixinggia]]''.<ref name=Lu2016>{{cite journal|last1=Lü|first1=J.|last2=Chen|first2=R.|last3=Brusatte|first3=S. L.|last4=Zhu|first4=Y.|last5=Shen|first5=C.|date=2016|title=A Late Cretaceous diversification of Asian oviraptorid dinosaurs: evidence from a new species preserved in an unusual posture|journal=Scientific Reports|volume=6|number=35780|bibcode=2016NatSR...635780L|doi=10.1038/srep35780|doi-access=free|pmc=5103654|pmid=27831542}}</ref><ref>{{cite journal|last1=Lü|first1=J.|last2=Li|first2=G.|last3=Kundrát|first3=M.|last4=Lee|first4=Y.-N.|last5=Sun|first5=Z.|last6=Kobayashi|first6=Y.|last7=Shen|first7=C.|last8=Teng|first8=F.|last9=Liu|first9=H.|date=2017|title=High diversity of the Ganzhou Oviraptorid Fauna increased by a new cassowary-like crested species|journal=Scientific Reports|volume=7|number=6393|bibcode=2017NatSR...7.6393L|doi=10.1038/s41598-017-05016-6|doi-access=free|pmc=5532250|pmid=28751667}}</ref> However, it is possible that some of these oviraptorosaurs did not actually live together given the poor stratigraphic analysis of the formation.<ref name=Lu2016/> Additional dinosaurs are represented by the fast running, long-snouted tyrannosaurids ''[[Qianzhousaurus]]'',<ref>{{cite journal|last1=Lü|first1=J.|last2=Yi|first2=L.|last3=Brusatte|first3=S. L.|last4=Yang|first4=L.|last5=Chen|first5=L.|date=2014|title=A new clade of Asian Late Cretaceous long-snouted tyrannosaurids|journal=Nature Communications|volume=5|number=3788|bibcode=2014NatCo...5.3788L|doi=10.1038/ncomms4788|doi-access=free|pmid=24807588}}</ref> the sauropods ''[[Gannansaurus]]'',<ref>{{cite journal|last1=Lü|first1=J.|last2=Yi|first2=L.|last3=Zhong|first3=H.|last4=Wei|first4=X.|date=2013 |title=A New Somphospondylan Sauropod (Dinosauria, Titanosauriformes) from the Late Cretaceous of Ganzhou, Jiangxi Province of Southern China|journal=Acta Geologica Sinica (English Edition)|volume=87|issue=3|pages=678−685|doi=10.1111/1755-6724.12079|url=http://www.geojournals.cn/dzxben/ch/reader/view_abstract.aspx?file_no=201303005&flag=1}}</ref> and the very sparce remains of hadrosaurid dinosaurs such as ''[[Microhadrosaurus]]'' (now a ''[[nomen dubium]]'').<ref name=Xing2017/> Other reptiles that composed the fauna were the terrestrial or semiaquatic [[nanhsiungchelyid]] turtles ''[[Nanhsiungchelys]]'' and ''[[Jiangxichelys]]'', [[squamates]] ''[[Chianghsia]]'' and ''[[Tianyusaurus]]'', and the [[crocodilians]] ''[[Jiangxisuchus]]''.<ref>{{cite journal |author1=Chun Li |author2=Xiao-chun Wu |author3=Scott Rufolo |year=2018 |title=A new crocodyloid (Eusuchia: Crocodylia) from the Upper Cretaceous of China |journal=Cretaceous Research |volume= 94|pages= 25–39|doi=10.1016/j.cretres.2018.09.015 }}</ref><ref name=Tong2019N/>
 ==See also==
 {{Portal|Dinosaurs}}
 * [[Timeline of therizinosaur research]]
-* [[Nanxiong Formation]]
 ==References==
 {{Reflist}}
-==External links==
-*[http://dml.cmnh.org/2000Oct/msg00117.html Details on "Nanshiungosaurus" bohlini]
 {{Theropoda|M.}}