Nemegtosauridae

Nemegtosauridae is a family of titanosaurian sauropod dinosaurs based on their diplodocid-like skulls.[1][2][3] Only three species are known:[4] Nemegtosaurus, Quaesitosaurus and Tapuiasaurus, each from the Cretaceous.

Nemegtosauridae Temporal range: Late Cretaceous, 120–66 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Cast of the skull of Nemegtosaurus, on a mounted Opisthocoelicaudia skeleton, Museum of Evolution of Polish Academy of Sciences, Warsaw
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Dinosauria
Clade:	Saurischia
Suborder:	†Sauropodomorpha
Clade:	†Sauropoda
Clade:	†Eusauropoda
Clade:	†Neosauropoda
Clade:	†Macronaria
Clade:	†Titanosauria
Clade:	†Eutitanosauria
Clade:	†Lithostrotia
Family:	†Nemegtosauridae Upchurch, 1995
Genera
†Nemegtosaurus †Quaesitosaurus †Tapuiasaurus

History of classification

Due to the diplodocid-like nature of the taxa placed in Nemegtosauridae, the systematic position of this family in Sauropoda was disputed until recently. McIntosh (1990) included both these animals in the family Diplodocidae, subfamily Dicraeosaurinae, as they resemble the skull of Dicraeosaurus, although differing in certain details. Although the skull of Nemegtosaurus was found in the same formation as the headless skeleton of Opisthocoelicaudia, McIntosh (1990) kept Nemegtosaurus in Diplodocoidea while keeping Opisthocoelicauda separate from the former, a position reiterated by Upchurch (1995,[5] 1999[6]), and Upchurch et al. (2004). A cladistic analysis published in 2002 transferred Nemegtosaurus and Opisthocoelicauda from Diplodocoidea to Titanosauria.[1]

Apesteguia (2004), in a paper describing a new Patagonian sauropod, Bonitasaura salgadoi, may have been the first to properly define the taxon, although without the use of cladistic analysis: the stemclade consisting of all titanosaurs more closely related to Nemegtosaurus than to Saltasaurus. He argued for a close relationship between Nemegtosaurus, Quaesitosaurus, Rapetosaurus, and Bonitasaura and referred to the previous phylogenetic analysis and use of Nemegtosauridae by Wilson (2002).[7]

In his redescription of the Nemegtosaurus holotype, Wilson (2005) elaborated on the titanosaurian nature of Nemegtosaurus, defining Nemegtosauridae as a stem-based clade that includes all titanosaurs more closely related to Nemegtosaurus than to Saltasaurus. He also suggested that Opisthocoelicaudia may eventually be shown to be a junior synonym of Nemegtosaurus.[8] For her part, Kristina Curry Rogers (see also Cuury Rogers and Forster [2001][9]) agreed with Wilson that both Nemegtosaurus and Quaesitosaurus were titanosaurs rather than diplodocoids, but rejected the validity of Nemegtosauridae and the clade concepts given under that name. Quaesitosaurus was placed in the Saltasaurinae and Nemegtosaurus in a new, unnamed "Rapetosaurus clade" (which, under ICZN rules, would, if named, be termed subfamily Nemegtosaurinae or tribe Nemegtosaurini, depending on its position). Opisthocoelicaudia was placed in a separate clade, the Opisthocoelicaudiinae. All three clades are included in the Saltasauridae (= Titanosauridae).[10]

In a paper discussing new anatomical data on the skull of Tapuiasaurus, Wilson and his colleagues cast doubt on the monophyly of Nemegtosauridae, judging from a rescoring of the Zaher et al. 2011 cladistic analysis regarding cranial characters. Tapuiasaurus was recovered as basal to Lithostrotia, rendering its position within Nemegtosauridae questionable.[11] A 2014 cladistic analysis gleaning new anatomical data from Diamantinasaurus also rendered Nemegtosauridae paraphyletic, with Rapetosaurus falling out as a member of Saltasauridae closer to Isisaurus than to Nemegtosaurus.[12] The cladistic analysis of Patagotitan recovered Tapuiasaurus as the sister taxon of Rapetosaurus and Isisaurus but not Nemegtosaurus.[13]

Phylogeny

Skull material of Tapuiasaurus

The cladogram below follows Zaher et al. (2011).[14]

Lithostrotia	Malawisaurus     Nemegtosauridae    Nemegtosaurus       Rapetosaurus     Tapuiasaurus           Isisaurus       Diamantinasaurus   Saltasauridae  Opisthocoelicaudiinae    Alamosaurus     Opisthocoelicaudia     Saltasaurinae    Neuquensaurus     Saltasaurus

References

1. Wilson, J.A. (13 September 2002). "Sauropod dinosaur phylogeny: critique and cladistic analysis" (PDF). Zoological Journal of the Linnean Society. The Linnean Society of London. 136 (2): 215–275. doi:10.1046/j.1096-3642.2002.00029.x.
2. McIntosh, J. S., 1990, "Sauropoda" in The Dinosauria, Edited by David B. Weishampel, Peter Dodson, and Halszka Osmólska. University of California Press, pp. 345–401.
3. Upchurch, P., Barrett, P.M. and Dodson, P. 2004. Sauropoda. In The Dinosauria, 2nd edition. D. Weishampel, P. Dodson, and H. Osmólska (eds.). University of California Press, Berkeley. pp. 259–322.
4. Re: Family Nemegtosauridae
5. Upchurch, P (1995). "The evolutionary history of sauropod dinosaurs". Philosophical Transactions of the Royal Society of London B. 349: 365–390. Bibcode:1995RSPTB.349..365U. doi:10.1098/rstb.1995.0125.
6. Upchurch, P (1999). "The phylogenetic relationships of the Nemegtosauridae (Saurischia, Sauropoda)". Journal of Vertebrate Paleontology. 19: 106–125. doi:10.1080/02724634.1999.10011127.
7. Apesteguía, S. (10 September 2004). "Bonitasaura salgadoi gen. et sp. nov.: a beaked sauropod from the Late Cretaceous of Patagonia". Naturwissenschaften. 91 (10): 493–497. Bibcode:2004NW.....91..493A. doi:10.1007/s00114-004-0560-6. PMID 15729763.
8. Wilson, J.A. (24 August 2005). "Redescription of the Mongolian sauropod Nemegtosaurus mongoliensis Nowinski (Dinosauria: Saurischia) and comments on Late Cretaceous sauropod diversity". Journal of Systematic Palaeontology. The Natural History Museum. 3 (3): 283–318. doi:10.1017/S1477201905001628 – via ResearchGate.
9. Rogers, K.C.; Forster, C.A. (2 August 2001). "The last of the dinosaur titans: a new sauropod from Madagascar". Nature. 412 (6846): 530–534. Bibcode:2001Natur.412..530C. doi:10.1038/35087566. PMID 11484051 – via ResearchGate.
10. Rogers, K.C.; Wilson, J. (2005). The Sauropods: Evolution and Paleobiology. University of California Press. pp. 50–103. ISBN 0520246233.
11. Wilson, J.A.; Pol, D.; Carvalho, A.B.; Zaher, H. (9 February 2016). "The skull of the titanosaur Tapuiasaurus macedoi (Dinosauria: Sauropoda), a basal titanosaur from the Lower Cretaceous of Brazil". Zoological Journal of the Linnean Society. The Linnean Society of London. 178 (3): 611–662. doi:10.1111/zoj.12420 – via ResearchGate.
12. Poropat, S.F.; Upchurch, P.; Mannion, P.D.; Hocknull, S.; Kear, B.P.; Sloan, T.; Sinapius, G.H.K.; Elliott, D.A. (18 April 2014). "Revision of the sauropod dinosaur Diamantinasaurus matildae Hocknull et al. 2009 from the mid-Cretaceous of Australia: Implications for Gondwanan titanosauriform dispersal". Gondwana Research. 27 (3): 995–1033. doi:10.1016/j.gr.2014.03.014 – via ResearchGate.
13. Jose, C.; Pol, D.; Otero, A.; Cerda, I.A.; Salgado, L.; Garrido, A.; Ramezani, J.; Cunéo, R.; Krause, J.M. (6 July 2017). "A new giant titanosaur sheds light on body mass evolution among sauropod dinosaurs". Proceedings of the Royal Society B: Biological Sciences. The Royal Society Publishing. 284 (1860): 20171219. doi:10.1098/rspb.2017.1219. PMC 5563814. PMID 28794222.
14. Hussam Zaher, Diego Pol, Alberto B. Carvalho, Paulo M. Nascimento, Claudio Riccomini, Peter Larson, Rubén Juarez-Valieri, Ricardo Pires-Domingues, Nelson Jorge da Silva Jr., Diógenes de Almeida Campos (2011). "A Complete Skull of an Early Cretaceous Sauropod and the Evolution of Advanced Titanosaurians". PLoS ONE. 6 (2): e16663. Bibcode:2011PLoSO...616663Z. doi:10.1371/journal.pone.0016663. PMC 3034730. PMID 21326881.CS1 maint: multiple names: authors list (link)