Tuesday, January 9, 2018

Is "Apatosaurus" minimus a camarasaurid?

With the recent erection of Galeamopus for the diplodocid "Diplodocus" hayi by Tschopp et al. (2015), but also growing understanding of basal somphospondylan evolution (e.g. Averianov et al. in press; D'Emic et al. 2013; Saegusa and Ikeda 2014) and a preliminary assessment of the systematic affinities of "Apatosaurus" minimus Mook, 1917 by Taylor and Wedel (2012), I have had the chance to compare this problematic sauropod from the Morrison Formation with members of Macronaria, especially non-titanosaurian taxa, to better interpret placement within Neosauropoda. Indeed, McIntosh (1990a.b) and Upchurch et al. (2004) rejected the placement of minimus in Apatosaurus by Mook (1917) because of the height of the neural spines, low ilia with preacetabular processes being directed strongly laterally, and an ischial articular surface of the pubis nearly 50 percent of the pubic length. They considered it likely that "Apatosaurus" minimus was a derived member of Macronaria. Taylor and Wedel (2012) elaborated further, noting that the taxon has a mosaic of basal diplodocoid and macronarian characters, including tall neural spines and flaring ilia, but unpublished cladistics results were inconclusive. 

In their cladistic analyses of Diplodocoidea, Tschopp et al. (2015) noted that "Apatosaurus" minimus shares with Camarasaurus and most somphospondyls six sacral vertebrae and widely splayed preacetabular lobes of the ilium, even as they noted that the pubic morphology of AMNH 675 resembles Camarasaurus. For example, six sacral vertebrae are present in non-titanosaur somphospondyls like Euhelopus (Wilson and Upchurch 2009), Huabeisaurus (D'Emic et al. 2013), and Tambatitanis (Saegusa and Ikeda 2014), and some specimens referred to Camarasaurus (AMNH 690, BYU 17465, GMNH-PV 101) also have six sacral vertebrae (Tidwell et al. 2005). However, Upchurch et al. (2004) noted that AMNH 675 differs from titanosauriforms in that the cranial part of the ilium has a subtriangular outline in lateral view. Moreover, the somphospondylan Sibirotitan has five sacral vertebrae rather than six, in contrast to all other basal Somphospondyli (Averianov et al., in press) and "Apatosaurus" minimus.   

Although Taylor and Wedel (2012) listed tall neural spines on the sacral vertebrae as a diplodocoid synapomorphy for "Apatosaurus" minimus, the non-neosauropod eusauropod Cetiosauriscus, previously classified as a diplodocoid following McIntosh (1990a) and Upchurch et al. (2004) but now placed outside Diplodocoidea following Heathcote & Upchurch (2003) and Rauhut et al. (2005) also possesses tall sacral neural spines (Upchurch et al. 2004). Moreover, Tschopp et al. (2015, Supplementary Table 108) report that tall sacral neural spines occur among non-diplodocoid neosauropods. Given that tall neural spines of the sacral vertebrae are present in non-neosauropod eusauropods, it could be parsiminous to interpret this character as having evolved in more than one eusauropod clade, suggesting that tall sacral neural spines are a reversal in "Apatosaurus" minimus within Macronaria.  

Judging from comparisons of AMNH 675 with non-titanosaurian macronarians and non-neosauropod eusauropods, and evaluation of the characters cited by Taylor and Wedel (2012), the best parsimonious conclusion to be drawn is that "Apatosaurus" minimus may be a derived macronarian closely related to Somphospondyli, but possibly phylogenetically intermediate between Camarasauridae and Titanosauriformes. The number of sacral vertebrae is distinct from Camarasaurus (except in AMNH 690, BYU 17465, GMNH-PV 101) and more closely matching members of Somphospondyli except Sibirotitan, but the subtriangular outline of the cranial portion of the ilium in lateral view excludes "A." minimus from Titanosauriformes, while the tall neural spines distinguish the species not just from camarasaurids but also from titanosauriforms.              

References:

Averianov, A., Ivantsov, S., Skutschas, P., Faingertz, A., and Leshchinskiy, S., in press. A new sauropod dinosaur from the Lower Cretaceous Ilek Formation, Western Siberia, Russia. Geobios DOI: https://doi.org/10.1016/j.geobios.2017.12.004


D'Emic, M.D., Mannion, P.D., Upchurch, P., Benson, R.B.J., Pang, Q., and Zhengwu, C., 2013. Osteology of Huabeisaurus allocotus (Sauropoda: Titanosauriformes) from the Upper Cretaceous of China. PLoS ONE 8(8): e69375. https://doi.org/10.1371/journal.pone.0069375
, and ., 2003. The relationships of Cetiosauriscus stewarti (Dinosauria; Sauropoda): implications for sauropod phylogeny. Journal of Vertebrate Paleontology 23:60A. 

., 1990a. Sauropoda. pp. 345-401. In: , , , (eds.) The Dinosauria, 1st edition. Berkeley: University of California Press

McIntosh J.S., 1990b. Species determination in sauropod dinosaurs with tentative suggestions for their classification. pp. 53-69. In: Carpenter K, Currie PJ, (eds.) Dinosaur systematics: perspectives and approaches. New York: Cambridge University Press.

., 1917. Criteria for the determination of species in the Sauropoda, with description of a new species of Apatosaurus. Bulletin of the American Museum of Natural History 37:355-358. 

, , , , ., 2005. Discovery of a short-necked sauropod dinosaur from the Late Jurassic period of Patagonia. Nature 435:670-672. 

Saegusa, H., and Ikeda, T., 2014. A new titanosauriform sauropod (Dinosauria: Saurischia) from the Lower Cretaceous of Hyogo, Japan. Zootaxa. 3848 (1): 1–66. doi:10.11646/zootaxa.3848.1.1

Taylor, Michael P., and Mathew J. Wedel. 2012. Re-evaluating "Apatosaurus" minimus, a bizarre Morrison Formation sauropod with diplodocoid and macronarian features. p. 23 in Matt Friedman and Graeme Lloyd (eds.), Programme and Abstracts, 60th Annual Symposium of Vertebrate Palaeontology and Comparative Anatomy, University of Oxford, Oxford, UK, September 10th-15th 2012. 33 pp.

, , and ., 2005. Age-related characteristics found in a partial pelvis of Camarasaurus. pp. 18-186. In: , Thunder-lizards: the sauropodomorph dinosaurs. Bloomington: Indiana University Press

Tschopp. E., Mateus, O., and Benson, R.B.J., 2015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda) PeerJ 3:e857

Upchurch, P., Barrett, P.M., Dodson, P., 2004. Sauropoda. pp. 259-322. In: Weishampel DB, Dodson P, Osmólska H, editors. The Dinosauria, 2nd ed. Berkeley: University of California Press. pp. 259–322.     

, and ., 2009. Redescription and reassessment of the phylogenetic affinities of Euhelopus zdanskyi (Dinosauria: Sauropoda) from the Early Cretaceous of China. Journal of Systematic Palaeontology 7:199-239. 

Friday, August 12, 2016

Are Europe's latest Cretaceous titanosaurs descended from a Central Asian ancestor?

Titanosaurian sauropods from the latest Cretaceous of Europe have been documented in the published literature since Matheron (1869) described Hypselosaurus priscus from fragmentary postcranial remains in the Provence region of southern France and Paul Gervais recorded titanosaur remains from marine deposits in the Aquitane region of southwestern France (Buffetaut et al. 1991). Although the discoveries of Ampelosaurus, Atsinganosaurus, Lirainosaurus, Magyarosaurus, Paludititan and now Lohuecotitan (Diaz et al. 2016), attest to the diversity of titanosaurs in the last few million years of the Cretaceous, few authors have attempted to discern the paleobiogeographical origins of Europe's latest Cretaceous titanosaur fauna by cladistic and non-cladistic means (Curry Rogers 2005; Garcia et al. 2010). However, the description of the aralosaurin lambeosaurine hadrosaurid Canardia from the same region as Ampelosaurus and Atsinganosaurus (Prieto-Marquez et al. 2013), the recognition of Pararhabdodon as a relative of the tsintaosaurin lambeosaurine Tsintaosaurus by Prieto-Marquez and Wagner (2009), and the discovery indeterminate titanosaur remains from the Bissekty Formation of Uzbekistan and the Dabrazinskaya Svita of Kazakhstan (Riabinin 1939; Sues et al. 2015), has led me to consider the possibility that either all titanosaur species from latest Cretaceous Europe, or at least some taxa, were descended from a titanosaur that immigrated to Europe from Central Asia during the latest Cretaceous.

In their description of titanosaur remains from the Bissekty Formation, Sues et al. (2015) note that the titanosaur braincase CCGME 628/12457 differs from the braincases of Lirainosaurus in lacking distal foramina on the basal tubera of the paroccipital processes and basal tubera separated by a wide depression ventral to the occipital condyle, with a round pit forming the center of the depression. Nevertheless, the presence of the abducens nerve VI extending lateral to the pituitary fossa is shared by both Lirainosaurus and CCGME 628/12457 along with other derived titanosaurs (cf. Sues et al. 2015, figs. 3-4 with Knoll et al. 2013), and the fact that the aforementioned features of the Bissekty braincase are also seen in several titanosaurs for which braincases are known (e.g. Jainosaurus, Muyelensaurus, Nemegtosaurus, Pitekunisaurus, and Rapetosaurus) may dampen the usefulness of braincase characters for determining the biogeographical origins of late Cretaceous European titanosaurs.

Although Garcia et al. (2010) suggested that Atsinganosaurus could be a European descendant of African lithostrotians based on comparisons with the caudal vertebrae of the basal lithostrotian Malawisaurus, they caution that a comprehensive phylogenetic analysis of Titanosauria is needed to confirm or refute the possibility of a Gondwanan origin for Atsinganosaurus. As a matter of fact, the near-lack of sauropod remains from pre-Turonian Cretaceous sediments in Central Asia (see Weishampel et al. 2004 for available records) suggests that a group of lithostrotians more primitive than Saltasauridae may have colonized Central Asia from Gondwana via rudimentary land bridges to Asia, because Garcia et al. (2010) note that Ampelosaurus and Lirainosaurus are more derived than Atsinganosaurus. Likewise, the placement of AmpelosaurusLirainosaurus, and Lohuecotitan by Diaz et al. (2013, 2016) and Garcia et al. (2013) bolsters the alternative hypothesis that even if some European titanosaurs are more primitive than others, they still could have evolved from a Central Asian ancestor because of the dearth of Early Cretaceous (Neocomian) non-avian dinosaur fossils from Central Asia.


References:

Buffetaut E, Cuny G, Le Loeuff J., 1991. French dinosaurs: The best record in Europe? Modern Geology 16: 17–42.

Curry Rogers, K. A., 2005. Titanosauria: A Phylogenetic Overview. pp. 50-103. In: Curry Rogers and Wilson (eds), The Sauropods: Evolution and Paleobiology. University of California Press: Berkeley.

Díez Díaz, V., Pereda Suberbiola, X., and Sanz, J.L. 2011. Braincase anatomy of the titanosaurian sauropod Lirainosaurus astibiae from the Late Cretaceous of the Iberian Peninsula. Acta Palaeontologica Polonica 56 (3): 521–533.

Díez Díaz, Verónica, Pereda Suberbiola, Xabier, and Sanz, José Luis. 2013. Appendicular skeleton and dermal armour of the Late Cretaceous titanosaur Lirainosaurus astibiae (Dinosauria: Sauropoda) from Spain, Palaeontologia Electronica Vol. 16, Issue 2; 19A; 18p; palaeo-electronica.org/content/2013/502-titanosaur-skeleton 

V. Díez Díaz; P. Mocho; A. Páramo; F. Escaso; F. Marcos-Fernández; J.L. Sanz; F. Ortega, 2016. A new titanosaur (Dinosauria, Sauropoda) from the Upper Cretaceous of Lo Hueco (Cuenca, Spain). Cretaceous Research. in press. doi:10.1016/j.cretres.2016.08.001.

Géraldine Garcia; Sauveur Amico; Francois Fournier; Eudes Thouand; Xavier Valentin, 2010. A new titanosaur genus (Dinosauria, Sauropoda) from the Late Cretaceous of southern France and its paleobiogeographic implications. Bulletin de la Societe Geologique de France. 181 (3): 269–277. doi:10.2113/gssgfbull.181.3.269.

Prieto-Márquez, A.; Dalla Vecchia, F. M.; Gaete, R.; Galobart, À., 2013. Diversity, Relationships, and Biogeography of the Lambeosaurine Dinosaurs from the European Archipelago, with Description of the New Aralosaurin Canardia garonnensis. PLoS ONE. 8 (7): e69835. doi:10.1371/journal.pone.0069835.

Matheron, P. (1869). Note sur les reptiles fossiles des dépôts fluvio-lacustres crétaces du bassin à lignite de Fuveau. Bulletin de la Société géologique de France. 26 (2): 781–795.


Riabinin, A.N. 1939. [The Upper Cretaceous vertebrate fauna of south Kazakhstan I. Reptilia. Pt. 1 Ornithischia]. Tsentral. Nauchno-issled. Geol. Inst. Trudy. 118: 1-40. [In Russian]

Sues, H.-D., A. Averianov, and R. C. Ridgely, and L. M. Witmer (2015) Titanosauria (Dinosauria, Sauropoda) from the Upper Cretaceous (Turonian) Bissekty Formation of Uzbekistan. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2014.889145
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004. Dinosaur Distribution. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second Edition. University of California Press. 517-606.

Wednesday, November 26, 2014

Is Caulodon a synonym of Camarasaurus?

The non-titanosauriform macronarian Camarasaurus is one of the American public's favorite macronarian sauropods from the Jurassic, being thought of as the most common macronarian from the Morrison Formation. Very few sauropod workers, however, have tackled the alpha-taxonomy of the genus and its various constituent taxonomy since the papers by Ikejiri (2005) and Osborn and Mook (1921). The result has been the tendency to treat Camarasaurus as a genus in numerous phylogenies of Sauropoda (Upchurch et al. 2004; Wilson 2002) without a comprehensive analysis of intrageneric variation of the genus.

In recent abstracts published at the SVP 2013 and 2014 meetings, Mateus and Tschopp (2013) and Tschopp et al. (2014) work to address the question of intraspecific variation in Camarasaurus. The abstract by Mateus and Tschopp (2013) deals with a newly discovered camarasaurid (SMA 0002) that is referred to Cathetosaurus lewisi based on shared characters with BYU 9047 (holotype of C. lewisi). On the other hand, Tschopp et al. (2013) conduct a specimen-level analysis of all Camarasaurus specimens including the type specimens of currently recognized species of the genus (C. grandis, C. lentus, and C. supremus). The phylogenetic results reported by Tschopp et al. (2014) seem to not only reaffirm the distinctness of Cathetosaurus from Camarasaurus, but they also support the validity of the three historical species of the latter genus.

These preliminary results have prompted me to revisit the synonymy of Caulodon with Camarasaurus performed by Osborn and Mook (1921) and followed by subsequent authors. When placing the two nominal species of Caulodon (C. diversidens and C. leptoganus) in synonymy with Camarasaurus supremus, Osborn and Mook noted that AMNH 5768 and AMNH 5769 were similar to the maxillary teeth of a referred specimen of Camarasaurus supremus (AMNH 5761) in being robust and spatulate-shaped, but were hesitant to rule out the possibility of two different sauropods with spatulate teeth from the Morrison Formation. However, they did not compare the holotypes of Caulodon diversidens and C. leptoganus with the teeth of Giraffatitan or USNM 5730 (referred to Brachiosaurus sp. by Carpenter and Tidwell 1998).

Since the monograph by Osborn and Mook, robust spatulate teeth have been described for numerous non-neosauropod and macronarian sauropods, including USNM 5730, Europasaurus, Mamenchisaurus, Turiasaurus, and Jobaria (Carpenter and Tidwell, 1998; Marpmann et al. 2014; Ouyang and He 2002; Royo-Torres and Upchurch 2012; Russell and Zheng 1994; Sereno et al. 1999). Because robust spatulate teeth are no longer considered diagnostic for Camarasaurus supremus or other camarasaurid species and are widely distributed among non-neosauropod and non-titanosauriform sauropods, Caulodon and its nominal species should not be considered synonymous with Camarasaurus supremus and instead must be considered nomina dubia at Eusauropoda indeterminate.


          

Holotype teeth of Caulodon diversidens (left) and C. leptoganus (right) (after Steel 1970)

     
References:

Carpenter, K., and Tidwell, V., 1998, Preliminary description of a Brachiosaurus skull from Felch Quarry 1, Garden Park, Colorado: In: The Upper Jurassic Morrison Formation: An Interdisciplinary Study. Edited by Carpenter, K., Chure, D. J., and Kirkland, J. I. Modern Geology 23 (2): 69-84.

Ikejiri, T., 2005, Distribution and biochronology of Camarasaurus (Dinosauria, sauropoda) from the Jurassic Morrison Formation of the Rocky Mountain Region: New Mexico Geological Society, 56th Field Conference Guidebook, Geology of the Chama Basin, 2005, pp. 367-379.

Marpmann, J., Carballido, J., Sander, P., Knötschke, N. 2014. Cranial anatomy of the Late Jurassic dward sauropod Europasaurus holgeri (Dinosauria, Camarasauromorpha): ontogenetic changes and size dimorphism. Journal of Systematic Paleontology. DOI: 10.1080/14772019.2013.875074

Mateus, O., & Tschopp E. (2013). Cathetosaurus as a valid sauropod genus and comparisons with Camarasaurus. Journal of Vertebrate Paleontology, Program and Abstracts, 2013. 173.

Osborn, H. F., and Mook, C. C., 1921, Camarasaurus, Amphicoelias, and other Sauropods of Cope: Memoris of the American Museum of Natural History, new series, v. 3, part 3, p. 249-387.

Ouyang, H., and Ye, Y., 2002, The first mamenchisaurian skeleton with complete skull, Mamenchisaurus youngi: 111pp.

Rafael Royo-Torres & Paul Upchurch, 2012, "The cranial anatomy of the sauropod Turiasaurus riodevensis and implications for its phylogenetic relationships", Journal of Systematic Palaeontology, 10(3): 553-583.

Russell, D. A., and Zheng, Z., 1994, A large mamenchisaurid from the Junggar Basin, Xinjiang, People’s Republic of China: In: Results from the Sino-Canadian Dinosaur Project. Canadian Journal of Earth Sciences, v. 30, p. 2082-2095.

Sereno, P, C., Beck, A. L., Dutheil, D. B., Larson, H. C. E., Lyon, G. H., Moussa, B., Sadleir, R. W., Sidor, C. A., Varricchio, D. J., Wilson, G. P., and Wilson, J. A., 1999, Cretaceous Sauropods from the Sahara and the Uneven Rate of Skeletal Evolution Among Dinosaurs: Science, v. 286, p. 1342-1347.

Steel, R., 1970, Saurischia: Handbuch der Palaoherpetology. Volume 14. Gustav Fischer Verlag.

Tschopp, E., Mateus O., Kosma R., Sander M., Joger U., & Wings O., 2014. A specimen-level cladistic analysis of Camarasaurus (Dinosauria, Sauropoda) and a revision of camarasaurid taxonomy. Journal of Vertebrate Paleontology. Program and Abstracts, 2014, 241-242.

Upchurch, P.M., Barrett, P.M., and Dodson, P. (2004). Sauropoda. In: Weishampel, D.B., Dodson, P., and Osmólska, H. (eds.). The Dinosauria (2nd edition). University of California Press:Berkeley 259-322. ISBN 0-520-24209-2

Wilson, J. A., 2002, Sauropod dinosaur phylogeny: critique and cladistic analysis: Zoological Journal of the Linnean Society, v. 136, p. 217-276.

Sunday, September 7, 2014

Dreadnoughtus and the true contender for the title of heaviest sauropod

Ever since the 1980s, many dinosaurologists familiar with sauropods have become fascinated by the specter of extravagantly colossal sauropod species, fortified by the discoveries of the giant diplodocids Seismosaurus hallorum (now Diplodocus hallorum) and Supersaurus vivianae in the 1970s and 1980s. When James Jensen announced the discovery of Supersaurus and its junior synonym Ultrasauros, the mainstream media outlets hailed Ultrasauros as the largest and heaviest sauropod that ever lived, and based on the holotype (BYU 9044) and the brachiosaurid scapulocoracoid BYU 9462, Jim Jensen put the estimates of the length and weight of Ultrasauros at 80 to 100 feet long (25 to 30 meters) and 75 tons (75,000 kilograms) respectively. On the other hand, the discoverer of Diplodocus hallorum, David Gillette, estimated the maximum length and weight of D. hallorum at 177 feet long (54 meters) and 125 tons (113,000 kilograms) respectively. However, the giant Morrison sauropods were eclipsed by the Patagonian titanosaurs as regards discussion of the heaviest sauropod taxa, and recent publications (Carpenter 2006; Foster 2003) have put Diplodocus hallorum and Supersaurus vivianae in the 33 to 50 ton range (30,000 to 45,000 kilograms), a far cry from the earlier weight estimates.

Now, the media has seized upon the discovery of the latest giant titanosaur to be described, Dreadnoughtus schrani (Lacovara et al. 2014), by hailing it as probably the biggest and heaviest sauropod ever to have roamed the earth. This discovery has provided a new window into the question of which sauropod was the heaviest that ever lived, but to compare Dreadnoughtus against other contenders for the title of heaviest sauropod (Argentinosaurus, Futalognkosaurus, Puertasaurus), I have the opportunity to discuss various weight and size estimates for the giant Patagonian titanosaurs with respect to Equation 1 devised by Campione and Evans (2012), which extrapolates the weight of a quadrupedal animal from the minimum circumference of the shaft of its humerus and femur.

When using Equation 1 of Campione and Evans (2012) in order to calculate the body mass of Dreadnoughtus, Lacovara et al. (2014) put the body mass estimate of this genus at 65.4 tons (59,291 kilograms), while noting that the body mass of Dreadnoughtus was probably greater taking into account their histological analysis indicating that the holotype (MPM-PV 1156) was not yet fully grown but still massive in terms of body mass. This is well above the weight estimates provided for Giraffatitan brancai by Benson et al. (2014) but less than the body mass estimate of 134.9 tons (122,400 kilograms) provided for the diplodocoid "Amphicoelias" fragillimus by Carpenter (2006), and less than the weight estimated by Mazetta et al. (2004) for the titanosaur "Antarctosaurus" giganteus. (Since the holotype remains of "Amphicoelias" fragillimus [AMNH 5777] have been lost, it is possible that Carpenter's weight estimate for this species is way off by several tons and "A." fragillimus was not as heavy as estimated by Carpenter.) By contrast, the primitive titanosaur Futalognkosaurus and the turiasaur Turiasaurus weighed 42 tons (38,139 kilograms) and 56.1 tons (50,923 kilograms) respectively.



Regarding the body mass of Argentinosaurus, Paul (1994) put the weight estimate of Argentinosaurus at 88 to 110 tons (80,000 to 100,000 kilograms), but Mazzetta et al. (2004) revised the weight to 80 tons (73,000 kilograms), while Sellers et al. (2013) put the weight estimate at 91 tons (83,000 kilograms). Although it is not implausible that Argentinosaurus was bigger and heavier than Dreadnoughtus judging from the size of the referred femur (MLP-DP 46-VIII-21-3), the known material of Argentinosaurus comprises only a tiny part of the postcranial skeleton (9.2 percent), insufficient to give a reliable estimate of the body mass of Argentinosaurus using the body mass formula devised by Campione and Evans (2012). Likewise, another giant titanosaur from Patagonia, Puertasaurus, may have been heavier and bigger than Dreadnoughtus, with an estimated weight of 88 to 110 tons (80,000 to 100,000 kilograms) based on the huge size of the dorsal vertebra (Novas et al. 2005), but the available material is too meagre to give a more accurate body mass estimate and more complete remains are needed to determine the size of Puertasaurus

In summary, any possible body mass estimates for "Amphicoelias" fragillimus, Argentinosaurus, and Puertasaurus should be treated with caution when taking these sauropods as candidates for the title of heaviest sauropod. Since "Antarctosaurus" giganteus has femora preserved in its type material, it could be a viable contender for the title of heaviest sauropod. Therefore, it may be parsimonious to treat "A." giganteus and Dreadnoughtus as the heaviest sauropods known to science given that the holotype of the former species was not yet fully mature.         

Update: A new paper by Bates et al. (2015) suggests that the original weight estimate for Dreadnoughtus was an overestimate and that Dreadnoughtus probably weighed 42.1 tons (38,225 kilograms), rather than 65.4 tons as originally claimed by Lacovara et al. (2014).

Benson RBJ, Campione NE, Carrano MT, Mannion PD, Sullivan C, et al., 2014. Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage. PLoS Biol 12(5): e1001853 doi:10.1371/journal.pbio.1001853.

Campione, N., and Evans, D., 2012. A universal scaling relationship between body mass and proximal limb bone dimensions in quadrupedal terrestrial tetrapods". BMC Biology: 15. doi:10.1186/1741-7007-10-60.

Carpenter, K., 2006. Biggest of the big: a critical re-evaluation of the mega-sauropod Amphicoelias fragillimus. In Foster, John R.; and Lucas, Spencer G. (eds.). Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin 36. Albuquerque: New Mexico Museum of Natural History and Science. pp. 131–138.

Foster, J.R., 2003. Paleoecological analysis of the vertebrate fauna of the Morrison Formation (Upper Jurassic), Rocky Mountain region, U.S.A. New Mexico Museum of Natural History and Science Bulletin, 23. Albuquerque, New Mexico: New Mexico Museum of Natural History and Science.

Lacovara, Kenneth J.; Ibiricu, L.M.; Lamanna, M.C.; Poole, J.C.; Schroeter, E.R.; Ullmann, P.V.; Voegele, K.K.; Boles, Z.M.; Egerton, V.M.; Harris, J.D.; Martínez, R.D.; Novas, F.E., 2014. A Gigantic, Exceptionally Complete Titanosaurian Sauropod Dinosaur from Southern Patagonia, Argentina. Scientific Reports. doi:10.1038/srep06196.

Mazzetta, G. V., Christiansen, P., Fariña, R. A., 2004. Giants and Bizarres: Body Size of Some Southern South American Cretaceous Dinosaurs. Historical Biology 16 (2-4): 71–83. doi:10.1080/08912960410001715132.

Novas, F.E.,  Salgado, L., Calvo, J., Agnolin, F., 2005. Giant titanosaur (Dinosauria, Sauropoda)from the Late Cretaceous of Patagonia. Revisto del Museo Argentino de Ciencias Naturales, n.s. 7 (1): 37–41.

Paul, G. S., 1994. Big Sauropods - Really, Really Big Sauropods. The Dinosaur Report. The Dinosaur Society. pp. 12–13.

Sellers, W. I.; Margetts, L.; Coria, R. A. ­B.; Manning, P. L., 2013. March of the Titans: The Locomotor Capabilities of Sauropod Dinosaurs. In Carrier, David. PLoS ONE 8 (10): e78733. doi:10.1371/journal.pone.0078733. PMC 3864407. PMID 24348896.  edit

Thursday, May 29, 2014

McPhee et al. (2014) paper on Antetonitrus, part 2: is Thotobolosaurus a relative of Antetonitrus?

In part 1 of my review of the McPhee et al. (2014) paper on the osteology of the basal sauropod Antetonitrus, I discussed the definition of Sauropodiformes and Sauropoda within in the context of the analysis by McPhee et al. (2014) regarding Antetonitrus. However, one overlooked aspect of the paper that didn't make anyone notice it was the discussion of the possible affinity of the Maphutseng sauropodomorph (aka "Thotobolosaurus") with Antetonitrus on page 157 of the paper.

The Maphutseng sauropodomorph, like the Proctor Lake hypsilophodont and the Dalton Wells quarry iguanodont, has been occasionally mentioned in the literature but has been only briefly described. It was first reported by Ellenberger and Ellenberger (1956) based on sauropodomorph remains pertaining to a minimum of six individuals collected from from the Late Triassic lower Elliot Formation of Maphutseng, Lesotho in the 1950s. The Maphutseng taxon was discussed by Charig et al. (1965), Ellenberger (1970), and Ellenberger & Ginsburg (1966), who noted its affinities with sauropods relative to other "prosauropods"; Ellenberger (1970) coined the nomen nudum "Thotobolosaurus mabeatae" for the Maphutseng sauropodomorph in his discussion of the fossil record of southern Africa for the Triassic-early Jurassic interval, but never described it in detail. Gauffre (1993) preliminarily re-assessed the Maphutseng sauropodomorph and tentatively referred it to the nomen dubium Euskelosaurus browni, but he later (1996) changed his mind about the attribution of this taxon and described it as a distinct sauropodomorph under the nomen ex dissertationae "Kholumolumosaurus ellenbergerorum".



CO 1069-209-59

The remains of "Thotobolosaurus"/"Kholumolumosaurus" being excavated in Maphutseng, Lesotho.

Note: Since this post was published, it has become clear (Peyre de Fabregues and Allain 2016) that Antetonitrus is not from the lower Elliot Formation as previously thought, but instead from the upper Elliot Formation, meaning that the Maphutseng sauropodomorph is clearly not congeneric with Antetonitrus


Charig, A.J., Attridge, J.& Crompton, A.W. 1965.On the origin of the sauropods and the classification of the Saurischia. Proceedings of the Linnean Society 176: 197–221.

Ellenberger, F., and Ellenberger, P. 1956. Le gisement de Dinosauriens de Maphutseng (Basutoland, Afrique du Sud) [The Maphutseng dinosaur locality (Basutoland, South Africa)]. Comptes Rendus de la Société géologique de France 1956:99-101.


F. Ellenberger and L. Ginsburg. 1966. Le gisement de Dinosauriens triasiques de Maphutseng (Basutoland) et l'origine des Sauropodes [The Triassic dinosaur locality of Maphutseng (Basutoland) and the origin of sauropods]. Comptes Rendus de l'Académie des Sciences à Paris, Série D 262:444-447.



Ellenberger, P.. 1970. Les niveaux paléontologiques de première apparition des mammifères primoridaux en Afrique du Sud et leur ichnologie. Establissement de zones stratigraphiques detaillees dans le Stormberg du Lesotho (Afrique du Sud) (Trias Supérieur à Jurassique) [The paleontological levels of the first appearance of primordial mammals in southern Africa and their ichnology. Establishment of detailed stratigraphic zones in the Stormberg of Lesotho (southern Africa) (Upper Triassic to Jurassic). In: S. H. Haughton (ed.), Second Symposium on Gondwana Stratigraphy and Paleontology, International Union of Geological Sciences. Council for Scientific and Industrial Research, Pretoria 343-370.



F.-X. Gauffre. 1993. Biochronostratigraphy of the lower Elliot Formation (southern Africa) and preliminary results on the Maphutseng dinosaur (Saurischia: Prosauropoda) from the same formation of Lesotho. In S. G. Lucas and M. Morales (eds.), The Nonmarine Triassic. New Mexico Museum of Natural History and Science Bulletin 3:147-149.

Gauffre, F. -X., 1996. Phylogenie des dinosaures prosauropodes et etude d’un prosauropode du Trias superieur d’Afrique australe. PhD thesis. Museum National d’Histoire Naturelle, Paris. 156 pp.

Peyre de Fabrègues, C., and Allain, R., 2016. New material and revision of Melanorosaurus thabanensis, a basal sauropodomorph from the Upper Triassic of Lesotho. PeerJ 4: e1639. doi:10.7717/peerj.1639.