Wednesday, January 2, 2019
Did lognkosaurs originate in Laurasia?
Last year yielded a number of surprises and new developments in terms of sauropod biogeography, including the description of the youngest sauropod from mainland Africa (Mansourasaurus), the erection of Maraapunisaurus for the apocryphal gigantic sauropod Amphicoelias fragillimus, and the description of the first diplodocoid from East Asia (Lingwulong). Therefore, why not start off the new year with a post about titanosaur biogeography?
Averianov and Efimov (2018) describe the new titanosaur taxon Volgatitan simbirskiensis from Early Cretaceous (Hauterivian) marine shale deposits along the banks of the Volga River in Ulyanovsk Province, European Russia, making this taxon the oldest record of a titanosaur from Laurasia. In their phylogenetic analysis, the authors recover this taxon as sister to the titanosaur clade Lognkosauria, which is exclusively known from Patagonia (the Asian form Ruyangosaurus is considered a member of Lognkosauria by Sassani and Bivens 2017) On the other hand, the same authors recover another titanosaur taxon from European Russia, Tengrisaurus, as a member of Saltasauridae.
The phylogenetic results for Volgatitan obtained by Averianov and Efimov (2018) raise questions about whether the clade Lognkosauria evolved in Gondwana during the early Cretaceous or branched out into Gondwana from Laurasia later in the Early Cretaceous. Until recently, all bonafide lognkosaurians were known from Late Cretaceous deposits in Patagonia, including the recently described taxon Patagotitan, but Sassani and Bivens (2017) extended the geographic range of Lognkosauria to Laurasia by recovering the Aptian-Albian form Ruyangosaurus from Henan Province, eastern China as a sister-taxon of Puertasaurus. When discussing the results of the phylogenetic analysis of Volgatitan, Averianov and Efimov considered it likely that Lognkosauria had a widespread geographic distribution until the Late Cretaceous, when representatives of the group became confined to South America. They discounted the likelihood of a South American origin for Lognkosauria given the absence of members of this clade from South America before the Albian period. At first glance, it may be prudent to accept the conclusions by Averianov and Efimov (2018) that early lognkosaurs had a Laurasian origin in the Early Cretaceous given the age of Volgatitan as well as Ruyangosaurus.
Although the East African taxon Malawisaurus is often recovered as the basalmost lithostrotian by multiple phylogenetic analyses (e.g. Upchurch et al. 2004; Poropat et al. 2014; Averianov and Efimov 2018), the cladogram of Sassani and Bivens (2017) recovers Malawisaurus as more closely related to members of Lognkosauria, rather than at the base of Lithostrotia, suggesting that early lognkosaurians, or at least close relatives of the group, were widespread globally during the Early Cretaceous. Averianov and Efimov (2018) put the body mass estimate of Volgatitan at 38,157 pounds (17,308 kg) and Paul (2010) gives the weight for Malawisaurus as 22,000 pounds (10,000 kg), far lower than the body mass estimates for Patagonian lognkosaurs, suggesting that some Early Cretaceous lognkosaur relative from Gondwana may have dispersed into Laurasia to either develop dwarfism in insular environments or resort to gigantism in response to phytobiotic factors like tall gymnosperms. Given the dearth of titanosaur remains from the earliest Cretaceous (the genus Triunfosaurus from the Berriasian-Valanginian of Brazil was assigned to Titanosauria by Carvalho et al. 2017, but its placement was questioned by Poropat et al. 2017), but also the age of Malawisaurus and Ruyangosaurus, it seems highly parsimonious to conclude that the earliest lognkosaurs originated in Gondwana during the Early Cretaceous and dispersed to other parts of the world in that time frame, eventually developing gigantism later in the Cretaceous due to environmental influences.
Averianov, A. and Efimov, V. 2018. The oldest titanosaurian sauropod
of the Northern Hemisphere. Bio. Comm. 63(3): 145–162. https://doi.org/10.21638/spbu03.2018.301
Carvalho, I.D.S., Salgado, L., Lindoso, R.M., Araujo-Junior, H.I.D., Noguiera, F.C.C. & Soares, J.A., 2017. A new basal titanosaur (Dinosauria, Sauropoda) from the Lower Cretaceous of Brazil. Journal of South American Earth Sciences 75: 74–84.
Paul, G.S, 2010. The Princeton Field Guide to Dinosaurs. Princeton: Princeton University Press.
Poropat, S.F.; Upchurch, P.;
Mannion, P.D.; Hocknull, S.A.; Kear, B.P.; Sloan, T.; Sinapius, G.H.K.;
Elliott, D.A., 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. doi:10.1016/j.gr.2014.03.014.
Poropat, S.F., Nair, J.P., Syme, C.E., Mannion, P.D., Upchurch, P., Hocknull, S.A., Cook, A.G., Tischler, T.R., and Holland, T., 2017. Reappraisal of Austrosaurus mckillopi Longman, 1933 from the Allaru Mudstone of Queensland, Australia’s first named Cretaceous sauropod dinosaur. Alcheringa: An Australasian Journal of Palaeontology 41:4, 543-580, DOI: 10.1080/03115518.2017.1334826
2017. The Chinese colossus: an evaluation of the phylogeny of Ruyangosaurus giganteus and its implications for titanosaur evolution. PeerJ Preprints
Upchurch, P., Barrett, P. M., and
Dodson, P. 2004. Sauropoda. pp. 259−322 in: The Dinosauria. Second Edition,
edited by Weishampel, D. B., Dodson, P., and Osmolska, H. Berkeley, Los
Angeles, London: University of California Press.
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. 2018; 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) found "Apatosaurus" minimus shares with Camarasaurus and most somphospondyls six sacral vertebrae and widely splayed preacetabular lobes of the ilium, while noting 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., 2018) 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, supp. tab. 108) report that tall sacral neural spines occur among non-diplodocoid neosauropods. Since tall neural spines of the sacral vertebrae are present in non-neosauropod eusauropods, it could be parsimonious 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.
References:
Sauropoda. pp. 345-401. In: Weishampel DB, Dodson P, Osmólska H, (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.
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.
Discovery of a short-necked sauropod dinosaur from the Late Jurassic period of Patagonia. Nature 435:670-672.
Taylor, M.P., and Wedel, M.J., 2012. Re-evaluating "Apatosaurus" minimus, a bizarre Morrison Formation sauropod with diplodocoid and macronarian features. p. 23. In: Friedman, M., and Lloyd, M. (eds.), Programme and Abstracts, 60th Annual Symposium of Vertebrate Palaeontology and Comparative Anatomy, University of Oxford, Oxford, UK, September 10th-15th 2012.
Age-related characteristics found in a partial pelvis of Camarasaurus. pp. 18-186. In: Tidwell V, Carpenter K. (eds.) Thunder-lizards: the sauropodomorph dinosaurs. Bloomington: Indiana University Press.
015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda) PeerJ 3:e857 https://doi.org/10.7717/peerj.857
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.
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.
In their cladistic analyses of Diplodocoidea, Tschopp et al. (2015) found "Apatosaurus" minimus shares with Camarasaurus and most somphospondyls six sacral vertebrae and widely splayed preacetabular lobes of the ilium, while noting 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., 2018) and "Apatosaurus" minimus.
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 is that "Apatosaurus" minimus may be a derived macronarian related to Somphospondyli, 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 present in most 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., 2018. A new sauropod dinosaur from the Lower Cretaceous Ilek Formation,
Western Siberia, Russia. Geobios 51 (1): 1-14. 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
The relationships of Cetiosauriscus stewarti (Dinosauria; Sauropoda): implications for sauropod phylogeny. Journal of Vertebrate Paleontology 23:60A. Sauropoda. pp. 345-401. In: Weishampel DB, Dodson P, Osmólska H, (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.
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.
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, M.P., and Wedel, M.J., 2012. Re-evaluating "Apatosaurus" minimus, a bizarre Morrison Formation sauropod with diplodocoid and macronarian features. p. 23. In: Friedman, M., and Lloyd, M. (eds.), Programme and Abstracts, 60th Annual Symposium of Vertebrate Palaeontology and Comparative Anatomy, University of Oxford, Oxford, UK, September 10th-15th 2012.
Age-related characteristics found in a partial pelvis of Camarasaurus. pp. 18-186. In: Tidwell V, Carpenter K. (eds.) Thunder-lizards: the sauropodomorph dinosaurs. Bloomington: Indiana University Press.
015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda) PeerJ 3:e857 https://doi.org/10.7717/peerj.857
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.
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 discovery of the aralosaurin lambeosaurine hadrosaurid Canardia from southern France (Prieto-Marquez et al. 2013), the recognition of Pararhabdodon from Spain as closely related to the lambeosaurine Tsintaosaurus by (Prieto-Marquez and Wagner 2009), and 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 titanosaur braincase are also seen in several titanosaur taxa 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 immigrant taxon from Africa 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. In fact, the near-lack of sauropod remains from pre-Turonian Cretaceous sediments in Central Asia (see Weishampel et al. 2004) suggests that lithostrotians more primitive than Saltasauridae might have colonized Central Asia from Gondwana via rudimentary land bridges to Asia, as Garcia et al. (2010) note that Ampelosaurus and Lirainosaurus are more derived than Atsinganosaurus. Likewise, the placement of Ampelosaurus, Lirainosaurus, 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, V., Pereda Suberbiola, X., and Sanz, J.L., 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
Díez Díaz, V., Mocho, P., Páramo, A., Escaso, F., Marcos-Fernández, F., Sanz, J.L., and Ortega, F., 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.
Garcia, G., Amico, S., Fournier, F., Thouand, E., and Valentin, X., 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.
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 titanosaur braincase are also seen in several titanosaur taxa 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 immigrant taxon from Africa 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. In fact, the near-lack of sauropod remains from pre-Turonian Cretaceous sediments in Central Asia (see Weishampel et al. 2004) suggests that lithostrotians more primitive than Saltasauridae might have colonized Central Asia from Gondwana via rudimentary land bridges to Asia, as Garcia et al. (2010) note that Ampelosaurus and Lirainosaurus are more derived than Atsinganosaurus. Likewise, the placement of Ampelosaurus, Lirainosaurus, 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, V., Pereda Suberbiola, X., and Sanz, J.L., 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
Díez Díaz, V., Mocho, P., Páramo, A., Escaso, F., Marcos-Fernández, F., Sanz, J.L., and Ortega, F., 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.
Garcia, G., Amico, S., Fournier, F., Thouand, E., and Valentin, X., 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.
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