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.
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