Bone histology of the titanosaur Lirainosaurus astibiae (Dinosauria: Sauropoda) from the Latest Cretaceous of Spain

The titanosaur Lirainosaurus astibiae is the only sauropod species known from the Late Cretaceous of the Iberian Peninsula. Lirainosaurus did not reach a gigantic body size and is one of the smallest sauropods discovered to date. Histological analysis of Lirainosaurus bones, focused on diaphyseal transverse sections of appendicular elements, reveals that Lirainosaurus did not exhibit the osseous microstructure typical for large sauropods, but is comparable with that of the coeval titanosaurs Alamosaurus sanjuanensis, Ampelosaurus atacis, and Magyarosaurus dacus, and also shares histological traits with other small to medium-sized sauropodomorph dinosaurs. Lirainosaurus limb bones exhibit a laminar fibrolamellar bone microstructure interrupted by growth marks, fully obliterated in adulthood by intense secondary remodeling processes which tend to replace completely the primary cortex. Lirainosaurus attained smaller sizes than typical sauropods reducing the rate of primary periosteal osteogenesis and developing an extensive secondary remodeling well before the adult size was reached. Histological organization of Lirainosaurus long bones is more mature than observed in basal neosauropods at similar ontogenetic stage, documenting a case of peramorphosis by pre-displacement. This heterochronic growth would be a reversal of the accelerated pattern of bone deposition typical for the sauropod lineage.