Life and reproduction of titanosaurians: Isotopic hallmark of mid-palaeolatitude eggshells and its significance for body temperature, diet, and nesting

Abstract Eggshells represent an important part of the fossil record of Titanosauria (Dinosauria – Neosauropoda) and their stable isotope compositions are valuable palaeoenvironmental proxies. A new set of conventional (δ18O and δ13C) and clumped (Δ47) stable isotope compositions of titanosaurian eggshells is presented, together with that of a bone and a single associated tooth, sampled in three Late Cretaceous nesting sites from La Rioja Province, NW Argentina. The preservation state of the fossils was first evaluated using optical and analytical techniques, such as transmitted light and optical cathodoluminescence (CL) microscopy, energy dispersive X-ray spectroscopy (EDX), and laser ablation-inductively coupled plasma-mass spectrometry (LA-ICP-MS). The isotopic compositions of the fossils were then compared to those of associated carbonate rocks and nodules, hydrothermal calcite and quartz, and those reported for eggshells from other nesting sites worldwide. This large, combined sample set allows us to define an isotopic hallmark (δ13CVPDB = −15 to −11‰; δ18OVSMOW = 27 to 33‰) typical for well-preserved mid-palaeolatitude titanosaurian eggshells. This hallmark is intended to identify the oological specimens best suited for palaeoenvironmental reconstructions, for instance in museum collection samples that may lack associated abiogenic materials such as host rocks. In addition, our isotopic data support that titanosaurians were animals with an elevated body temperature, mainly feeding on C3 plants, and reproducing under conditions more arid than the long-term average. The data are in excellent agreement with the isotopic data reported from other mid-palaeolatitude nesting sites around the world, indicating that titanosaurians needed similar environmental conditions to reproduce, regardless of the palaeogeographic location of their habitat. Finally, we raise the question whether titanosaurians experienced reproduction-related fasting, as noted for several extant vertebrates, and discuss the complexity of interpreting Δ47-derived temperatures, despite very consistent bulk isotopic data.