Evidence for heterothermic endothermy and reptile-like eggshell mineralization inTroodon, a non-avian maniraptoran theropod

Abstract

The dinosaur–bird transition involved several anatomical, biomechanical, and physiological modifications of the theropod bauplan. Non-avian maniraptoran theropods, such as Troodon , are key to better understand changes in thermophysiology and reproduction occurring during this transition. Here, we applied dual clumped isotope (Δ 47 and Δ 48 ) thermometry, a technique that resolves mineralization temperature and other nonthermal information recorded in carbonates, to eggshells from Troodon , modern reptiles, and modern birds. Troodon eggshells show variable temperatures, namely 42 and 29 ± 2 °C, supporting the hypothesis of an endothermic thermophysiology with a heterothermic strategy for this extinct taxon. Dual clumped isotope data also reveal physiological differences in the reproductive systems between Troodon , reptiles, and birds. Troodon and modern reptiles mineralize their eggshells indistinguishable from dual clumped isotope equilibrium, while birds precipitate eggshells characterized by a positive disequilibrium offset in Δ 48 . Analyses of inorganic calcites suggest that the observed disequilibrium pattern in birds is linked to an amorphous calcium carbonate (ACC) precursor, a carbonate phase known to accelerate eggshell formation in birds. Lack of disequilibrium patterns in reptile and Troodon eggshells implies these vertebrates had not acquired the fast, ACC-based eggshell calcification process characteristic of birds. Observation that Troodon retained a slow reptile-like calcification suggests that it possessed two functional ovaries and was limited in the number of eggs it could produce; thus its large clutches would have been laid by several females. Dual clumped isotope analysis of eggshells of extinct vertebrates sheds light on physiological information otherwise inaccessible in the fossil record.

Resolving and correcting for kinetic biases on methane seep paleotemperature using carbonate ∆ 47 /∆ 48 analysis

“…Because of the variable expression of KIEs, calculating paleotemperatures and interpreting the physical-chemical state of relic seep systems would yield ambiguous results dependent upon many assumptions regarding the magnitude of these offsets. Recent advances in clumped isotopes ( 30 ) have shown that with the analyses of multiple isotopologues at mass/charge ratio ( m / z ) = 47 (∆ 47 ) and m / z = 48 (∆ 48 ), the effects of these kinetic processes can be resolved ( 31 – 33 ), allowing for more confident reconstruction of temperatures ( 34 – 36 ), as well as identifying diagenetic effects ( 33 , 37 , 38 ), and fingerprinting the processes that caused the kinetic bias ( 26 , 27 , 32 , 35 , 38 , 39 ).…”