Microstructural and crystallographic evolution of palaeognath (Aves) eggshells.

The avian palaeognath phylogeny has been recently revised significantly due to the advancement of genome-wide comparative analyses and provides the opportunity to trace the evolution of the microstructure and crystallography of modern dinosaur eggshells. Here, eggshells of all major clades of Palaeognathae (including extinct taxa) and selected eggshells of Neognathae and non-avian dinosaurs are analysed with electron backscatter diffraction. Our results show the detailed microstructures and crystallographies of (previously) loosely categorized ostrich-, rhea-, and tinamou-style morphotypes of palaeognath eggshells. All rhea-style eggshell appears homologous, while respective ostrich-style and tinamou-style morphotypes are best interpreted as homoplastic morphologies (independently acquired). Ancestral state reconstruction and parsimony analysis additionally show that rhea-style eggshell represents the ancestral state of palaeognath eggshells both in microstructure and crystallography. The ornithological and palaeontological implications of the current study are not only helpful for the understanding of evolution of modern and extinct dinosaur eggshells, but also aid other disciplines where palaeognath eggshells provide useful archive for comparative contrasts (e.g. palaeoenvironmental reconstructions, geochronology, and zooarchaeology).

About 50 species of birds on the planet today do not belong to the same group as the other 10,000 currently in existence. Known as the paleognaths, this small clade features many of the largest and heaviest avian specimens on Earth, bringing together ostriches and their distant South American relatives the rheas, as well as emus and cassowaries. Kiwis and ground-dwelling species known as tinamous complete the family. None of these birds can fly, except for the tinamous. Paleognath eggs are also somewhat distinct from the rest of the avian population, being larger and sporting thicker shells. Advanced genetic analyses in the late 2000's have upended researchers' understanding of in what sequence these birds have evolved, and how they are related to each other. The new phylogenetic family tree offers the opportunity to re-evaluate previous conclusions about this group, which could in turn clarify the evolution and lifestyle of flightless modern and extinct dinosaurs. Choi et al. decided to use this updated genetic information to better understand how paleognath eggs have evolved. Traditionally, these have been loosely classified into three types (rhea-style, ostrich-style and tinamou-style) based on various morphological features. Their microstructure, however, remains poorly studied, and it is unclear whether this categorisation reflects evolutionary processes. Aiming to fill this gap, Choi et al. employed electron microscopy approaches to examine the microstructure of the eggshell in all groups of paleognath birds (including the now extinct moas from New Zealand and elephant birds from Madagascar), as well as in selected species of flying birds and non-avian dinosaurs. Combined with the new evolutionary tree and additional analyses, these experiments suggest that the ancestor of the paleognaths laid rhea-style eggs, which are still the most common type amongst the family. In fact, several non-paleognath bird eggs also showed these features. In contrast, ostrich-style and tinamou-style eggs seem to have evolved independently in several distantly related species within the group. Equipped with this knowledge, it may become possible for ornithologists to decipher how eggshells evolved in other lineages of flightless birds, and for palaeontologists to better interpret fossil bird and other dinosaur eggs.

A comparative study of eggshells of Gekkota with morphological, chemical compositional and crystallographic approaches and its evolutionary implications.

The Gekkota is an important clade in the evolution of calcified eggshells in that some of its families lay rigid eggshells like archosaurs. However, the fundamental differences and similarities between the mechanism of rigid eggshell formation of the Gekkota and Archosauria have not been investigated thoroughly due to the lack of knowledge of gekkotan eggshells. Here, we report for the first time a comprehensive analysis of morphological, chemical compositional, and crystallographic features of rigid and soft gekkotan eggshells. Exhaustive morphological description provided common characters for gekkotan eggshells, as well as unique features of each species. We found that elemental distribution of rigid gekkotan eggshells is different from that of avian eggshells, especially in the case of Mg and P. In addition, the crystallographic features (size, shape, and alignment of calcite grains) of gekkotan eggshells are completely different from those of archosaur eggshells. The result of this study suggests that soft gekkotan eggshells are morphologically more similar to tuatara eggshells rather than soft eggshells of derived squamates. The chemical compositional analysis suggests that the eggshell may act as a mineral reservoir for P and F as well as Ca. More importantly, all chemical compositions and crystallographic features imply that the gekkotan eggshell formation may begin at the outer surface and growing down to the inner surface, which is opposite to the direction of the archosaur eggshell formation. This character would be crucial for identifying fossil gekkotan eggs, which are poorly known in paleontology. All these lines of evidence support that soft gekkotan and tuatara eggshells share the primitive characters of all lepidosaurid eggshells. Finally, gekkotan and archosaur rigid eggshells represent a typical example of convergent evolution in the lineage of the Sauropsida.