Browsing by Author "Choi, Seung"

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The diverse terminology of reptile eggshell microstructure and its effect on phylogenetic comparative analyses
(Wiley, 2022-06) Legendre, Lucas J.; Choi, Seung; Clarke, Julia A.
Reptile eggshell ensures water and gas exchange during incubation and plays a key role in reproductive success. The diversity of reptilian incubation and life history strategies has led to many clade-specific structural adaptations of their eggshell, which have been studied in extant taxa (i.e. birds, crocodilians, turtles, and lepidosaurs). Most studies on non-avian eggshells were performed over 30 years ago and categorized reptile eggshells into two main types: “hard” and “soft” – sometimes with a third intermediate category, “semi-rigid.” In recent years, however, debate over the evolution of eggshell structure of major reptile clades has revealed how definitions of hard and soft eggshells influence inferred deep-time evolutionary patterns. Here, we review the diversity of extant and fossil eggshell with a focus on major reptile clades, and the criteria that have been used to define hard, soft, and semi-rigid eggshells. We show that all scoring approaches that retain these categories discretize continuous quantitative traits (e.g. eggshell thickness) and do not consider independent variation of other functionally important microstructural traits (e.g. degree of calcification, shell unit inner structure). We demonstrate the effect of three published approaches to discretizing eggshell type into hard, semi-rigid, and soft on ancestral state reconstructions using 200+ species representing all major extant and extinct reptile clades. These approaches result in different ancestral states for all major clades including Archosauria and Dinosauria, despite a difference in scoring for only 1–4% of the sample. Proposed scenarios of reptile eggshell evolution are highly conditioned by sampling, tree calibration, and lack of congruence between definitions of eggshell type. We conclude that the traditional “soft/hard/semi-rigid” classification of reptilian eggshells should be abandoned and provide guidelines for future descriptions focusing on specific functionally relevant characteristics (e.g. inner structures of shell units, pores, and membrane elements), analyses of these traits in a phylogenetic context, and sampling of previously undescribed taxa, including fossil eggs.
Fossil turtle eggs from the Upper Cretaceous Gaogou Formation, Xiaguan-Gaoqiu Basin, Neixiang County, Henan Province, China: Interpretation of the transformation from aragonite to calcite in fossil turtle eggshell
(Elsevier BV, 2022-06) Xu, Li; Xie, Junfang; Zhang, Shukang; Choi, Seung; Kim, Noe-Heon; Gao, Diansong; Jin, Xingsheng; Jia, Songhai; Gao, Yongli
In 2007, a clutch of approximately 30 fossil eggs reported from the Upper Cretaceous Gaogou Formation, Xiaguan-Gaoqiu Basin, Neixiang County, Henan Province, China was assigned to the oofamily “Crocodiloolithidae”. After 11 years, another clutch of 15 eggs of the same type associated with Mosaiceratops azumai and from the same county was identified as turtle eggs (Testudoolithus), based on the needle-like crystals interpreted as aragonite from their eggshells. Our detailed study of the clutch of 15 eggs and six clutches of the same ootaxon using transmitted/polarized light microscope, followed by scanning electron microscope observations combined with electron backscatter diffraction analysis showed that their eggshells are dominated by secondary calcite; with a few pristine aragonite crystals that unequivocally support the turtle affinity of the eggs. Furthermore, superimposed cone-shaped structural units with radial ultrastructures, combined with the extremely thick eggshell challenges the assignment to the oogenus Testudoolithus. Hence, we suggest that this ootaxon could represent a new oogenus of Testudoolithidae. Although additional eggshell units were reported in the abnormal fossil turtle egg with multilayered eggshell from the Judith River Formation in Montana, U.S.A, the relatively constant eggshell thickness indicates that the eggs from Neixiang are not pathological. Therefore, this is the first report of superimposed structural units inside normal turtle eggshells, revealing the unique eggshell formation mechanism of the extinct turtle from Neixiang. Moreover, the aragonite discovered in the eggs may extend the temporal record of confirmed aragonite in turtle eggs. The crystallographic comparison between the eggs from Neixiang and the fossil turtle eggshells from Zhejiang Province and Montana revealed the transformation process from aragonite to calcite in fossil turtle eggshell.
Microstructural and crystallographic evolution of palaeognath (Aves) eggshells
(eLife Sciences Publications, Ltd, 2023-01) Choi, Seung; Hauber, Mark E.; Legendre, Lucas J.; Kim, Noe-Heon; Lee, Yuong-Nam; Varricchio, David J.
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).
Tiny, ornamented eggs and eggshell from the Upper Cretaceous of Utah represent a new ootaxon with theropod affinities
(Springer Science and Business Media LLC, 2021-05) Oser, Sara E.; Chin, Karen; Sertich, Sertich; Varricchio, David J.; Choi, Seung; Rifkin, Jeffrey
A new Cretaceous ootaxon (eggshell type) from the Kaiparowits Formation of Grand Staircase-Escalante National Monument is among a growing number of very small eggs described from the Mesozoic. Analyses of two partial eggs (~ 17.7 mm in diameter) and 29 eggshell fragments reveal that this new ootaxon exhibits nodose ornamentation with distinctive branching pore canals that open atop the nodes. Its two-layered microstructure consists of a mammillary layer and a continuous layer with rugged grain boundaries between calcite grains. Although the exact identity of the egg producer is unknown, the eggshell microstructure and small size is consistent with a small-bodied avian or non-avian theropod. The specific combination of small egg size, branching pores, two-layered microstructure, and dispersituberculate ornamentation preserved in this new ootaxon is unique among theropod eggs. This underscores that both eggshell and skeletal fossils of Cretaceous theropods can display a mosaic of transitional morphological and behavioural features characteristic of both avian and non-avian taxa. As such, this new ootaxon increases the diversity of Cretaceous eggs and informs our understanding of the evolution of theropod eggshell microstructure and morphology.