Gerald Grellet-Tinner

Digital preparation of a probable neoceratopsian preserved within an egg, with comments on microstructural anatomy of ornithischian eggshells

We describe the first known embryo of a neoceratopsian dinosaur, perhaps the most ubiquitous Laurasian group of Cretaceous dinosaurs, which is preserved completely enclosed within an egg. This specimen was collected from Late Cretaceous beds of southern Mongolia, which commonly preserve fossils of the neoceratopsian, Yamaceratops dorngobiensis. The small egg was scanned using high-resolution X-ray computed tomography and digitally prepared from the matrix. The preserved and imaged elements support a diagnosis of the embryo to Neoceratopsia and allow preliminary observations of ontogenetic transformations within this group. The addition of an embryo also adds another important data point to the already impressive postnatal ontogenetic series that are available for this clade.

Phylogenetic interpretation of eggs and eggshells: implications for phylogeny of Palaeognathae

This oological analysis supports Palaeognathae monophyly, and places Dromornithidae within Neognathae, the sister clade of Paleognathae. This Dromornithidae relationship is congruent with a recent phylogenetic hypothesis, based on cranial characters, that differs from the original assignment of this group of extinct birds to the Palaeognathae. Minor differences exist between previous palaeognath phylogenetic hypotheses and that obtained from this research. Some of the topological disparities might be influenced solely by inclusion of more fossil taxa in this analysis compared with those considering either molecular or skeletal characters. Surprisingly, oological data differentiate the Australasian palaeognath taxa from those of Africa and South America; this phylogenetic topology conforms to the paleognath geographic distribution. Regardless of the issues related to Paleognathae, results show that oological characters permit the differentiation of groups at a generic level, thus validating the use of eggs and eggshells as a source of evolutionary characters.

Palaeobiology: Argentinian unhatched pterosaur fossil

Our knowledge of the eggs and embryos of pterosaurs, the Mesozoic flying reptiles, is sparse. Until now, the recent discovery of an ornithocheirid embryo from 121-million-year-old rocks in China constituted the only reliable evidence of an unhatched pterosaur. Here we describe an embryonic fossil of a different pterosaur from the Early Cretaceous lacustrine deposits of Loma del Pterodaustro (the Lagarcito Formation, which is about 100 million years old) in central Argentina. This new fossil provides insight into the eggshell morphology, early growth and nesting environments of pterosaurs.

A new Argentinean nesting site showing neosauropod dinosaur reproduction in a Cretaceous hydrothermal environment

Although several late Cretaceous sauropod colonial nesting sites have been discovered nearly on every continent during the last few decades, no studies have been performed to determine the factors that underpinned the choice of these specific sites. Here, we report the first definitive evidence of a group of sauropods that nested repetitively and purposely at a Cretaceous hydrothermal site at Sanagasta, La Rioja Province, Argentina. The discovery of this new colonial nesting locality shows nest fidelity over a long time, and a symbiotic relationship between egg clutches and a peculiar hydrothermal environment that favoured their incubation. Sedimentary and geochemical analyses of 80 clutches and their large eggs with thick eggshells substantiate that the Sanagasta sauropods were specifically using the soil moisture and thermoradiance to incubate their eggs, similar to a few extant species, namely, the megapode, Megapodius pritchardii, which is known to lay its egg clutches in burrows at volcanically heated nesting grounds.

Water Vapor Conductance of the Lower Cretaceous Dinosaurian Eggs from Sanagasta, La Rioja, Argentina: Paleobiological and Paleoecological Implications for South American Faveoloolithid and Megaloolithid Eggs

ABSTRACT The water vapor conductance (GH2O) of the neosauropod eggs from the Lower Cretaceous Sanagasta nesting site in La Rioja Province, Argentina, was examined and compared with other Cretaceous Argentinean oological material. The 2900 mgH2O/day·Torr GH2O of the Sanagasta eggshells confirms an extremely moist nesting environment and supports field observations of dug-out nests in a geothermal setting. The observed thinning of the outer eggshell surface during incubation increases gas conductance and concomitantly decreases eggshell mechanical resistance during the late ontogenetic stages, thus facilitating embryonic development and hatching. The Sanagasta and Entre Ríos Province faveoloolithid eggs display the highest and comparable GH2O values and share several morphological and diagenetic characters, indicating comparable nesting strategy in geothermal settings. However, the faveoloolithid Yaminué and La Pampa Province specimens cluster together with lower GH2O values closer to the megaloolithid eggs. The GH2O of the megaloolithid egg Megaloolithus patagonicus was reconsidered and new results are now congruent with other reported megaloolithid GH2O values. Additionally, we hypothesize that Y-shaped pore canals of M. patagonicus, which upper sections reach only the top third or half eggshell thickness and, a wider section in the middle would not compromise the overall egg mechanical resistance like vertical pores connecting directly the outer to the inner eggshell surfaces. Such pore spatial arrangement and geometry would enhance, as the eggshell thins during incubation, a greater GH2O, GO2 and GCO2 and facilitate embryonic development in high moisture nesting contents. Overall, data suggests that neosauropod nesting and brooding behaviors were dependent on elevated moisture nesting environments.

First Evidence of Reproductive Adaptation to “Island Effect” of a Dwarf Cretaceous Romanian Titanosaur, with Embryonic Integument In Ovo

Background The Cretaceous vertebrate assemblages of Romania are famous for geographically endemic dwarfed dinosaur taxa. We report the first complete egg clutches of a dwarf lithostrotian titanosaur, from Toteşti, Romania, and its reproductive adaptation to the “island effect”. Methodology/Findings The egg clutches were discovered in sequential sedimentary layers of the Maastrichtian Sânpetru Formation, Toteşti. The occurrence of 11 homogenous clutches in successive strata suggests philopatry by the same dinosaur species, which laid clutches averaging four ∼12 cm diameters eggs. The eggs and eggshells display numerous characters shared with the positively identified material from egg-bearing level 4 of the Auca Mahuevo (Patagonia, Argentina) nemegtosaurid lithostrotian nesting site. Microscopic embryonic integument with bacterial evidences was recovered in one egg. The millimeter-size embryonic integument displays micron size dermal papillae implying an early embryological stage at the time of death, likely corresponding to early organogenesis before the skeleton formation. Conclusions/Significance The shared oological characters between the Haţeg specimens and their mainland relatives suggest a highly conservative reproductive template, while the nest decrease in egg numbers per clutch may reflect an adaptive trait to a smaller body size due to the “island effect”. The combined presence of the lithostrotian egg and its embryo in the Early Cretaceous Gobi coupled with the oological similarities between the Haţeg and Auca Mahuevo oological material evidence that several titanosaur species migrated from Gondwana through the Haţeg Island before or during the Aptian/Albian. It also suggests that this island might have had episodic land bridges with the rest of the European archipelago and Asia deep into the Cretaceous.

Minute theropod eggs and embryo from the Lower Cretaceous of Thailand and the dinosaur-bird transition

We report on very small fossil eggs from the Lower Cretaceous of Thailand, one of them containing a theropod embryo, which display a remarkable mosaic of characters. While the surficial ornamentation is typical of non-avian saurischian dinosaurs, the three-layered prismatic structure of the eggshell is currently known only in extant and fossil eggs associated with birds. These eggs, about the size of a goldfinchs, mirror at the reproductive level the retention of small body size that was paramount in the transition from non-avian theropods to birds. The egg-layer may have been a small feathered theropod similar to those recently found in China.

Isotopic ordering in eggshells reflects body temperatures and suggests differing thermophysiology in two Cretaceous dinosaurs

Our understanding of the evolutionary transitions leading to the modern endothermic state of birds and mammals is incomplete, partly because tools available to study the thermophysiology of extinct vertebrates are limited. Here we show that clumped isotope analysis of eggshells can be used to determine body temperatures of females during periods of ovulation. Late Cretaceous titanosaurid eggshells yield temperatures similar to large modern endotherms. In contrast, oviraptorid eggshells yield temperatures lower than most modern endotherms but ∼ 6 °C higher than co-occurring abiogenic carbonates, implying that this taxon did not have thermoregulation comparable to modern birds, but was able to elevate its body temperature above environmental temperatures. Therefore, we observe no strong evidence for end-member ectothermy or endothermy in the species examined. Body temperatures for these two species indicate that variable thermoregulation likely existed among the non-avian dinosaurs and that not all dinosaurs had body temperatures in the range of that seen in modern birds.

The First Occurrence in the Fossil Record of an Aquatic Avian Twig-Nest with Phoenicopteriformes Eggs: Evolutionary Implications

Background We describe the first occurrence in the fossil record of an aquatic avian twig-nest with five eggs in situ (Early Miocene Tudela Formation, Ebro Basin, Spain). Extensive outcrops of this formation reveal autochthonous avian osteological and oological fossils that represent a single taxon identified as a basal phoenicopterid. Although the eggshell structure is definitively phoenicopterid, the characteristics of both the nest and the eggs are similar to those of modern grebes. These observations allow us to address the origin of the disparities between the sister taxa Podicipedidae and Phoenicopteridae crown clades, and traces the evolution of the nesting and reproductive environments for phoenicopteriforms. Methodology/Principal Findings Multi-disciplinary analyses performed on fossilized vegetation and eggshells from the eggs in the nest and its embedding sediments indicate that this new phoenicopterid thrived under a semi-arid climate in an oligohaline (seasonally mesohaline) shallow endorheic lacustine environment. High-end microcharacterizations including SEM, TEM, and EBSD techniques were pivotal to identifying these phoenicopterid eggshells. Anatomical comparisons of the fossil bones with those of Phoenicopteriformes and Podicipediformes crown clades and extinct palaelodids confirm that this avian fossil assemblage belongs to a new and basal phoenicopterid. Conclusions/Significance Although the Podicipediformes-Phoenicopteriformes sister group relationship is now well supported, flamingos and grebes exhibit feeding, reproductive, and nesting strategies that diverge significantly. Our multi-disciplinary study is the first to reveal that the phoenicopteriform reproductive behaviour, nesting ecology and nest characteristics derived from grebe-like type strategies to reach the extremely specialized conditions observed in modern flamingo crown groups. Furthermore, our study enables us to map ecological and reproductive characters on the Phoenicopteriformes evolutionary lineage. Our results demonstrate that the nesting paleoenvironments of flamingos were closely linked to the unique ecology of this locality, which is a direct result of special climatic (high evaporitic regime) and geological (fault system) conditions.

A note on pterosaur nesting behavior

Based on examination of eggshell structure and predicted vapor conductances in eggshells in recently described material from Argentina and China we conclude that pterosaurs buried their eggs. Egg-burying imposes theoretical restrictions on the distribution of pterosaurs, both geographically and spatially, raises the possibility of thermal sex determination and supports previous suggestions that they exhibited nesting fidelity. Some features associated with egg-burying, such as weight savings, are likely to have been fortuitous pre-adaptations for these flying reptiles, but others may have disadvantaged them relative to avian competitors or increased their vulnerability to extinction in a cooling climate.