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Item The egg-thief architect: experimental oviraptorosaur nesting physiology, the possibility of adult-mediated incubation, and the feasibility of indirect contact incubation(Cambridge University Press, 2023-08) Hogan, Jason D.Numerous, high-quality reproduction-related oviraptorosaur fossils have been described. However, oviraptorosaur-style nests are unknown among extant animals, and their curious construction makes nesting behavior difficult to interpret. Experiments were undertaken to better understand oviraptorosaur nesting strategies. A surrogate was constructed and placed atop mock-oviraptorosaur nests built from sand and 36 infertile emu eggs (as Macroolithus approximations) arranged according to the most current nest reconstructions. Thermometers, placed within each egg and throughout the experimental area, recorded energy flow from the surrogate dinosaur into the nesting microenvironment. One experiment examined a basic open nest warmed from above; the second, a fully buried clutch warmed from above; and the third, a nest open like the first but with heating elements (representing hindlimbs) extending down into the nest. It was found that egg temperatures in each scenario surpassed ambient temperatures without requiring excessive energy input. Final clutch temperatures were below most avian values, closer to crocodilian incubation, but are likely conservative, considering experimental parameters. These results may support the idea that an oviraptorosaur could use adult-generated energy to warm a clutch above ambient conditions. Additionally, egg tiers would be warmer and more uniform in temperature if heated by elements within the nest, such as hindlimbs, instead of solely from above. Results from the second experiment indicate that an endothermic adult could possibly warm a clutch fully buried beneath itself despite a barrier. Although not likely a behavior exhibited by oviraptorosaurs, such results suggest an important evolutionary step bridging guarded subterranean eggs and contact-incubated subaerial eggs.Item Chthonic severance: dinosaur eggs of the Mesozoic, the significance of partially buried eggs and contact incubation precursors(The Royal Society, 2023-07) Hogan, Jason D.; Varricchio, David J.For most dinosaurs, clutches consisted of a single layer of spherical to sub-spherical, highly porous eggs that were probably fully buried. Both eggs and clutch form change drastically with pennaraptoran theropods, the clade that includes birds. Here, far less porous, more elongate eggs are arranged with additional complexity, and only partially buried. While partial egg burial seems to be effective for an extremely small group of modern birds, the behaviour's overall rarity complicates our understanding of Mesozoic analogies. Recent experimental examination of pennaraptoran nesting thermodynamics suggests that partial egg burial, combined with contact incubation, may be more efficacious than has been presumed. We propose that nest guarding behaviour by endothermic archosaurs may have led to an indirect form of contact incubation using metabolic energy to affect temperature change in a buried clutch through a barrier of sediment, which in turn may have selected for shallower clutch burial to increasingly benefit from adult-generated energy until partial egg exposure. Once partially exposed, continued selection pressure may have aided a transition to fully subaerial eggs. This hypothesis connects the presence of partially buried dinosaurian clutches with the transition from basal, crocodile-like nesting (buried clutches guarded by adults) to the dominant avian habit of contact incubating fully exposed eggs.Item Do paleontologists dream of electric dinosaurs? Investigating the presumed inefficiency of dinosaurs contact incubating partially buried eggs(Cambridge University Press, 2021-01) Hogan, Jason D.; Varricchio, David J.Troodon formosus, a theropod from the Late Cretaceous, is one of the few species of dinosaurs with multiple nest sites uncovered. It has been consistently demonstrated that eggs within these nests would have been partially buried in life—an exceedingly rare state in modern vertebrates. There has been debate over Troodon's capacity to engage in thermoregulatory contact incubation, especially regarding an adult's ability to efficiently supply partially buried eggs with energy. An actualistic investigation was undertaken to determine the thermodynamic efficiency of contact incubating partially buried eggs. An efficient system would keep eggs at temperatures closer to the surrogate parent than the ambient, without prohibitively high energy input. For the experiment, a surrogate dinosaur was created and used in both indoor controlled ambient temperature trials and in an outdoor variant. Even with ambient temperatures that were likely cooler than Cretaceous averages, the results showed that contact incubating partially buried eggs did seem to confer an energetic advantage; egg temperatures remained closer to the surrogate than ambient in both indoor and outdoor tests. Still, critics of contact incubating partially buried eggs are correct in that there is a depth at which adult energy would fail to make much of an impact—perhaps more relevant to buried eggs, as partially buried eggs would be in contact with an adult and likely above the thermal input threshold. Additionally, results from this experiment provide evidence for a possible evolutionary path from guarding behavior to thermoregulatory contact incubation.Item Do paleontologists dream of electric dinosaurs? Investigating the presumed inefficiency of dinosaurs contact incubating partially buried eggs(2020-10) Hogan, Jason D.; Varricchio, David J.Troodon formosus, a theropod from the Late Cretaceous, is one of the few species of dinosaurs with multiple nest sites uncovered. It has been consistently demonstrated that eggs within these nests would have been partially buried in life—an exceedingly rare state in modern vertebrates. There has been debate over Troodon's capacity to engage in thermoregulatory contact incubation, especially regarding an adult's ability to efficiently supply partially buried eggs with energy. An actualistic investigation was undertaken to determine the thermodynamic efficiency of contact incubating partially buried eggs. An efficient system would keep eggs at temperatures closer to the surrogate parent than the ambient, without prohibitively high energy input. For the experiment, a surrogate dinosaur was created and used in both indoor controlled ambient temperature trials and in an outdoor variant. Even with ambient temperatures that were likely cooler than Cretaceous averages, the results showed that contact incubating partially buried eggs did seem to confer an energetic advantage; egg temperatures remained closer to the surrogate than ambient in both indoor and outdoor tests. Still, critics of contact incubating partially buried eggs are correct in that there is a depth at which adult energy would fail to make much of an impact—perhaps more relevant to buried eggs, as partially buried eggs would be in contact with an adult and likely above the thermal input threshold. Additionally, results from this experiment provide evidence for a possible evolutionary path from guarding behavior to thermoregulatory contact incubation.