In 1980, Steven J. Gould published an essay on the emergence of paleobiology as a nomothetic discipline (Gould 1980), nomothetism referring to the search for general laws or principles. Gould contrasted this with the foundation of paleontology, the idiographic tradition of detailing the history of life from the description of new fossil taxa to the elucidation of the long-term patterns of change through time. Among the pioneers of this nomothetic expansion was Jim Valentine. Here I pay tribute to Jim as one of the first paleobiologists, a colleague, coauthor, and friend, emphasizing his intellectual style and insights as much as his lasting contributions. I have written this in part as a eulogy, a remembrance for those who knew him, but also as an introduction to the continuing relevance of his work for those who may be unfamiliar with it.

Data compilations expand the scope of research; however, data citation practice lags behind advances in data use. It remains uncommon for data users to credit data producers in professionally meaningful ways. In paleontology, databases like the Paleobiology Database (PBDB) enable assessment of patterns and processes spanning millions of years, up to global scale. The status quo for data citation creates an imbalance wherein publications drawing data from the PBDB receive significantly more citations (median: 4.3 ± 3.5 citations/year) than the publications producing the data (1.4 ± 1.3 citations/year). By accounting for data reuse where citations were neglected, the projected citation rate for data-provisioning publications approached parity (4.2 ± 2.2 citations/year) and the impact factor of paleontological journals (n = 55) increased by an average of 13.4% (maximum increase = 57.8%) in 2019. Without rebalancing the distribution of scientific credit, emerging “big data” research in paleontology—and science in general—is at risk of undercutting itself through a systematic devaluation of the work that is foundational to the discipline.

The fossil record is spatiotemporally heterogeneous: taxon occurrence data have patchy spatial distributions, and this patchiness varies through time. Large-scale quantitative paleobiology studies that fail to account for heterogeneous sampling coverage will generate uninformative inferences at best and confidently draw wrong conclusions at worst. Explicitly spatial methods of standardization are necessary for analyses of large-scale fossil datasets, because nonspatial sample standardization, such as diversity rarefaction, is insufficient to reduce the signal of varying spatial coverage through time or between environments and clades. Spatial standardization should control both geographic area and dispersion (spread) of fossil localities. In addition to standardizing the spatial distribution of data, other factors may be standardized, including environmental heterogeneity or the number of publications or field collecting units that report taxon occurrences. Using a case study of published global Paleobiology Database occurrences, we demonstrate strong signals of sampling; without spatial standardization, these sampling signatures could be misattributed to biological processes. We discuss practical issues of implementing spatial standardization via subsampling and present the new R package divvy to improve the accessibility of spatial analysis. The software provides three spatial subsampling approaches, as well as related tools to quantify spatial coverage. After reviewing the theory, practice, and history of equalizing spatial coverage between data comparison groups, we outline priority areas to improve related data collection, analysis, and reporting practices in paleobiology.

Women are underrepresented in paleontology. Despite more women students, representation at senior levels remains low. To advance professionally, scientists must disseminate their research through peer-reviewed publications. We examine gendered authorship patterns in Paleobiology to ask whether the publishing infrastructure supports the Paleontological Society's gender-equity goals. We reviewed all papers published in Paleobiology from its inception in 1975 through 2021. For each paper, we recorded each author, the author's position in the author list, and the total number of authors on each paper. We coded gender based on a combination of personal communication and pronouns used in publicly available information. We compared author demographics with anonymized membership data from the Paleontological Society. Over the journal's run, the number of authors per paper increased due to cultural shifts toward collaborative work and acknowledging student contributions with coauthorship. These trends contribute to proportionally more women authors, beginning in the early 2000s. Despite these increases, women remain chronically underrepresented. In 2018, 2019, and 2021, the proportion of women authors in Paleobiology paralleled membership in the Paleontological Society. However, in 2020, Paleobiology published fewer women authors than expected based on society membership. This echoes declines in women's scholarly productivity in the first year of the COVID-19 pandemic observed across many disciplines. We offer four recommendations: (1) practice double-anonymous peer review; (2) recruit editors from diverse backgrounds who invite reviewers with diverse backgrounds; (3) democratize manuscript review by selecting reviewers from a disaggregated reviewer database; and (4) gather and analyze demographic data for both submissions and publications.

Biological variation fuels evolutionary change. Across longer timescales, however, polymorphisms at both the genomic and phenotypic levels often persist longer than would be expected under standard population genetic models such as positive selection or genetic drift. Explaining the maintenance of this variation within populations across long time spans via balancing selection has been a major triumph of theoretical population genetics and ecology. Although persistent polymorphisms can often be traced in fossil lineages over long periods through the rock record, paleobiology has had little to say about either the long-term maintenance of phenotypic variation or its macroevolutionary consequences. I explore the dynamics that occur when persistent polymorphisms maintained over long lineage durations are filtered into descendant lineages during periods of demographic upheaval that occur at speciation. I evaluate these patterns in two lineages: Ectocion, a genus of Eocene mammals, and botryocrinids, a Mississippian cladid crinoid family. Following origination, descendants are less variable than their ancestors. The patterns by which ancestral variation is sorted cannot be distinguished from drift. Maintained and accumulated polymorphisms in highly variable ancestral lineages such as Barycrinus rhombiferus Owen and Shumard, 1852 may fuel radiations as character states are sorted into multiple descendant lineages. Interrogating the conditions under which trans-specific polymorphism is either maintained or lost during periods of demographic and ecological upheaval can explain how population-level processes contribute to the emergent macroevolutionary dynamics that shape the history of life as preserved in the fossil record.

The process of evolution and the structures it produces are best understood in the light of hierarchy theory. The biota traditionally is described by either the genealogical Linnaean hierarchy or economic hierarchies of communities or ecosystems. Here we describe the Bretskyan hierarchy—a hybrid eco-genealogical hierarchy that consists of nested sets of different-sized, usually polyphyletic communities of interacting individuals separated from other such communities in space and time at multiple scales. The Bretskyan hierarchy consists of elements that have both genealogical and economic properties and functions—situated between, and connecting the elements of, the economic hierarchies (Vernadskyan) and the genealogical (Linnaean) hierarchy. The described hierarchy at lower tiers is populated by holobionts, individuals composed of multiple polyphyletic lineages integrated by functional interactions or biotically fabricated structures, such as membranes. At larger spatial tiers and longer time scales, the members of the Bretskyan hierarchy are of a more diffuse nature, partially due to the small size and relatively short duration of us as observers of larger and longer-lasting structures, here described as geobiomes. Their individuality is externally forced and directly tied to the spatial and temporal physical structures of our planet. These are sub-bioprovinces and bioprovinces—large and effectively isolated spatiotemporal structures of biota integrated internally by coevolution and individuated externally by a hierarchy of barriers. Gaia is here understood as the largest eco-genealogical individual compartmentalized by the outer space of the Earth and integrated at long time scales by biotic interactions and plate tectonic mixing of biota. The existence of a hierarchy of barriers and multilevel allopatry suggests that geographic isolation takes part not only in individuating species lineages, but also in producing coherent complexes of separate lineages forming bioprovinces at multiple space and time scales. The sizes, configurations, and durations of Bretskyan units are directly tied to geodynamics, demonstrating the central role of the physical planet in the processes of individuation and merging of geobiomes and the control of coevolution, and all its ramifications, at multiple space and time scales. The Bretskyan hierarchy also allows the integration of previously unconnected themes—“egalitarian” major transitions in individuality (e.g., eukaryogenesis) and some of the megatrajectories in the history of life—into a single theoretical framework of spatial and temporal scaling of eco-genealogy. The pervasive scaling of geodynamical processes and the direct connection of geodynamics to the dynamics of Bretskyan units allows us to formulate conjectures on the scales and limits of spatial and temporal contingency and competitiveness of biotas in evolution.

We know the fossil record is incomplete, but just how much biodiversity does it miss? We produce the first geographically controlled estimate by comparing the geographic ranges of 34,266 modern tetrapods with a map of the world's sedimentary basins. By modeling which tetrapods live within sedimentary basins, we produce a first-order estimate of what might be found in the fossil record of the future. In this record, nearly 30% of tetrapod species have almost no chance of fossilizing, and more stringent criteria for fossilization exclude far more diversity. This geographically structured fossil record preserves disparate patterns of taxonomic and phylogenetic diversity in different tetrapod groups and underpreserves projected extinctions. For the globally threatened amphibians, the magnitude of the extinction of all endangered species would be underestimated by 66–98% in our future record. These results raise profound questions about the structure of the fossil record. Is it capable of recording major origination and extinction events on land? Have swaths of terrestrial diversity gone unrecorded based on geography alone? There are chapters of Earth history that paleontologists can never hope to know, but what is missing, and why?

The first size reduction (FSR) in the Reticulofenestra-Gephyrocapsa-Emiliania (RGE) lineage (order Isochrysidales), which occurred in the early Oligocene (~32 Ma), is of great significance for understanding the Lilliput effect that has affected coccolithophore communities from the late Eocene to this day. We conducted a morphologic analysis on the coccoliths of Reticulofenestra species that lived during the late middle Eocene to early Oligocene (~40–31 Ma), using marine sediments from the South Atlantic Ocean. Our data show increasing size and decreasing abundance of the large species during the late Eocene, leading to their disappearance at the FSR, and a concurrent decrease in the size variability of the small- to medium-sized coccoliths whose central opening diameter had become very reduced. Although the cosmopolitan late Paleogene through Neogene size decrease in coccolithophores has been linked to the concomitant long-term decline in global pCO2, we suggest here that the FSR was the result of environmental destabilization caused by the expansion of eutrophic environments following the late Eocene establishment of overturning circulation associated with ice buildup on Antarctica. This study also leads us to propose a hypothetical model that links coccolith morphology of species of the RGE lineage and trophic resources in the upper ocean: the small- to medium-sized, r-selected coccolithophores with smaller coccolith central openings live in nutrient-rich waters where they rely mostly on photosynthesis and little on mixotrophy, whereas the larger, K-selected species with larger coccolith central openings live in oligotrophic waters where they are more dependent on mixotrophy.

The Fezouata Shale Formation has dramatically impacted our understanding of Early Ordovician marine ecosystems before the great Ordovician biodiversification event (GOBE), thanks to the abundance and quality of exceptionally preserved animals within it. Systematic work has noted that the shelly fossil subassemblages of the Fezouata Shale biota are typical of open-marine deposits from the Lower Ordovician, but no studies have tested the quantitative validity of this statement. We extracted 491 occurrences of recalcitrant fossil genera from the Paleobiology Database to reconstruct 31 subassemblages to explore the paleoecology of the Fezouata Shale and other contemporary, high-latitude (66°S–90°S) deposits from the Lower Ordovician (485.4–470 Ma) and test the interpretation that the Fezouata Shale biota is typical for an Ordovician open-marine environment. Sørensen's dissimilarity metrics and Wilcoxon tests indicate that the subassemblages of the Tremadocian-aged lower Fezouata Shale are approximately 20% more heterogenous than the Floian-aged upper Fezouata Shale. Dissimilarity metrics and visualization suggest that while the lower Fezouata and upper Fezouata share faunal components, the two sections have distinct faunas. We find that the faunal composition of the lower Fezouata Shale is comparable with other Tremadocian-aged subassemblages from high latitudes, suggesting that it is typical for an Early Ordovician open-marine environment. We also find differences in faunal composition between Tremadocian- and Floian-aged deposits. Our results corroborate previous field-based and qualitative systematic studies that concluded that the shelly assemblages of the Fezouata Shale are comparable with those of other Lower Ordovician deposits from high latitudes. This establishes the first quantitative baseline for examining the composition and variability within the assemblages of the Fezouata Shale and will be key to future studies attempting to discern the degree to which it can inform our understanding of marine ecosystems just before the start of the GOBE.

Corallite sizes reflect a continuum in the efficacy of photosymbiosis in colonial reef corals, with smaller corallite sizes generally associated with higher autotrophy. Using a large compilation of reef-coral traits and corallite diameters as a proxy, we test here the hypothesis that photosymbiotic efficacy has increased over the evolutionary history of scleractinian corals. To gain a more comprehensive understanding of the evolutionary versus ecological patterns of corallite sizes of reef corals, we used three analytical methods: (1) occurrences-weighted within-bin analyses as a proxy for abundance or ecological dominance to depict ecological patterns; (2) unweighted range-through analyses; and (3) unweighted sampled-in-bin analyses to represent diversity in terms of taxonomic richness, enabling us to trace evolutionary patterns. By-genus, range-through analysis indicates a slightly positive trend of corallite sizes toward the Recent. However, the occurrences-weighted assessment shows a pronounced negative trend of corallite sizes in colonial corals since the Mesozoic. Random walk and directional evolution are both statistically supported to explain this long-term decrease. A driven trend is evolutionarily plausible, giving reef corals a selective advantage in the oligotrophic environments they largely occupy today.

Radiodonta is a clade of stem euarthropods of central importance to our understanding of the evolution of this phylum. Radiodonts include some of the largest early Paleozoic animals; however, little is known about their ontogeny. We present an analysis of molting patterns and ontogeny in the radiodont Stanleycaris based on 265 exceptionally preserved specimens from the mid-Cambrian (Wuliuan) Burgess Shale. Ranging in size from 10 to 83 mm, they constitute the most extensive radiodont ontogenetic series known. Using a novel morphospace approach, we show that putative carcasses and exuviae can be quantitatively distinguished by the particular suites of structures preserved and their modes of preservation. We propose that Stanleycaris, and probably other radiodonts, molted via a suture near the anterior of the trunk. Similar anterior molting strategies, with a suture located at the head–trunk boundary, are shared with some Cambrian euarthropods and are potentially ancestral. Allometric analyses suggest that as Stanleycaris body size increased, the head sclerite and neck became relatively broader, while the trunk and flaps became slightly longer. The eyes developed precociously, indicating an important role of visual processing in juveniles. Finally, we find evidence for an initial anamorphic developmental phase, where segment number increased at least from 11 or 12 up to 17, followed by an epimorphic phase, in which growth continued without segment addition. This is consistent with the hypothesis that finite postembryonic segment addition (hemianamorphosis) is ancestral for arthropods and refines the timing of the origin of this important developmental mode.

A traditional typological approach to taxonomy often does not adequately account for intraspecific variation and can result in taxonomic oversplitting. For many groups, including ammonoids of the Placenticeras genus, intraspecific variation documented in recent studies (e.g., ontogenetic changes, sexual dimorphism, polymorphism) challenges the historic proliferation of species names. Here, we used a population approach to taxonomy and quantitatively evaluated morphometric variation in a sample of Late Cretaceous (Santonian–Campanian) Placenticeras from Alabama and adjacent counties.

We used linear mixed models (LMMs) to characterize how morphological variables scale with conch size across the sample, exploiting mixed longitudinal data to evaluate individual variation in growth and inform interpretations of multivariate analyses. Extended LMMs incorporating geological formation evaluated morphological changes through time. Principal component analysis and clustering analysis were then used to evaluate the number of distinct clusters that emerged in multivariate morphospace independent of previous taxon name assignments.

Discontinuous scaling relationships and distinct clusters in multivariate space suggest sexual dimorphism characterized by differences in adult size and, secondarily, shape. Previous Stantonoceras and Placenticeras assignments broadly overlap in our morphospace, failing to justify this historic distinction (as sexual dimorphs or as genera or subgenera). Placenticeras conch morphology and ornament placement changed through time, suggesting a potential utility for coarse (stage-level) biostratigraphy. However, temporal changes were not associated with distinct clusters in morphospace, and our data fail to support the plethora of reported species names. As few as one or two (successive) species may be present in our sample (representing 130 years of collection effort). In addition to highlighting the need for a significant taxonomic revision of the Placenticeras genus, this study demonstrates the utility of LMMs for distinguishing between different sources of morphological variation, improving interpretations of morphospace under a population approach to taxonomy, and maximizing the amount of ontogenetic information that can be obtained nondestructively.

The Paleozoic represents a key time interval in the origins and early diversification of chondrichthyans (cartilaginous fishes), but their diversity and macroevolution are largely obscured by heterogenous spatial and temporal sampling. The predominantly cartilaginous skeletons of chondrichthyans pose an additional limitation on their preservation potential and hence on the quality of their fossil record. Here, we use a newly compiled genus-level dataset and the application of sampling standardization methods to analyze global total-chondrichthyan diversity dynamics through time from their first appearance in the Ordovician through to the end of the Permian. Subsampled estimates of chondrichthyan genus richness were initially low in the Ordovician and Silurian but increased substantially in the Early Devonian. Richness reached its maximum in the middle Carboniferous before dropping across the Carboniferous/Permian boundary and gradually decreasing throughout the Permian. Sampling is higher in both the Devonian and Carboniferous compared with the Silurian and most of the Permian stages. Shark-like scales from the Ordovician are too limited to allow for some of the subsampling techniques. Our results detect two Paleozoic radiations in chondrichthyan diversity: the first in the earliest Devonian, led by acanthodians (stem-group chondrichthyans), which then decline rapidly by the Late Devonian, and the second in the earliest Carboniferous, led by holocephalans, which increase greatly in richness across the Devonian/Carboniferous boundary. Dispersal of chondrichthyans, specifically holocephalans, into deeper-water environments may reflect a niche expansion following the faunal displacement in the aftermath of the Hangenberg extinction event at the end of the Devonian.

The Mississippi River watershed drains 40% of the continental United States, and the tremendous primary productivity in the adjacent north-central Gulf of Mexico has created one of the most extensive dead zones on Earth. In contrast, smaller watersheds deliver fewer nutrients to the northeastern gulf, and consequently, productivity is limited and hypoxia is uncommon. How has variation in primary productivity, oxygen availability, and sea-surface temperature affected coastal food webs? Here, we investigate environmental controls on the size of molluscan predators and prey in the northern Gulf of Mexico using Holocene death assemblages. Linear mixed models indicate that bivalve size and the frequency of drilling predation are affected by dissolved oxygen concentrations; drilling frequency declines with declining oxygen, whereas bivalve size increases. In contrast, sea-surface temperature is positively associated with the size of molluscan predators and prey. Net primary productivity contributes relatively little to predator or prey size, and predator-to-prey size ratios do not vary consistently with environmental conditions across the northern gulf. Larger bivalves in areas of oxygen limitation may be due to decreased predation pressure and, consequently, greater prey longevity. The larger size of bivalves and predatory gastropods in warmer waters may reflect enhanced growth under these conditions, provided dissolved oxygen concentrations exceed a minimum threshold. Holocene death assemblages can be used to test long-standing hypotheses regarding environmental controls on predator−prey body-size distributions through geologic time and provide baselines for assessing the ongoing effects of anthropogenic eutrophication and warming on coastal food webs.

The Late Triassic conodont species Mockina ex gr. carinata and Mockina ex gr. englandi were exceptionally prevalent among the marine fauna of the Panthalassan realm from the middle Norian through to the Rhaetian. Leading into the complete extinction of conodonts near the Triassic/Jurassic boundary, a significant turnover event occurred in conodont fauna across the Norian/Rhaetian boundary (NRB), with the pectiniform elements of common Rhaetian genera from Tethys exhibiting minimal or absent platforms. This intergeneric trend of platform reduction is not as evident in Panthalassa, where these genera are very rare, but morphometric analyses of M. ex gr. carinata and M. ex gr. englandi specimens from across the Canadian Cordillera demonstrate that comparable shifts in morphology occurred intraspecifically in Panthalassa across the NRB, confirming the global extent of these trends. Pectiniform elements of M. ex gr. carinata display a sequential reduction of platform width from the middle Norian to late Norian to Rhaetian, whereas pectiniform elements of M. ex gr. englandi display a reduction of platform width only from the late Norian to Rhaetian. Specimens of both species that have a mid-platform length to breadth ratio greater than 3:1 are restricted to the Rhaetian. Specimens from the Kennecott Point section on Haida Gwaii, British Columbia, demonstrate that this morphological shift occurred somewhat later than other biostratigraphic proxies for the NRB. The global trend of platform width reduction in many conodont pectiniform elements may reflect a change in primary diet away from hard food sources, perhaps suggesting some degree of carbonate biomineralization suppression beginning around the NRB. This interpretation would support CO2 outgassing as the causal mechanism of the environmental disturbance at the NRB and identify the NRB as a significant turning point for Late Triassic ecosystems, marking the beginning of a protracted, multiphase end-Triassic mass extinction.

Crocodylomorpha is a large and diverse clade with a long evolutionary history now restricted to modern crocodilians. Tethysuchia is a less-inclusive clade of semi-amphibious taxa that crossed two biological crises: the second Oceanic Anoxic Event (OAE 2) and the Cretaceous/Paleogene (K/Pg) crisis. Numerous studies have sought to find the driving factors explaining crocodylomorph evolution, producing contradictory conclusions. Studies of included groups may be useful. Here, we study factors driving tethysuchian evolution using phylogenetically informed statistical analyses. First, we tested the phylogenetic structure of tethysuchian extinction at the OAE 2 and K/Pg crises. We then used phylogenetic comparative methods to test the influence of intrinsic (body size, snout proportion) and extrinsic (temperature, paleolatitude) factors on the evolution of tethysuchian diversity at the OAE 2 and the K/Pg crises. Finally, we tested whether temperature influenced the evolution of body size. We conclude that (1) extinction was not random in regard to phylogeny for Tethysuchia at the OAE 2 and K/Pg crises; (2) while an important tethysuchian turnover follows OAE 2, the K/Pg crisis was followed by an explosion in diversity of tethysuchians, probably linked to the colonization of emptied ecological niches; (3) tethysuchians lived in warmer environments after the OAE 2 crisis, possibly because of both global warming and latitudinal distribution shifts; (4) there is a significant change of snout proportion after the OAE 2 and the K/Pg crises, likely caused by niche partitioning; and (5) there is a positive correlation between body size and temperature, possibly because of a longer growth season.

The unique body plan of frogs (Lissamphibia: Anura) has been largely conserved from at least 200 Myr, and its evolution from a more generalized tetrapod condition is still poorly understood, in part due to the scarce early fossil record of Salientia, the anuran total-group. The origin of the anuran Bauplan has been classically explained as an adaptation to jumping, but recent studies incorporating new data in a phylogenetic context have challenged the popular jumping hypothesis. Here we revisit and test this hypothesis from a paleobiological perspective by integrating limb data from a wide range of extant and fossil frogs. We first explored the evolution of limb proportions from the Jurassic to the Paleogene to understand when the present limb diversity originated and whether, and to what extent, limb proportions have been conserved over the last 200 Myr. We then inferred the locomotor capabilities of extinct species by phylogenetic flexible discriminant analysis, and from these inferences, we studied the locomotor diversity of frogs over geological time and reconstructed the ancestral state for frog-like salientians. The evolution of limb proportions is characterized by an early diversification that was underway in the Jurassic, followed by a repeated convergence over a limited area of the morphospace that was already explored by the Early Cretaceous. In agreement with this early limb diversity, the Jurassic stem species were also locomotory diverse, and their inferred locomotor modes do not support the jumping hypothesis. We propose that the patterns found herein of repeated convergent evolution of both limb proportions and locomotor capabilities over geological time hamper any attempt to confidently infer the ancestral locomotion mode and, it therefore might be time to start focusing on other hypotheses on the origin of the anuran Bauplan that are not related to locomotion.

Theropods are obligate bipedal dinosaurs that appeared 230 Ma and are still extant as birds. Their history is characterized by extreme variations in body mass, with gigantism evolving convergently between many lineages. However, no quantification of hindlimb functional morphology has shown whether these body mass increases led to similar specializations between distinct lineages. Here we studied femoral shape variation across 41 species of theropods (n = 68 specimens) using a high-density 3D geometric morphometric approach. We demonstrated that the heaviest theropods evolved wider epiphyses and a more distally located fourth trochanter, as previously demonstrated in early archosaurs, along with an upturned femoral head and a mediodistal crest that extended proximally along the shaft. Phylogenetically informed analyses highlighted that these traits evolved convergently within six major theropod lineages, regardless of their maximum body mass. Conversely, the most gracile femora were distinct from the rest of the dataset, which we interpret as a femoral specialization to “miniaturization” evolving close to Avialae (bird lineage). Our results support a gradual evolution of known “avian” features, such as the fusion between lesser and greater trochanters and a reduction of the epiphyseal offset, independent from body mass variations, which may relate to a more “avian” type of locomotion (more knee than hip driven). The distinction between body mass variations and a more “avian” locomotion is represented by a decoupling in the mediodistal crest morphology, whose biomechanical nature should be studied to better understand the importance of its functional role in gigantism, miniaturization, and higher parasagittal abilities.

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.

Electron backscatter diffraction (EBSD) has been widely used in recent studies of eggshells for its convenience in collecting in situ crystallographic information. China has a wide variety of dinosaur eggshells, although nearly none have been studied with this technique. Elongatoolithid eggs include many oogenera, although the microstructural differences of some were not highly appreciated, leading to several parataxonomic problems. In this paper, we surveyed seven elongatoolithid oogenera in China using EBSD in order to acquire more information about their microstructural variation. It is shown in this paper that in some elongatoolithid eggshells, scaly calcite grains that form the squamatic ultrastructure are not the only form of calcite in the continuous layer. Large columnar grains separated by high-angled grain boundaries and slender subgrains separated by radially arranged low-angled grain boundaries could exist in certain areas of the eggshells such as Macroolithus and Macroelongatoolithus. This paper discusses the criteria for identifying squamatic ultrastructure and proposes type I (rich in rugged high-angled grain boundaries) and type II (rich in both rugged high- and low-angled grain boundaries) squamatic ultrastructures. A pathological layer is found in Undulatoolithus pengi. An external zone is identified in the eggshell of Heishanoolithus changii, which does not support its position within the oofamily Elongatoolithidae. We argue that Paraelongatoolithus no longer belongs to Elongatoolithidae based on a combination of reticulated ornamentation, columnar continuous layer, and acicular mammillae. The high structural variation in elongatoolithid eggshells also implies that it may be inappropriate to relate all previous elongatoolithid eggshells to oviraptorosaurs and assume they are non-monophyletic.