Pterosaurs, a Mesozoic group of flying archosaurs, have become a focal point for debates pertaining to the impact of sampling biases on our reading of the fossil record, as well as the utility of sampling proxies in palaeodiversity reconstructions. The completeness of the pterosaur fossil specimens themselves potentially provides additional information that is not captured in existing sampling proxies, and might shed new light on the group's evolutionary history. Here we assess the quality of the pterosaur fossil record via a character completeness metric based on the number of phylogenetic characters that can be scored for all known skeletons of 172 valid species, with averaged completeness values calculated for each geological stage. The fossil record of pterosaurs is observed to be strongly influenced by the occurrence and distribution of Lagerstätten. Peaks in completeness correlate with Lagerstätten deposits, and a recovered correlation between completeness and observed diversity is rendered non-significant when Lagerstätten species are excluded. Intervals previously regarded as potential extinction events are shown to lack Lagerstätten and exhibit low completeness values: as such, the apparent low diversity in these intervals might be at least partly the result of poor fossil record quality. A positive correlation between temporal patterns in completeness of Cretaceous pterosaurs and birds further demonstrates the prominent role that Lagerstätten deposits have on the preservation of smaller bodied organisms, contrasting with a lack of correlation with the completeness of large-bodied sauropodomorphs. However, we unexpectedly find a strong correlation between sauropodomorph and pterosaur completeness within the Triassic-Jurassic, but not the Cretaceous, potentially relating to a shared shift in environmental preference and thus preservation style through time. This study highlights the importance of understanding the relationship between various taphonomic controls when correcting for sampling bias, and provides additional evidence for the prominent role of sampling on observed patterns in pterosaur macroevolution. KeywordsFossil record completeness; Lagerstätten; Mesozoic; Pterodactyloidea; Pterosauria; Sampling bias PTEROSAURS were a group of Mesozoic flying archosaurs that went extinct at the Cretaceous/Paleogene (K/Pg) mass extinction 66 Ma, approximately 150 myr after their first appearance in the fossil record (Wellnhofer 1991;Unwin 2003Unwin , 2005Barrett et al. 2008; Dalla Vecchia 2013;Witton 2013). They show high taxonomic (Unwin 2005;Butler et al. 2009Butler et al. , 2013Witton 2013) and morphological diversity (Prentice et al. 2011; Foth et al. Europe PMC Funders GroupAuthor Manuscript Palaeontology. Author manuscript; available in PMC 2017 January 14. Published in final edited form as:Palaeontology. 2016 January 14; 59(2): 225-247. doi:10.1111/pala.12225. Europe PMC Funders Author ManuscriptsEurope PMC Funders Author Manuscripts 2012;Butler et al. 2012), with a global distribution b...

The advent of palaeontological occurrence databases has allowed for detailed reconstruction and analyses of species richness through deep time. While a substantial literature has evolved ensuring that taxa are fairly counted within and between different time periods, how time itself is divided has received less attention. Stage-level or equalinterval age bins have frequently been used for regional and global studies in vertebrate palaeontology. However, when assessing diversity at a regional scale, these resolutions can prove inappropriate with the available data. Herein, we propose a new method of binning geological time for regional studies that intrinsically incorporates the chronostratigraphic heterogeneity of different rock formations to generate unique stratigraphic bins. We use this method to investigate the diversity dynamics of dinosaurs from the Late Cretaceous of the Western Interior of North America prior to the Cretaceous-Palaeogene mass extinction.

The latitudinal biodiversity gradient (LBG), in which species richness decreases from tropical to polar regions, is a pervasive pattern of the modern biosphere. Although the distribution of fossil occurrences suggests this pattern has varied through deep time, the recognition of palaeobiogeographic patterns is hampered by geological and anthropogenic biases. In particular, spatial sampling heterogeneity has the capacity to impact upon the reconstruction of deep time LBGs. Here we use a simulation framework to test the detectability of three different types of LBG (flat, unimodal and bimodal) over the last 300 Myr. We show that heterogeneity in spatial sampling significantly impacts upon the detectability of genuine LBGs, with known biodiversity patterns regularly obscured after applying the spatial sampling window of fossil collections. Sampling-standardization aids the reconstruction of relative biodiversity gradients, but cannot account for artefactual absences introduced by geological and anthropogenic biases. Therefore, we argue that some previous studies might have failed to recover the ‘true’ LBG type owing to incomplete and heterogeneous sampling, particularly between 200 and 20 Ma. Furthermore, these issues also have the potential to bias global estimates of past biodiversity, as well as inhibit the recognition of extinction and radiation events.

Pterosaurs, a Mesozoic group of flying archosaurs, have become a focal point for debates pertaining to the impact of sampling biases on our reading of the fossil record, as well as the utility of sampling proxies in palaeodiversity reconstructions. The completeness of the pterosaur fossil specimens themselves potentially provides additional information that is not captured in existing sampling proxies, and might shed new light on the group’s evolutionary history. Here we assess the quality of the pterosaur fossil record via a character completeness metric based on the number of phylogenetic characters that can be scored for all known skeletons of 172 valid species, with averaged completeness values calculated for each geological stage. The fossil record of pterosaurs is observed to be strongly influenced by the occurrence and distribution of Lagerstätten. Peaks in completeness correlate with Lagerstätten deposits, and a recovered correlation between completeness and observed diversity is rendered non-significant when Lagerstätten species are excluded. Intervals previously regarded as potential extinction events are shown to lack Lagerstätten and exhibit low completeness values: as such, the apparent low diversity in these intervals might be at least partly the result of poor fossil record quality. A positive correlation between temporal patterns in completeness of Cretaceous pterosaurs and birds further demonstrates the prominent role that Lagerstätten deposits have on the preservation of smaller bodied organisms, contrasting with a lack of correlation with the completeness of large-bodied sauropodomorphs. However, we unexpectedly find a strong correlation between sauropodomorph and pterosaur completeness within the Triassic–Jurassic, but not the Cretaceous, potentially relating to a shared shift in environmental preference and thus preservation style through time. This study highlights the importance of understanding the relationship between various taphonomic controls when correcting for sampling bias, and provides additional evidence for the prominent role of sampling on observed patterns in pterosaur macroevolution.

Despite extensive outcrop and previous sedimentologic study, the role of tidal processes along sandy, wave- and river-dominated shorelines of the North American Cretaceous Western Interior Seaway remains uncertain, particularly for the extensive mid-Campanian (ca. 75–77.5 Ma) tidal deposits of Utah and Colorado, USA. Herein, paleotidal modeling, paleogeographic reconstructions, and interpretations of depositional process regimes are combined to evaluate the regional-scale (hundreds to thousands of kilometers) basin physiographic controls on tidal range and currents along these regressive shorelines in the “Utah Bight”, southwestern Western Interior Seaway. Paleotidal modeling using a global and astronomically forced tidal model, combined with paleobathymetric sensitivity tests, indicates the location of stratigraphic units preserving pronounced tidal influence only when the seaway had a deep center (∼400 m) and southern entrance (>100 m). Maximum tidal velocity vectors under these conditions suggest a dominant southeasterly ebb tide within the Utah Bight, consistent with the location and orientation of paleocurrent measurements in regressive, tide-influenced deltaic units. The modeled deep paleobathymetry increased tidal inflow into the basin and enhanced local-scale (tens to hundreds of kilometers) resonance effects in the Utah Bight, where an amphidromic cell was located. However, the preservation of bidirectional, mudstone-draped cross-stratification in fine- to medium-grained sandstones requires tides in combination with fluvial currents and/or local tidal amplification below the maximum resolution of model meshes (∼10 km). These findings suggest that while regional-scale controls govern tidal potential within basins, localized physiography exerts an important control on the preservation of tidal signatures in the geologic record.