Inferring the Total-Evidence Timescale of Marattialean Fern Evolution in the Face of Model Sensitivity.

Phylogenetic divergence-time estimation has been revolutionized by two recent developments: 1) total-evidence dating (or "tip-dating") approaches that allow for the incorporation of fossils as tips in the analysis, with their phylogenetic and temporal relationships to the extant taxa inferred from the data and 2) the fossilized birth-death (FBD) class of tree models that capture the processes that produce the tree (speciation, extinction, and fossilization) and thus provide a coherent and biologically interpretable tree prior. To explore the behavior of these methods, we apply them to marattialean ferns, a group that was dominant in Carboniferous landscapes prior to declining to its modest extant diversity of slightly over 100 species. We show that tree models have a dramatic influence on estimates of both divergence times and topological relationships. This influence is driven by the strong, counter-intuitive informativeness of the uniform tree prior, and the inherent nonidentifiability of divergence-time models. In contrast to the strong influence of the tree models, we find minor effects of differing the morphological transition model or the morphological clock model. We compare the performance of a large pool of candidate models using a combination of posterior-predictive simulation and Bayes factors. Notably, an FBD model with epoch-specific speciation and extinction rates was strongly favored by Bayes factors. Our best-fitting model infers stem and crown divergences for the Marattiales in the mid-Devonian and Late Cretaceous, respectively, with elevated speciation rates in the Mississippian and elevated extinction rates in the Cisuralian leading to a peak diversity of ${\sim}$2800 species at the end of the Carboniferous, representing the heyday of the Psaroniaceae. This peak is followed by the rapid decline and ultimate extinction of the Psaroniaceae, with their descendants, the Marattiaceae, persisting at approximately stable levels of diversity until the present. This general diversification pattern appears to be insensitive to potential biases in the fossil record; despite the preponderance of available fossils being from Pennsylvanian coal balls, incorporating fossilization-rate variation does not improve model fit. In addition, by incorporating temporal data directly within the model and allowing for the inference of the phylogenetic position of the fossils, our study makes the surprising inference that the clade of extant Marattiales is relatively young, younger than any of the fossils historically thought to be congeneric with extant species. This result is a dramatic demonstration of the dangers of node-based approaches to divergence-time estimation, where the assignment of fossils to particular clades is made a priori (earlier node-based studies that constrained the minimum ages of extant genera based on these fossils resulted in much older age estimates than in our study) and of the utility of explicit models of morphological evolution and lineage diversification. [Bayesian model comparison; Carboniferous; divergence-time estimation; fossil record; fossilized birth-death; lineage diversification; Marattiales; models of morphological evolution; Psaronius; RevBayes.].

Corrigendum: The Extended Specimen Network: A Strategy to Enhance US Biodiversity Collections, Promote Research and Education.

[This corrects the article DOI: 10.1093/biosci/biz140.].

The first juvenile dromaeosaurid (Dinosauria: Theropoda) from Arctic Alaska.

Compared to the osteological record of herbivorous dinosaurs from the Late Cretaceous Prince Creek Formation of northern Alaska, there are relatively fewer remains of theropods. The theropod record from this unit is mostly comprised of isolated teeth, and the only non-dental remains known can be attributed to the troodontid cf. Troodon and the tyrannosaurid Nanuqsaurus. Thus far, the presence of members of Dromaeosauridae has been limited to isolated teeth. Here we describe a symphyseal portion of a small dentary with two ziphodont teeth. Based on tooth shape, denticle morphology, and the position of the Meckelian groove, we attribute this partial dentary to a saurornitholestine dromaeosaurid. The fibrous bone surface, small size, and higher number of mesial denticles compared to distal ones point to a juvenile growth stage for this individual. Multivariate comparison of theropod teeth morphospace by means of principal component analysis reveals an overlap between this dentary and Saurornitholestinae dromaeosaurid morphospace, a result supported by phylogenetic analyses. This is the first confirmed non-dental fossil specimen from a member of Dromaeosauridae in the Arctic, expanding on the role of Beringia as a dispersal route for this clade between Asia and North America. Furthermore, the juvenile nature of this individual adds to a growing body of data that suggests Cretaceous Arctic dinosaurs of Alaska did not undergo long-distance migration, but rather they were year-round residents of these paleopolar latitudes.

A workflow and protocol describing the field to digitization process for new project-based fossil leaf collections.

PREMISE OF THE STUDY: This article provides a workflow and protocol for paleobotanical researchers that integrates project-based fossil leaf specimen and data collection with curation and digitization. The methods aim to facilitate efficient digitization of new collections by researchers during the course of their study and promote public databasing of new specimen and project data. METHODS AND RESULTS: The workflow was developed and refined to facilitate a project reconstructing an extensive fossil forest from leaf impressions/compressions. The workflow consists of field, museum, and data mobilization components. Customizing a workspace and streamlining all steps of specimen data collection, curation, and digitization into an integrated processing pipeline resulted in faster accumulation of specimen data and images. CONCLUSIONS: These protocols provide paleobotanists with logistics-focused methods for integrating research with digitization, and are particularly applicable at institutions with limited collection support staff or when specimen images are needed for project purposes.