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

   https://doi.org/10.1093/sysbio/syab020  

   May, Michael R; Contreras, Dori L; Sundue, Michael A; Nagalingum, Nathalie S; Looy, Cindy V; Rothfels, Carl J (March 2021, Systematic Biology)

   Folk, Ryan (Ed.)

   Abstract 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.] 

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2. Fossils Do Not Substantially Improve, and May Even Harm, Estimates of Diversification Rate Heterogeneity

   https://doi.org/10.1093/sysbio/syac049  

   Beaulieu, Jeremy M.; O’Meara, Brian C.; Wright, ed., April (July 2022, Systematic Biology)

   Abstract The fossilized birth–death (FBD) model is a naturally appealing way of directly incorporating fossil information when estimating diversification rates. However, an important yet often overlooked property of the original FBD derivation is that it distinguishes between two types of sampled lineages. Here, we first discuss and demonstrate the impact of severely undersampling, and even not including fossils that represent samples of lineages that also had sampled descendants. We then explore the benefits of including fossils, generally, by implementing and then testing two types of FBD models, including one that converts a fossil set into stratigraphic ranges, in more complex likelihood-based models that assume multiple rate classes across the tree. Under various simulation scenarios, including a scenario that exists far outside the set of models we evaluated, including fossils rarely outperform analyses that exclude them altogether. At best, the inclusion of fossils improves precision but does not influence bias. Similarly, we found that converting the fossil set to stratigraphic ranges, which is one way to remedy the effects of undercounting the number of k-type fossils, results in turnover rates and extinction fraction estimates that are generally underestimated. Although fossils remain essential for understanding diversification through time, in the specific case of understanding diversification given an existing, largely modern tree, they are not especially beneficial. [Fossilized birth–death; fossils; MiSSE; state speciation extinction; stratigraphic ranges; turnover rate.] 

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3. Putting the F into FBD analysis: tree constraints or morphological data?

   https://doi.org/10.1111/pala.12679  

   Barido‐Sottani, Joëlle; Pohle, Alexander; De_Baets, Kenneth; Murdock, Duncan; Warnock, Rachel_C_M (November 2023, Palaeontology)

   Abstract The fossilized birth–death (FBD) process provides an ideal model for inferring phylogenies from both extant and fossil taxa. Using this approach, fossils are directly integrated into the tree, leading to a statistically coherent prior on divergence times. Since fossils are typically not associated with molecular sequences, additional information is required to place fossils in the tree. We use simulations to evaluate two different approaches to handling fossil placement in FBD analyses: using topological constraints, where the user specifies monophyletic clades based on established taxonomy, or using total‐evidence analyses, which use a morphological data matrix in addition to the molecular alignment. We also explore how rate variation in fossil recovery or diversification rates impacts these approaches. We find that the extant topology is well recovered under all methods of fossil placement. Divergence times are similarly well recovered across all methods, with the exception of constraints which contain errors. We see similar patterns in datasets which include rate variation, however, relative errors in extant divergence times increase when more variation is included in the dataset, for all approaches using topological constraints, and particularly for constraints with errors. Finally, we show that trees recovered under the FBD model are more accurate than those estimated using non‐time calibrated inference. Overall, we show that both fossil placement approaches are reliable even when including uncertainty. Our results underscore the importance of core taxonomic research, including morphological data collection and species descriptions, irrespective of the approach to handling phylogenetic uncertainty using the FBD process. 

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4. Fossils improve extinction-rate estimates under state-dependent diversification models

   https://doi.org/10.1098/rstb.2023.0313  

   do_Rosario_Petrucci, Bruno; May, Michael R; Heath, Tracy A (February 2025, Philosophical Transactions of the Royal Society B: Biological Sciences)

   The effect of traits on diversification rates is a major topic of study in the fields of evolutionary biology and palaeontology. Many researchers investigating these macroevolutionary questions currently make use of the extensive suite of state-dependent speciation and extinction (SSE) models. These models were developed for, and are almost exclusively used with, phylogenetic trees of extant species. However, analyses considering only extant taxa are limited in their power to estimate extinction rates. Furthermore, SSE models can erroneously detect associations between neutral traits and diversification rates when the true associated trait is not observed. In this study, we examined the impact of including fossil data on the accuracy of parameter estimates under the binary-state speciation and extinction (BiSSE) model. This was achieved by combining SSE models with the fossilized birth–death process. We show that the inclusion of fossils improves the accuracy of extinction-rate estimates for analyses applying the BiSSE model in a Bayesian inference framework, with no negative impact on speciation-rate and state transition-rate estimates when compared with estimates from trees of only extant taxa. However, even with the addition of fossil data, analyses under the BiSSE model continued to incorrectly identify correlations between diversification rates and neutral traits. This article is part of the theme issue ‘“A mathematical theory of evolution”: phylogenetic models dating back 100 years’. 

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5. Assessing the impact of incomplete species sampling on estimates of speciation and extinction rates

   https://doi.org/10.1017/pab.2020.12  

   Warnock, Rachel C.; Heath, Tracy A.; Stadler, Tanja (May 2020, Paleobiology)

   Abstract Estimating speciation and extinction rates is essential for understanding past and present biodiversity, but is challenging given the incompleteness of the rock and fossil records. Interest in this topic has led to a divergent suite of independent methods—paleontological estimates based on sampled stratigraphic ranges and phylogenetic estimates based on the observed branching times in a given phylogeny of living species. The fossilized birth–death (FBD) process is a model that explicitly recognizes that the branching events in a phylogenetic tree and sampled fossils were generated by the same underlying diversification process. A crucial advantage of this model is that it incorporates the possibility that some species may never be sampled. Here, we present an FBD model that estimates tree-wide diversification rates from stratigraphic range data when the underlying phylogeny of the fossil taxa may be unknown. The model can be applied when only occurrence data for taxonomically identified fossils are available, but still accounts for the incomplete phylogenetic structure of the data. We tested this new model using simulations and focused on how inferences are impacted by incomplete fossil recovery. We compared our approach with a phylogenetic model that does not incorporate incomplete species sampling and to three fossil-based alternatives for estimating diversification rates, including the widely implemented boundary-crosser and three-timer methods. The results of our simulations demonstrate that estimates under the FBD model are robust and more accurate than the alternative methods, particularly when fossil data are sparse, as the FBD model incorporates incomplete species sampling explicitly. 

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