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

   May, M.R. ; Contreras, D.L. ; Sundue, M.A. ; Nagalingum, N.S. ; Looy, C.V. ; Rothfels, C.J. ( March 2021 , Systematic biology)

   Ryan Folk (Ed.)

   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 ∼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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3. Phylogeny of gracillariid leaf‐mining moths: evolution of larval behaviour inferred from phylogenomic and Sanger data

   https://doi.org/10.1111/cla.12490  

   Li, Xuankun ; St Laurent, Ryan ; Earl, Chandra ; Doorenweerd, Camiel ; van Nieukerken, Erik J. ; Davis, Donald R. ; Johns, Chris A. ; Kawakita, Atsushi ; Kobayashi, Shigeki ; Zwick, Andreas ; et al ( October 2021 , Cladistics)

   Gracillariidae is the most taxonomically diverse cosmopolitan leaf‐mining moth family, consisting of nearly 2000 named species in 105 described genera, classified into eight extant subfamilies. The majority of gracillariid species are internal plant feeders as larvae, creating mines and galls in plant tissue. Despite their diversity and ecological adaptations, their phylogenetic relationships, especially among subfamilies, remain uncertain. Genomic data (83 taxa, 589 loci) were integrated with Sanger data (130 taxa, 22 loci), to reconstruct a phylogeny of Gracillariidae. Based on analyses of both datasets combined and analyzed separately, monophyly of Gracillariidae and all its subfamilies, monophyly of the clade “LAMPO” (subfamilies: Lithocolletinae, Acrocercopinae, Marmarinae, Phyllocnistinae, and Oecophyllembiinae) and relationships of its subclade “AMO” (subfamilies: Acrocercopinae, Marmarinae, and Oecophyllembiinae) were strongly supported. A sister‐group relationship of Ornixolinae to the remainder of the family, and a monophyletic leaf roller lineage (CallicercopsVári + Parornichinae) + Gracillariinae, as sister to the “LAMPO” clade were supported by the most likely tree. Dating analyses indicate a mid‐Cretaceous (105.3 Ma) origin of the family, followed by a rapid diversification into the nine subfamilies predating the Cretaceous–Palaeogene extinction. We hypothesize that advanced larval behaviours, such as making keeled or tentiform blotch mines, rolling leaves and galling, allowed gracillariids to better avoid larval parasitoids allowing them to further diversify. Finally, we stabilize the classification by formally re‐establishing the subfamily ranks of Marmarinaestat.rev., Oecophyllembiinaestat.rev. and Parornichinaestat.rev., and erect a new subfamily, Callicercopinae Li, Ohshima and Kawahara to accommodate the enigmatic genusCallicercops.

    

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4. A well‐resolved transcriptomic phylogeny of the mite harvestman family Pettalidae (Arachnida, Opiliones, Cyphophthalmi) reveals signatures of Gondwanan vicariance

   https://doi.org/10.1111/jbi.13828  

   Baker, Caitlin M. ; Boyer, Sarah L. ; Giribet, Gonzalo ( March 2020 , Journal of Biogeography)

   We explored the extent to which Gondwanan vicariance contributed to the circum‐Antarctic distribution of the mite harvestman family Pettalidae, a group of small, dispersal‐limited arachnids whose phylogeny has been poorly resolved, precluding rigorous biogeographic hypothesis testing.

   Continental landmasses of former temperate Gondwana (Chile, South Africa, Sri Lanka, Australia and New Zealand).

   Pettalidae, Opiliones.

   We generated transcriptomes for a phylogeny of 16 pettalids, spanning 9 genera. Data were analysed using maximum likelihood, Bayesian inference and coalescence methods. The phylogenetic position of the Sri Lankan genusPettaluswas further explored using quartet likelihood mapping and changes in gene likelihood scores. We also estimated divergence times and looked for signatures of extinction across Antarctica and central Australia using previously published phylogenies with near‐complete species sampling constrained to match our transcriptomic results. Finally, we estimated ancestral ranges and inferred instances of vicariance.

   We recovered a well‐supported topology with a division between taxa from landmasses that made up East Gondwana, and a grade of taxa from West Gondwana.Pettaluswas resolved either as the sister group of the Queensland‐endemicAustropurcellia, or as the sister group to a larger clade from East Gondwana, though favouringPettalus + Austropurcellia. Divergence times for multiple vicariance events coincided with Gondwana's breakup. Speciation–extinction analysis found one diversification process for the family: an initial burst of cladogenesis that slowed down through time.

   Given that the order of cladogenesis corresponds to the order in which Gondwana fragmented, and the concurrent timing of vicariance and rifting, Gondwanan breakup explains major biogeographic patterns in Pettalidae. Some divergences predate initial rifting, but there is no evidence oftrans‐oceanic dispersal. The Sri Lanka–eastern Australia relationship makes sense in the light of large‐scale extinction across Antarctica and central Australia; however, we find no clear signatures of mass extinction.

    

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5. Testing the phylogeny of Periptychidae and "archaic" Paleocene mammals uner different optimality criteria

   Puschel, H. P. ; Shelley, S. L. ; Williamson, T. E. ; Wible, J. R. ; Brusatte, S. L. ( October 2019 , Journal of Vertebrate Paleontology, Abstracts and Program)

   It is now well established that the end-Cretaceous mass extinction had enormous repercussions for mammalian evolution. Following the extinction, during the Paleocene, mammals started to radiate, occupying new and diverse ecological niches. However, the phylogenetic relationships between the socalled “archaic” mammals of this time, and their position within Placentalia, remain contentious. The Periptychidae are a clade of distinctive “archaic” ungulates, composed of ~17 genera of small to large bodied, highly bunodont, terrestrial herbivores that were among the first placental mammals to appear after the end-Cretaceous mass extinction. Although the Periptychidae has been historically considered a distinctive “condylarth” subgroup, their higherlevel relationships have been rarely tested. Here, we present an inclusive cladistic analysis to determine and test the phylogenetic affinities of Periptychidae and other key Paleocene groups within Placentalia under different cladistic optimality criteria. We scored 140 taxa for 503 dental, cranial and postcranial characters, incorporating new morphological and taxonomic data. The data were then subject to parsimony and Bayesian tree of morphological evolution, running 5000000 generations with samples every 200 generations and discarding 25% of the samples as burn-in. Stationarity was achieved and a 50 percent majority rule consensus tree from the sampled trees was obtained. The parsimony analysis recovered 48 most parsimonious trees. The two consensus trees derived from the different analyses are largely congruent and recover a monophyletic Periptychidae, although the parsimony consensus tree is better resolved. These results are consistent with simulation studies showing that parsimony tends to be more precise (more nodes reconstructed) than Bayesian analyses, although less accurate. The main topological differences between the results relate to the position of poorly known Puercan (earliest Paleocene) species. Our results affirm the monophyly of Periptychidae and its nesting within a group of “condylarths” positioned at the base of Laurasiatheria and closely related to Artiodactyla. Within Periptychidae we found support for the three major subfamilial divisions in both analyses. These results highlight the importance of using different optimality criteria when resolving a phylogeny and provide a new insight into how placental mammals were evolving after the end-Cretaceous extinction. Grant Information: CONICYT PFCHA/DOCTORADO BECAS CHILE/2018, European Research Council Starting Grant (ERC StG 2017, 756226, PalM), National Science Foundation (NSF EAR 1654952, DEB 1654949) 

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