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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’. 

Abstract Recent fossil discoveries from New Zealand have revealed a remarkably diverse assemblage of Paleocene stem group penguins. Here, we add to this growing record by describing nine new penguin specimens from the late Paleocene (upper Teurian local stage; 55.5–59.5 Ma) Moeraki Formation of the South Island, New Zealand. The largest specimen is assigned to a new species,Kumimanu fordycein. sp., which may have been the largest penguin ever to have lived. Allometric regressions based on humerus length and humerus proximal width of extant penguins yield mean estimates of a live body mass in the range of 148.0 kg (95% CI: 132.5 kg–165.3 kg) and 159.7 kg (95% CI: 142.6 kg–178.8 kg), respectively, forKumimanu fordycei. A second new species,Petradyptes stonehousein. gen. n. sp., is represented by five specimens and was slightly larger than the extant emperor penguinAptenodytes forsteri. Two small humeri represent an additional smaller unnamed penguin species. Parsimony and Bayesian phylogenetic analyses recoverKumimanuandPetradyptescrownward of the early Paleocene mainland NZ taxaWaimanuandMuriwaimanu, but stemward of the Chatham Island taxonKupoupou. These analyses differ, however, in the placement of these two taxa relative toSequiwaimanu,Crossvallia, andKaiika. The massive size and placement ofKumimanu fordyceiclose to the root of the penguin tree provide additional support for a scenario in which penguins reached the upper limit of sphenisciform body size very early in their evolutionary history, while still retaining numerous plesiomorphic features of the flipper. UUID:https://zoobank.org/15b1d5b2-a5a0-4aa5-ba0a-8ef3b8461730 

Myriad branches in the tree of life are intertwined through ecological relationships. Biologists have long hypothesized that intimate symbioses between lineages can influence diversification patterns to the extent that it leaves a topological imprint on the phylogenetic trees of interacting clades. Over the past few decades, cophylogenetic methods development has provided a toolkit for identifying such histories of codiversification, yet it is often difficult to determine which tools best suit the task at hand. In this review, we organize currently available cophylogenetic methods into three categories—pattern-based statistics, event-scoring methods, and more recently developed generative model–based methods—and discuss their assumptions and appropriateness for different types of cophylogenetic questions. We classify cophylogenetic systems based on their biological properties to provide a framework for empiricists investigating the macroevolution of symbioses. In addition, we provide recommendations for the next generation of cophylogenetic models that we hope will facilitate further methods development. 

Abstract Bayesian total-evidence approaches under the fossilized birth-death model enable biologists to combine fossil and extant data while accounting for uncertainty in the ages of fossil specimens, in an integrative phylogenetic analysis. Fossil age uncertainty is a key feature of the fossil record as many empirical data sets may contain a mix of precisely dated and poorly dated fossil specimens or deposits. In this study, we explore whether reliable age estimates for fossil specimens can be obtained from Bayesian total-evidence phylogenetic analyses under the fossilized birth-death model. Through simulations based on the example of the Baltic amber deposit, we show that estimates of fossil ages obtained through such an analysis are accurate, particularly when the proportion of poorly dated specimens remains low and the majority of fossil specimens have precise dates. We confirm our results using an empirical data set of living and fossil penguins by artificially increasing the age uncertainty around some fossil specimens and showing that the resulting age estimates overlap with the recorded age ranges. Our results are applicable to many empirical data sets where classical methods of establishing fossil ages have failed, such as the Baltic amber and the Gobi Desert deposits. [Bayesian phylogenetic inference; fossil age estimates; fossilized birth-death; Lagerstätte; total-evidence.] 

Abstract Gene flow is increasingly recognized as an important macroevolutionary process. The many mechanisms that contribute to gene flow (e.g. introgression, hybridization, lateral gene transfer) uniquely affect the diversification of dynamics of species, making it important to be able to account for these idiosyncrasies when constructing phylogenetic models. Existing phylogenetic‐network simulators for macroevolution are limited in the ways they model gene flow.We presentSiPhyNetwork, an R package for simulating phylogenetic networks under a birth–death‐hybridization process.Our package unifies the existing birth–death‐hybridization models while also extending the toolkit for modelling gene flow. This tool can create patterns of reticulation such as hybridization, lateral gene transfer, and introgression.Specifically, we model different reticulate events by allowing events to either add, remove or keep constant the number of lineages. Additionally, we allow reticulation events to be trait dependent, creating the ability to model the expanse of isolating mechanisms that prevent gene flow. This tool makes it possible for researchers to model many of the complex biological factors associated with gene flow in a phylogenetic context. 

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. 

No abstract available. 

New Zealand is a globally significant hotspot for seabird diversity, but the sparse fossil record for most seabird lineages has impeded our understanding of how and when this hotspot developed. Here, we describe multiple exceptionally well-preserved specimens of a new species of penguin from tightly dated (3.36–3.06 Ma) Pliocene deposits in New Zealand. Bayesian and parsimony analyses place Eudyptes atatu sp. nov. as the sister species to all extant and recently extinct members of the crested penguin genus Eudyptes . The new species has a markedly more slender upper beak and mandible compared with other Eudyptes penguins. Our combined evidence approach reveals that deep bills evolved in both crested and stiff-tailed penguins ( Pygoscelis ) during the Pliocene. That deep bills arose so late in the greater than 60 million year evolutionary history of penguins suggests that dietary shifts may have occurred as wind-driven Pliocene upwelling radically restructured southern ocean ecosystems. Ancestral area reconstructions using BioGeoBEARS identify New Zealand as the most likely ancestral area for total-group penguins, crown penguins and crested penguins. Our analyses provide a timeframe for recruitment of crown penguins into the New Zealand avifauna, indicating this process began in the late Neogene and was completed via multiple waves of colonizing lineages.