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Bayesian Estimation of Species Divergence Times Integrating Fossil and Molecular Information

Lead Research Organisation: UNIVERSITY COLLEGE LONDON
Department Name: Genetics Evolution and Environment

Abstract

If our genes accumulate changes over time at a constant rate, the genetic distance between two species, measured by the number of changes accumulated, will be proportional to the time of species divergence. Thus molecules can serve as a clock, keeping time of species divergence by the accumulated changes. If fossil records or geological events can be used to assign an absolute geological time to a species divergence event on the phylogenetic tree, one can convert all calculated genetic distances into absolute geological times. This rationale for molecular clock dating has recently been extended to deal with local variation in evolutionary rate. Critical to molecular dating is the use of fossil information to calibrate the clock. In this project, we will develop statistical models and computer programs to analyze fossil and molecular data to accurately represent and incorporate the information in the fossil record in molecular dating analysis. The new methods will be applied to analyze datasets to estimate divergence times among mammals and to date viral transmission events.

Technical Summary

Two major improvements have recently been made to molecular clock dating methods: (i) relaxation of the clock assumption through local-clock models and (ii) incorporation of uncertainties in fossil calibrations. Nevertheless, representation of errors and uncertainties in the fossil record in a molecular dating analysis remains a challenging task. In this project, we will use models of clade divergences, fossil preservation and discovery, and morphological character evolution to derive statistical distributions of divergence times, which will be used as calibration densities for molecular clock dating. We will develop new models to describe the change in the evolutionary rate. The new models and methods will be applied to large datasets to date divergences among mammals and to date the host-switching events of the influenza virus.

Planned Impact

Accurate estimation of species divergence times is important to assessing the current biodiversity, and the impact of geological and environmental changes on biodiversity. The research results from this project will thus be useful for providing advice on decision making concerning biodiversity management and conservation policies.

Knowledge of absolute times of divergence between viral subtypes and of viral transmissions across species boundaries (such as the host switch of the flu virus from birds to humans) is important for understanding viral transmission and evolutionary dynamics. Such information may be useful to advise on public-health decisions and prevention of flu pandemics.

Publications

10 25 50

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De Baets K (2016) Tectonic blocks and molecular clocks. in Philosophical transactions of the Royal Society of London. Series B, Biological sciences

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Donoghue PC (2016) The evolution of methods for establishing evolutionary timescales. in Philosophical transactions of the Royal Society of London. Series B, Biological sciences

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Dos Reis M (2016) Notes on the birth-death prior with fossil calibrations for Bayesian estimation of species divergence times. in Philosophical transactions of the Royal Society of London. Series B, Biological sciences

 
Description Estimation of species divergence times allows biologists to place the speciation events into the correct geological and environmental framework, and to test hypotheses concerning species divergences and extinctions. Geological times of species divergences are estimated by using fossil information to calibrate the molecular species phylogeny, and the inference involve considerable uncertainties, because the molecular data provide information about the genetic distances but not absolute times and rates. We have developed a theory to characterize the uncertainties in the estimation of species divergence times, which can be used to determine whether the uncertainty in the estimates is mainly due to fossils or to molecules.
Exploitation Route Our new methods are used by many empirical biologists who analyze data from different groups of species such as mammals and flowering plants.
Sectors Education

Environment

URL http://abacus.gene.ucl.ac.uk/divtimes/index.html
 
Title Data from: Bayesian estimation of species divergence times using correlated quantitative characters 
Description Discrete morphological data have been widely used to study species evolution, but the use of quantitative (or continuous) morphological characters is less common. Here, we implement a Bayesian method to estimate species divergence times using quantitative characters. Quantitative character evolution is modelled using Brownian diffusion with character correlation and character variation within populations. Through simulations, we demonstrate that ignoring the population variation (or population "noise") and the correlation among characters leads to biased estimates of divergence times and rate, especially if the correlation and population noise are high. We apply our new method to the analysis of quantitative characters (cranium landmarks) and molecular data from carnivoran mammals. Our results show that time estimates are affected by whether the correlations and population noise are accounted for or ignored in the analysis. The estimates are also affected by the type of data analysed, with analyses of morphological characters only, molecular data only, or a combination of both; showing noticeable differences among the time estimates. Rate variation of morphological characters among the carnivoran species appears to be very high, with Bayesian model selection indicating that the independent-rates model fits the morphological data better than the autocorrelated-rates model. We suggest that using morphological continuous characters, together with molecular data, can bring a new perspective to the study of species evolution. Our new model is implemented in the MCMCtree computer program for Bayesian inference of divergence times. 
Type Of Material Database/Collection of data 
Year Produced 2019 
Provided To Others? Yes  
URL https://datadryad.org/stash/dataset/doi:10.5061/dryad.q7rf263
 
Title Data from: Tips and nodes are complimentary not competing approaches to the calibration of molecular clocks 
Description Molecular clock methodology provides the best means of establishing evolutionary timescales, the accuracy and precision of which remain reliant on calibration, traditionally based on fossil constraints on clade (node) ages. Tip calibration has been developed to obviate undesirable aspects of node calibration, including the need for maximum age constraints that are invariably very difficult to justify. Instead, tip calibration incorporates fossil species as dated tips alongside living relatives, potentially improving the accuracy and precision of divergence time estimates. We demonstrate that tip calibration yields node calibrations that violate fossil evidence, contributing to unjustifiably young and ancient age estimates, less precise and (presumably) accurate than conventional node calibration. However, we go on to show that node and tip calibrations are complementary, producing meaningful age estimates, with node minima enforcing realistic ages and fossil tips interacting with node calibrations to objectively define maximum age constraints on clade ages. Together, tip and node calibrations may yield evolutionary timescales that are better justified, more precise and accurate than either calibration strategy can achieve alone. 
Type Of Material Database/Collection of data 
Year Produced 2016 
Provided To Others? Yes  
URL https://datadryad.org/stash/dataset/doi:10.5061/dryad.2q3k2
 
Title Data from: Using phylogenomic data to explore the effects of relaxed clocks and calibration strategies on divergence time estimation: primates as a test case 
Description Primates have long been a test case for the development of phylogenetic methods for divergence time estimation. Despite a large number of studies, however, the timing of origination of crown Primates relative to the K-Pg boundary and the timing of diversification of the main crown groups remain controversial. Here we analysed a dataset of 372 taxa (367 Primates and 5 outgroups, 3.4 million aligned base pairs) that includes nine primate genomes. We systematically explore the effect of different interpretations of fossil calibrations and molecular clock models on primate divergence time estimates. We find that even small differences in the construction of fossil calibrations can have a noticeable impact on estimated divergence times, especially for the oldest nodes in the tree. Notably, choice of molecular rate model (auto-correlated or independently distributed rates) has an especially strong effect on estimated times, with the independent rates model producing considerably more ancient age estimates for the deeper nodes in the phylogeny. We implement thermodynamic integration, combined with Gaussian quadrature, in the program MCMCTree, and use it to calculate Bayes factors for clock models. Bayesian model selection indicates that the auto-correlated rates model fits the primate data substantially better, and we conclude that time estimates under this model should be preferred. We show that for eight core nodes in the phylogeny, uncertainty in time estimates is close to the theoretical limit imposed by fossil uncertainties. Thus, these estimates are unlikely to be improved by collecting additional molecular sequence data. All analyses place the origin of Primates close to the K-Pg boundary, either in the Cretaceous or straddling the boundary into the Palaeogene. 
Type Of Material Database/Collection of data 
Year Produced 2018 
Provided To Others? Yes  
URL https://datadryad.org/stash/dataset/doi:10.5061/dryad.c020q