Zombi: A phylogenetic simulator of trees, genomes and sequences that accounts for dead lineages

International audienceHere we present Zombi, a tool to simulate the evolution of species, genomes and sequences in silico, that considers for the first time the evolution of genomes in extinct lineages. It also incorporates various features that have not to date been combined in a single simulator, such as the possibility of generating species trees with a pre-defined variation of speciation and extinction rates through time, simulating explicitly intergenic sequences of variable length and outputting gene tree - species tree reconciliations

Megaphylogeny resolves global patterns of mushroom evolution

Mushroom-forming fungi (Agaricomycetes) have the greatest morphological diversity and complexity of any group of fungi. They have radiated into most niches and fulfil diverse roles in the ecosystem, including wood decomposers, pathogens or mycorrhizal mutualists. Despite the importance of mushroom-forming fungi, large-scale patterns of their evolutionary history are poorly known, in part due to the lack of a comprehensive and dated molecular phylogeny. Here, using multigene and genome-based data, we assemble a 5,284-species phylogenetic tree and infer ages and broad patterns of speciation/extinction and morphological innovation in mushroom-forming fungi. Agaricomycetes started a rapid class-wide radiation in the Jurassic, coinciding with the spread of (sub)tropical coniferous forests and a warming climate. A possible mass extinction, several clade-specific adaptive radiations and morphological diversification of fruiting bodies followed during the Cretaceous and the Paleogene, convergently giving rise to the classic toadstool morphology, with a cap, stalk and gills (pileate-stipitate morphology). This morphology is associated with increased rates of lineage diversification, suggesting it represents a key innovation in the evolution of mushroom-forming fungi. The increase in mushroom diversity started during the Mesozoic-Cenozoic radiation event, an era of humid climate when terrestrial communities dominated by gymnosperms and reptiles were also expanding.Fil: Varga, Torda. Hungarian Academy Of Sciences; HungríaFil: Krizsán, Krisztina. Hungarian Academy Of Sciences; HungríaFil: Földi, Csenge. Hungarian Academy Of Sciences; HungríaFil: Dima, Bálint. Eötvös Loránd University; HungríaFil: Sánchez-García, Marisol. Clark University; Estados UnidosFil: Lechner, Bernardo Ernesto. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Micología y Botánica. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Micología y Botánica; ArgentinaFil: Sánchez-Ramírez, Santiago. University of Toronto; CanadáFil: Szöllosi, Gergely J.. Eötvös Loránd University; HungríaFil: Szarkándi, János G.. University Of Szeged; HungríaFil: Papp, Viktor. Szent István University; HungríaFil: Albert, László. Hungarian Mycological Society; HungríaFil: Andreopoulos, William. United States Department Of Energy. Joint Genome Institute; Estados UnidosFil: Angelini, Claudio. Jardin Botanico Nacional Ma. Moscoso; República DominicanaFil: Antonín, Vladimír. Moravian Museum; República ChecaFil: Barry, Kerrie W.. United States Department Of Energy. Joint Genome Institute; Estados UnidosFil: Bougher, Neale L.. Western Australian Herbarium; AustraliaFil: Buchanan, Peter. Manaaki Whenua-landcare Research; Nueva ZelandaFil: Buyck, Bart. Muséum National d'Histoire Naturelle; FranciaFil: Bense, Viktória. Hungarian Academy Of Sciences; HungríaFil: Catcheside, Pam. State Herbarium Of South Australia; AustraliaFil: Chovatia, Mansi. United States Department Of Energy. Joint Genome Institute; Estados UnidosFil: Cooper, Jerry. Manaaki Whenua-landcare Research; Nueva ZelandaFil: Dämon, Wolfgang. Oberfeldstrasse 9; AustriaFil: Desjardin, Dennis. San Francisco State University; Estados UnidosFil: Finy, Péter. Zsombolyai U. 56.; HungríaFil: Geml, József. Naturalis Biodiversity Center; Países BajosFil: Haridas, Sajeet. United States Department Of Energy. Joint Genome Institute; Estados UnidosFil: Hughes, Karen. University of Tennessee; Estados UnidosFil: Justo, Alfredo. Clark University; Estados UnidosFil: Karasinski, Dariusz. Polish Academy of Sciences; Poloni

Lateral Gene Transfer from the Dead.

International audienceIn phylogenetic studies, the evolution of molecular sequences is assumed to have taken place along the phylogeny traced by the ancestors of extant species. In the presence of lateral gene transfer (LGT), however, this may not be the case, because the species lineage from which a gene was transferred may have gone extinct or not have been sampled. Because it is not feasible to specify or reconstruct the complete phylogeny of all species, we must describe the evolution of genes outside the represented phylogeny by modelling the speciation dynamics that gave rise to the complete phylogeny. We demonstrate that if the number of sampled species is small compared to the total number of existing species, the overwhelming majority of gene transfers involve speciation to, and evolution along extinct or unsampled lineages. We show that the evolution of genes along extinct or unsampled lineages can to good approximation be treated as those of independently evolving lineages described by a few global parameters. Using this result, we derive an algorithm to calculate the probability of a gene tree and recover the maximum likelihood reconciliation given the phylogeny of the sampled species. Examining 473 near universal gene families from 36 cyanobacteria, we find that nearly a third of transfer events - 28% - appear to have topological signatures of evolution along extinct species, but only approximately 6% of transfers trace their ancestry to before the common ancestor of the sampled cyanobacteria

Dating with transfers

National audienceTo reconstruct the timing of the diversification of life on Earth, biologists combine fossil evidence with inferences drawn from the comparison of genome sequences. These inferences are based on the molecular clock or on a softer version of it, the relaxed molecular clock. This approach consists of estimating the divergence between sequences and then, assuming that mutations occur clockwise , trying to determine the age of the ancestral sequence. This method can be refined by using diverse models that relax the hypothesis of constant pace of evolution and consider that sequences can evolve at different speeds. Some models assume that these rates of evolution are independent along the different branches of the tree relating the different sequences of DNA; some others consider that the rates are correlated among related branches, so the rate of a given branch is inherited to some extent from the parental one. Which is of these methods perform best is still heavily debated. In spite of the sophistication of the different models, calculating these rates is not a trivial problem and the best estimates of divergence time have usually very wide confidence intervals. To overcome this problem scientists can use fossils, that can be independently dated using methods such as stratigraphy or radiometry. Fossils are useful because they provide external information that can be used to constrain the positions of the nodes in a species tree, improving the accuracy of the estimates of the molecular clock. Combining relaxed molecular clock estimates and fossil is in active field of research in phylogenetics [1]. However, fossils are extremely scarce in the geological record. For about 80 % of the history of life, all organisms were unicellular, which means that finding fossils becomes an almost impossible task. Bones and hard shells are easy to be preserved but they just became frequent after the Cambrian explosion, when all the major animal clades appear at sudden. Before that Earth was dominated by bacteria and to a minor extent, small eukaryotes. These organisms are extremely small organisms with no hard parts that can fossilize easily. On top of that, for the few existent fossils we have there is very little certainty about the clades to which they belong, since morphological features cannot be used to place them in a phylogenetic tree. This means that if we are interested in studying what happened in the distant past, we have very little help coming from fossils and we must rely almost exclusively in the information conveyed by the DNA. As we previously stated, this is a hard problem since the estimates of the molecular clock can vary widely. We need accurate calculations if we want to know for example when did Eukaryotes diversify or when did cyanobacteria appear on Earth. To overcome these problems, we propose a new method of dating, based on the DNA sequence complementary to the molecular clock. Lateral gene transfer (LGT) is a common and almost universal phenomenon in nature, where different species (sometimes even species belonging to different domains) exchange genes. This can be detected using differences between species trees and gene trees. We do this using ALE, a method to reconcile species trees and gene trees that allows

Dating with transfers

National audienceTo reconstruct the timing of the diversification of life on Earth, biologists combine fossil evidence with inferences drawn from the comparison of genome sequences. These inferences are based on the molecular clock or on a softer version of it, the relaxed molecular clock. This approach consists of estimating the divergence between sequences and then, assuming that mutations occur clockwise , trying to determine the age of the ancestral sequence. This method can be refined by using diverse models that relax the hypothesis of constant pace of evolution and consider that sequences can evolve at different speeds. Some models assume that these rates of evolution are independent along the different branches of the tree relating the different sequences of DNA; some others consider that the rates are correlated among related branches, so the rate of a given branch is inherited to some extent from the parental one. Which is of these methods perform best is still heavily debated. In spite of the sophistication of the different models, calculating these rates is not a trivial problem and the best estimates of divergence time have usually very wide confidence intervals. To overcome this problem scientists can use fossils, that can be independently dated using methods such as stratigraphy or radiometry. Fossils are useful because they provide external information that can be used to constrain the positions of the nodes in a species tree, improving the accuracy of the estimates of the molecular clock. Combining relaxed molecular clock estimates and fossil is in active field of research in phylogenetics [1]. However, fossils are extremely scarce in the geological record. For about 80 % of the history of life, all organisms were unicellular, which means that finding fossils becomes an almost impossible task. Bones and hard shells are easy to be preserved but they just became frequent after the Cambrian explosion, when all the major animal clades appear at sudden. Before that Earth was dominated by bacteria and to a minor extent, small eukaryotes. These organisms are extremely small organisms with no hard parts that can fossilize easily. On top of that, for the few existent fossils we have there is very little certainty about the clades to which they belong, since morphological features cannot be used to place them in a phylogenetic tree. This means that if we are interested in studying what happened in the distant past, we have very little help coming from fossils and we must rely almost exclusively in the information conveyed by the DNA. As we previously stated, this is a hard problem since the estimates of the molecular clock can vary widely. We need accurate calculations if we want to know for example when did Eukaryotes diversify or when did cyanobacteria appear on Earth. To overcome these problems, we propose a new method of dating, based on the DNA sequence complementary to the molecular clock. Lateral gene transfer (LGT) is a common and almost universal phenomenon in nature, where different species (sometimes even species belonging to different domains) exchange genes. This can be detected using differences between species trees and gene trees. We do this using ALE, a method to reconcile species trees and gene trees that allows

Genome-scale coestimation of species and gene trees.

International audienceComparisons of gene trees and species trees are key to understanding major processes of genome evolution such as gene duplication and loss. Because current methods to reconstruct phylogenies fail to model the two-way dependency between gene trees and the species tree, they often misrepresent gene and species histories. We present a new probabilistic model to jointly infer rooted species and gene trees for dozens of genomes and thousands of gene families. We use simulations to show that this method accurately infers the species tree and gene trees, is robust to misspecification of the models of sequence and gene family evolution and provides a precise historic record of gene duplications and losses throughout genome evolution. We simultaneously reconstruct the history of mammalian species and their genes, based on 36 completely sequenced genomes, and use the reconstructed gene trees to infer the gene content and organization of ancestral mammalian genomes. We show that our method yields a more accurate picture of ancestral genomes than the trees available in the authoritative database Ensembl