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. 2009 Mar 10;106(10):3859-64.
doi: 10.1073/pnas.0807880106. Epub 2009 Feb 23.

Phylogenomic analyses support the monophyly of Excavata and resolve relationships among eukaryotic "supergroups"

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Phylogenomic analyses support the monophyly of Excavata and resolve relationships among eukaryotic "supergroups"

Vladimir Hampl et al. Proc Natl Acad Sci U S A. .

Abstract

Nearly all of eukaryotic diversity has been classified into 6 suprakingdom-level groups (supergroups) based on molecular and morphological/cell-biological evidence; these are Opisthokonta, Amoebozoa, Archaeplastida, Rhizaria, Chromalveolata, and Excavata. However, molecular phylogeny has not provided clear evidence that either Chromalveolata or Excavata is monophyletic, nor has it resolved the relationships among the supergroups. To establish the affinities of Excavata, which contains parasites of global importance and organisms regarded previously as primitive eukaryotes, we conducted a phylogenomic analysis of a dataset of 143 proteins and 48 taxa, including 19 excavates. Previous phylogenomic studies have not included all major subgroups of Excavata, and thus have not definitively addressed their interrelationships. The enigmatic flagellate Andalucia is sister to typical jakobids. Jakobids (including Andalucia), Euglenozoa and Heterolobosea form a major clade that we name Discoba. Analyses of the complete dataset group Discoba with the mitochondrion-lacking excavates or "metamonads" (diplomonads, parabasalids, and Preaxostyla), but not with the final excavate group, Malawimonas. This separation likely results from a long-branch attraction artifact. Gradual removal of rapidly-evolving taxa from the dataset leads to moderate bootstrap support (69%) for the monophyly of all Excavata, and 90% support once all metamonads are removed. Most importantly, Excavata robustly emerges between unikonts (Amoebozoa + Opisthokonta) and "megagrouping" of Archaeplastida, Rhizaria, and chromalveolates. Our analyses indicate that Excavata forms a monophyletic suprakingdom-level group that is one of the 3 primary divisions within eukaryotes, along with unikonts and a megagroup of Archaeplastida, Rhizaria, and the chromalveolate lineages.

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Conflict of interest statement

The authors declare no conflict of interest.

Figures

Fig. 1.
Fig. 1.
The phylogenetic tree estimated from the main dataset. This topology received the highest likelihood in the exhaustive search of unconstrained nodes using the WAG+ Γ model; branch-lengths were calculated in RAxML using the WAG+ Γ model. The representatives of the 6 supergroups are color-coded. Asterisks indicate the nodes that were not constrained during the exhaustive search. The numbers at the nodes indicate bootstrap support calculated by RAxML bootstrapping/PhyloBayes posterior probability. At nodes that were not constrained during the exhaustive search in the separate analysis (asterisks), the third number indicates the RELL bootstrap value. Branches that received maximum possible support by all methods are indicated by full circles. Dashes indicate bootstrap values
Fig. 2.
Fig. 2.
LB taxon removal. (A) The support for the nodes of interest calculated by RAxML bootstrapping is plotted against the number of long-branch taxa that were removed from the concatenate. The support for the unikont bipartition (X) is used as a control. A root position at the midpoint of the branch connecting unikonts with the rest of eukaryotes was used to calculate root-to-tip distances of taxa. The order of taxon removal is given on the x axis. (B) A maximum likelihood tree after removal of 14 taxa (gray box in the part A). The tree was constructed in RAxML using WAG+ Γ model and colored as described in Fig. 1. The numbers at the nodes indicate bootstrap support calculated by RAxML bootstrapping. Branches that received maximum support by all methods are indicated by full circles.
Fig. 3.
Fig. 3.
LB gene sequence removal. (A) The support for the nodes of interest calculated by RAxML bootstrapping is plotted against the number of longest-branch gene sequences that were removed from the concatenate. The support for unikonts (X) is used as a control. (B) A maximum likelihood tree after the removal of 1,750 of the longest-branch gene sequences (gray box in the A). The tree was constructed in RAxML using the WAG+ Γ model and colored as described in Fig. 1. The numbers at the nodes indicate bootstrap support calculated by RAxML bootstrapping. Branches that received maximum support by all methods are indicated by full circles, dashes indicate bootstrap values <50%.

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