Mitochondria are ATP-generating organelles whose endosymbiotic origin has been a key event in the evolution of eukaryotic cells. Despite overwhelming phylogenetic evidence for an alphaproteobacterial ancestry, efforts to pinpoint the closest relatives among the sampled alphaproteobacteria have been conflicting, complicating detailed inferences about the identity and nature of the mitochondrial ancestor. While most studies support that mitochondria evolved from an ancestor related to Rickettsiales, an order that includes several host-associated pathogenic and endosymbiotic lineages, others suggest that mitochondria evolved from a free-living group. Here we re-evaluate the phylogenetic placement of mitochondria. We employ genome-resolved binning of oceanic metagenome datasets and increase the genomic sampling of Alphaproteobacteria with twelve novel divergent clades, and one novel clade representing a sister group to all Alphaproteobacteria. Subsequent phylogenomic analyses that specifically address long branch attraction and compositional bias artefacts suggest that mitochondria did not evolve from Rickettsiales or any other currently recognized alphaproteobacterial lineage. Rather, our analyses indicate that mitochondria evolved from a proteobacterial lineage that branched off prior to the divergence of all sampled alphaproteobacteria. In light of this new result, previous hypotheses on the nature of the mitochondrial ancestor should be re-evaluated.
AtlanticOcean_100m_contigs.fa
Complete assembly of the Moca1 metagenome (SRA accession: SRP081826). Metagenome was derived from Atlantic Ocean sampled at 100m depth. See the associated publication for technical details.
AtlanticOcean_776m_contigs.fa
Complete assembly of the Moca2 metagenome (SRA accession: SRP081826). Metagenome was derived from Atlantic Ocean sampled at 776m depth. See the associated publication for technical details.
AtlanticOcean_2745m_contigs.fa
Complete assembly of the Moca3 metagenome (SRA accession: SRP081826). Metagenome was derived from Atlantic Ocean sampled at 2745m depth. See the associated publication for technical details.
AtlanticOcean_5002m_contigs.fa
Complete assembly of the Moca4 metagenome (SRA accession: SRP081826). Metagenome was derived from Atlantic Ocean sampled at 5002m depth. See the associated publication for technical details.
PacificOcean_115m_contigs.fasta
Complete assembly of the Tara Oceans metagenome (SRA accession: ERR594304). Metagenome was derived from Pacific Ocean sampled at 115m depth. See the associated publication for technical details.
PacificOcean_5m_contigs.fa
Complete assembly of the Tara Oceans metagenome (SRA accession: ERR594339). Metagenome was derived from Pacific Ocean sampled at 5m depth. See the associated publication for technical details.
PacificOcean_140m_contigs.fa
Complete assembly of the Tara Oceans metagenome (SRA accession: ERR599156). Metagenome was derived from Pacific Ocean sampled at 140m depth. See the associated publication for technical details.
alphaproteobacteria_untreated.aln
Concatenated supermatrix alignment of 72 genes conserved among alphaproteobacteria. See associated publication for technical details on how this alignment was prepared.
alphaproteobacteria_stationarytrimmed.aln
Stationary trimmed, concatenated supermatrix alignment of 72 genes conserved among alphaproteobacteria. See associated publication for technical details on how this alignment was prepared.
alphaproteobacteria_SR4recoded.aln
SR4 recoded, concatenated supermatrix alignment of 72 genes conserved among alphaproteobacteria. See associated publication for technical details on how this alignment was prepared.
alphaproteobacteria_mitochondria_untreated.aln
Concatenated supermatrix alignment of 24 genes conserved among alphaproteobacteria and mitochondrial genomes. See associated publication for technical details on how this alignment was prepared.
alphaproteobacteria_mitochondria_stationarytrimmed.aln
Stationary-trimmed, concatenated supermatrix alignment of 24 genes conserved among alphaproteobacteria and mitochondrial genomes. See associated publication for technical details on how this alignment was prepared.
alphaproteobacteria_mitochondria_SR4recoded.aln
SR4 recoded, concatenated supermatrix alignment of 24 genes conserved among alphaproteobacteria and mitochondrial genomes. See associated publication for technical details on how this alignment was prepared.
alphamito38.concat.untreated.aln
Concatenated supermatrix alignment of 38 genes conserved among alphaproteobacteria and mitochondrial genomes. See associated publication for technical details on how this alignment was prepared.
alphamito38.concat.stattrim.aln
Stationary-trimmed, concatenated supermatrix alignment of 38 genes conserved among alphaproteobacteria and mitochondrial genomes. See associated publication for technical details on how this alignment was prepared.
nuclearEncoded29_concat_Composite.untreated.aln
Concatenated supermatrix alignment of 29 genes conserved among alphaproteobacteria and mitochondrial genes encoded on nuclear eukaryotic genomes. See associated publication for technical details on how this alignment was prepared.
nuclearEncoded29_concat_Composite.stattrim.aln
Stationary-trimmed, concatenated supermatrix alignment of 29 genes conserved among alphaproteobacteria and mitochondrial genes encoded on nuclear eukaryotic genomes. See associated publication for technical details on how this alignment was prepared.
alphamito24.otherMitoTaxa.concat.untreated.aln
Concatenated supermatrix alignment of 24 genes conserved among alphaproteobacteria and mitochondrial genomes. Here, a more diverse set of mitochondria has been selected. See associated publication for technical details on how this alignment was prepared.
alphamito24.otherMitoTaxa.concat.stattrim.aln
Stationary-trimmed, concatenated supermatrix alignment of 24 genes conserved among alphaproteobacteria and mitochondrial genomes. Here, a more diverse set of mitochondria has been selected. See associated publication for technical details on how this alignment was prepared.
alphamito24.concat.NoOutgroups.untreated.aln
Concatenated supermatrix alignment of 24 genes conserved among alphaproteobacteria and mitochondrial genomes. Here, the outgroups MarineProteo1, Magnetococcales, Beta- and Gammaproteobacteria have been removed. See associated publication for technical details on how this alignment was prepared.
alphamito24.concat.NoOutgroups.stattrim.aln
Stationary-trimmed, concatenated supermatrix alignment of 24 genes conserved among alphaproteobacteria and mitochondrial genomes. Here, the outgroups MarineProteo1, Magnetococcales, Beta- and Gammaproteobacteria have been removed. See associated publication for technical details on how this alignment was prepared.