bis2c: lecture 10 extras on "new view of the tree of life" paper
TRANSCRIPT
Hug et al 2016
!1
Hug et al. 2016 Tree of Life
92 Bacterial Phyla25 Archaeal Phyla5 Eukaryotic Supergroups
Hug et al. Nature Microbiology. A new view of the tree of life. http://dx.doi.org/10.1038/nmicrobiol.2016.48
Laura Hug U. Waterloo
Jill Banfield UC Berkeley
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Hug et al 2016Hug et al. 2016 Bacteria
Hug et al. Nature Microbiology. A new view of the tree of life. http://dx.doi.org/10.1038/nmicrobiol.2016.48
Taxa Covered in Textbook
!3Hug et al. Nature Microbiology. A new view of the tree of life. http://dx.doi.org/10.1038/nmicrobiol.2016.48
!4
Hug et al 2016Phyla Never Grown in the Lab
Hug et al. Nature Microbiology. A new view of the tree of life. http://dx.doi.org/10.1038/nmicrobiol.2016.48
Hug et al 2016
!5
Hug et al. 2016 Archaea and Eukaryotes
Hug et al. Nature Microbiology. A new view of the tree of life. http://dx.doi.org/10.1038/nmicrobiol.2016.48
Hug et al 2016
!6
Hug et al. 2016 Archaea Phyla Never Grown in the Lab
Hug et al. Nature Microbiology. A new view of the tree of life. http://dx.doi.org/10.1038/nmicrobiol.2016.48
Major Groups by Evolutionary Distance
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The Dark Matter of Biology
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0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
© 2016 Macmillan Publishers Limited. All rights reserved
0.2
Korarchaeota
DiapherotritesNanohaloarchaeota
Unclassified archaea Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciArchaeoglobi, Methanomicrobia, Halobacteria
Aciduliprofundum, ThermoplasmataUncultured Thermoplasmata
ThermoplasmataOpisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
Cyanobacteria, Melainabacteria
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
Microgenomates Microgenomates Curtissbacteria Microgenomates Daviesbacteria
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates Amesbacteria
Microgenomates Shapirobacteria Microgenomates Beckwithbacteria, Pacebacteria, Collierbacteria
Microgenomates Gottesmanbacteria
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, MagasanikbacteriaParcubacteriaParcubacteria
Parcubacteria
ParcubacteriaParcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, KaiserbacteriaParcubacteria
Parcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomi Thermotogae
Omnitrophica Omnitrophica
Spirochaetes Spirochaetes
Hydrogenedentes NKB19 Deltaproteobacteria
Epsilonproteobacteria TM6
Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot.
NC10, Rokubacteria, Aminicenantes, Acidobacteria
Planctomycetes Chlamydiae
Lentisphaerae Verrucomicrobia
Verrucomicrobia
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32)
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
Latescibacteria WS3 Gemmatimonadetes, WOR-3, TA06
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)
Aigarchaeota, Cand. Caldiarchaeum subterraneum
Unclassified archaea
Parcubacteria
Candidate Phyla Radiation
Cyanobacteria, Melainabacteria
Deinococcus-Thermus Aquificae, Calescamantes EM19
Caldiserica, Dictyoglomiq ,q ,
ThermotogaeA ifi C
Omnitrophica Omnitrophica
pp
Spirochaetes SpirochaetesS i h t
Hydrogenedentes NKB19 Deltaproteobacteria
H d d t N
Epsilonproteobacteriab
TM6Alphaproteobacteria, Zetaproteobacteria, Betaproteobacteria, Gammaproteobacteria Chrysiogenetes, Deferribacteres
Modulibacteria, Tectomicrobia, Nitrospinae, Nitrospirae, Dadabacteria, Thermodesulfobacteria, Deltaprot. NC10, Rokubacteria, Aminicenantes, Acidobacteria
D f bD f b, , p ,, , p ,
Planctomycetespp
Chlamydiaey
LentisphaeraeC a yd Ch
Verrucomicrobia Verrucomicrobia
pp
RBX-1 WOR-1
Firmicutes, Tenericutes, Armatimonadetes, Chloroflexi, Actinobacteria Fusobacteria, Synergistetes
Uncultured bacteria (CP RIF32), y g, y g
Zixibacteria, Marinimicrobia, Caldithrix, Chlorobi, Ignavibacteria, Bacteroidetes Fibrobacteres
Cloacamonetes
Atribacteria (OP9) BRC1, Poribacteria
( )Latescibacteria WS3
Gemmatimonadetes, WOR-3, TA06b M
ElusimicrobiaUncultured bacteria
Uncultured bacteria (CP RIF1)O h
Dojkabacteria WS6 CPR3
Katanobacteria WWE3 Katanobacteria WWE3
Microgenomates RoizmanbacteriaMicrogenomates Roizmanbacteria
MicrogenomatesMicrogenomates Curtissbacteria
ggMicrogenomates Daviesbacteria
gg
Microgenomates Levybacteria
Microgenomates Woesebacteria Microgenomates AmesbacteriaMi t L b t i
Microgenomates ShapirobacteriaMi t W bMi t
Microgenomates Beckwithbacteria, Pacebacteria, CollierbacteriaMi Sh i b i
Microgenomates Gottesmanbacteriat R i b t i
g yg y
KAZAN CPR2, Saccharibacteria TM7
Berkelbacteria
Berkelbacteria
Berkelbacteria Berkelbacteria
CPR Uncultured unclassified bacteria Peregrinibacteria
Peregrinibacteria
Absconditabacteria SR1Gracilibacteria BD1-5 / GNO2
SM2F11Parcubacteria
Parcubacteria Kuenenbacteria, Falkowbacteria, Uhrbacteria, Magasanikbacteriate a
ParcubacteriaParcubacteria
Parcubacteria
Parcubacteria AbscAbs
Parcubacteria Azambacteria, Jorgensenbacteria, Wolfebacteria, Giovannonibacteria, Nomurabacteria, Campbellbacteria, Adlerbacteria, Kaiserbacteria
ggParcubacteriaParcubacteria Moranbacteria
ParcubacteriaParcubacteria Yanofskybacteria
P b i
Candidate Phyla Radiation
DiapherotritesNanohaloarchaeota
Unclassified archaea
Pacearchaeota
Woesearchaeota, Nanoarchaeota
Woesearchaeota
AltiarchaealesZ7ME43
Methanopyri, Methanococci, Methanobacteria, Hadesarchaea, ThermococciE43
Archaeoglobi, Methanomicrobia, Halobacteria, , ,, , ,
Aciduliprofundum, Thermoplasmatagg
Uncultured Thermoplasmatap ,p ,
Thermoplasmatap
Unclassified archaea
Korarchaeota
,
CrenarchaeotaCrenarchaeota
Thorarchaeota Lokiarchaeota
YNPFFA
Thaumarchaeota Thaumarchaeota
b l b
Aigarchaeota,FFA
Cand. Caldiarchaeum subterraneum
C b t i M l i b t i
Opisthokonta, Excavata, Archaeplastida Chromalveolata, Amoebozoa
Th h,
Th h tEukaryotes
Bacteria
Archaea
Katanobacteria WWE3
Bootstrap ≥ 85% 85% > Bootstrap ≥ 50%
Woesearchaeota, Nanoarchaeota
Figure 2 | A reformatted view of the tree in Fig. 1 in which each major lineage represents the same amount of evolutionary distance. The threshold forgroups (coloured wedges) was an average branch length of <0.65 substitutions per site. Notably, some well-accepted phyla become single groups andothers are split into multiple distinct groups. We undertook this analysis to provide perspective on the structure of the tree, and do not propose the resultinggroups to have special taxonomic status. The massive scale of diversity in the CPR and the large fraction of major lineages that lack isolated representatives(red dots) are apparent from this analysis. Bootstrap support values are indicated by circles on nodes—black for support of 85% and above, grey for supportfrom 50 to 84%. The complete ribosomal protein tree is available in rectangular format with full bootstrap values as Supplementary Fig. 1 and in Newickformat in Supplementary Dataset 2.
NATURE MICROBIOLOGY DOI: 10.1038/NMICROBIOL.2016.48 LETTERS
NATURE MICROBIOLOGY | www.nature.com/naturemicrobiology 3
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