Bacterial phyla
Phylogenetic tree showing the diversity of Bacteria, Archaea, and Eukaryota.[ 1] Major lineages are assigned arbitrary colours and named, with well-characterized lineage names, in italics. Lineages lacking an isolated representative are highlighted with non-italicized names and red dots.
Phyla or divisions of the domain Bacteria
Bacterial phyla constitute the major lineages of the domain Bacteria . While the exact definition of a bacterial phylum is debated, a popular definition is that a bacterial phylum is a monophyletic lineage of bacteria whose 16S rRNA genes share a pairwise sequence identity of ~75% or less with those of the members of other bacterial phyla.[ 2]
It has been estimated that ~1,300 bacterial phyla exist.[ 2] As of May 2020, 41 bacterial phyla are formally accepted by the LPSN ,[ 3] 89 bacterial phyla are recognized on the Silva database , dozens more have been proposed,[ 4] [ 5] and hundreds likely remain to be discovered.[ 2] As of 2017, approximately 72% of widely recognized bacterial phyla were candidate phyla [ 6] (i.e. have no cultured representatives).
The rank of phylum has been included in the rules of the International Code of Nomenclature of Prokaryotes , using the ending –ota for phylum names that must be based on the name of a genus as its nomenclatural type.[ 7] [ 8]
List of bacterial phyla
The following is a list of bacterial phyla that have been validly published (not current).
Phylum
Alternative names
Group
Cultured representative
Notes
10bav-F6[ 9]
No
"Abawacabacteria "[ 4] [ 10]
RIF46
CPR; Gracilibacteria-related CPR
No
"Abditibacteriota "[ 11]
FBP
Yes[ 11]
"Absconditabacteria "[ 12] [ 10]
SR1
CPR; Gracilibacteria-related CPR
No
ABY1[ 13]
OD1-ABY1[ 14]
CPR; Parcubacteria
No
"Bipolaricaulota "[ 15]
OP1, "Acetothermia"
Acidobacteriota
"Acidobacteria"
Yes[ 16]
Actinomycetota
"Actinobacteria"
Terrabacteria
Yes[ 17]
"Adlerbacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 4
No
"Aerophobota " / "Aerophobetes"
CD12, BHI80-139
"Amesbacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Andersenbacteria "[ 4]
RIF9
CPR; Parcubacteria; Parcubacteria 4-related
No
Armatimonadota [ 15]
"Armatimonadetes", OP10
Terrabacteria
Yes[ 19]
"Aminicenantes "[ 15]
OP8
AncK6[ 9]
Apal-E12[ 9]
Atribacterota [ 15]
OP9, JS1
No
Aquificota
"Aquificae"
"Azambacteria " i[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; unclassified Parcubacteria
No
split by Anantharaman et al. …
"Azambacteria " ii[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; unclassified Parcubacteria
No
… (Oct 2016) as being polyphyletic
Bacteroidota
"Bacteroidetes"
FCB group
Yes
Balneolota [ 20]
Yes
Bdellovibrionota
"Beckwithbacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Berkelbacteria "[ 21] [ 10]
ACD58
CPR; Saccharibacteria-related CPR
No
BHI80-139[ 9]
"Blackburnbacteria "[ 4]
RIF35
CPR; Microgenomates
No
"Brennerbacteria "[ 4] [ 10]
RIF18
CPR; Parcubacteria; Parcubacteria 3
No
"Brownbacteria "[ 22]
CPR; Parcubacteria; unclassified Parcubacteria
No
"Buchananbacteria "[ 4] [ 10]
RIF37
CPR; Parcubacteria; Parcubacteria 1
No
Caldisericota [ 15]
OP5,[ 23] "Caldiserica"
FCB group
Yes[ 24]
Calditrichota [ 25]
FCB group[ 26]
"Calescamantes "
EM19
"Campbellbacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 4
No
seem to be polyphyletic: two clades
Campylobacterota
Chlamydiota
"Chlamydiae"[ 27]
PVC group
Chlorobiota
"Chlorobi"
FCB group
Chloroflexota
"Chloroflexi"
"Terrabacteria"
Chisholmbacteria [ 4]
RIF36
CPR; "Microgenomates"
No
Chrysiogenota
"Chrysiogenetes"
"Cloacimonetes "[ 28]
WWE1
FCB group[ 26]
"Coatesbacteria "[ 4]
RIF8
No
"Collierbacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Colwellbacteria "[ 4] [ 10]
RIF41
CPR; Parcubacteria; Parcubacteria 3
No
Coprothermobacterota
"Curtissbacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
CPR-1[ 1]
CPR
No
CPR-3[ 1]
CPR
No
"Cyanobacteria "
Terrabacteria
"Dadabacteria "[ 29]
No
"Daviesbacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Delphibacteria "[ 6]
FCB group
No
"Delongbacteria "[ 4]
RIF26, H-178
No
Deferribacterota
Deferribacteres
Deinococcota
Deinococcus–Thermus
Terrabacteria
"Dependentiae "[ 30]
TM6
Dictyoglomota
Dictyoglomi[ 31]
"Dojkabacteria "[ 10]
WS6
CPR; Microgenomates-related CPR
"Dormibacteraeota "[ 32]
AD3
No
"Doudnabacteria "[ 18] [ 10]
SM2F11
CPR; Parcubacteria; Parcubacteria 1-related
No
"Edwardsbacteria "[ 5] [ 4]
RIF29, UBP-2 [ 33]
No
"Eisenbacteria "[ 4]
RIF28
FCB group
No
Elusimicrobiota
Elusimicrobia, OP7, Termite Group 1 (TG1)[ 23]
Yes[ 34]
"Eremiobacteraeota "[ 35] [ 32]
WPS-2, Palusbacterota[ 36]
No
"Falkowbacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 1
No
"Fermentibacteria "[ 37]
Hyd24-12
No
"Fertabacteria "[ 6]
CPR; Gracilibacteria-related CPR
No
Fibrobacterota
"Fibrobacteres"
FCB group
"Firestonebacteria "[ 4]
RIF1
No
"Fervidibacteria "
OctSpa1-106
"Fischerbacteria "[ 4]
RIF25
No
Bacillota
"Firmicutes"
Terrabacteria
"Fraserbacteria "[ 4]
RIF31
No
Fusobacteriota
"Fusobacteria"
Gemmatimonadota
Gemmatimonadetes[ 38]
FCB group[ 26]
Yes[ 38]
"Glassbacteria "[ 4]
RIF5
No
"Giovannonibacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 4-related
No
"Gottesmanbacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Gracilibacteria "[ 39] [ 10]
GN02, BD1-5, SN-2
CPR; Patescibacteria; Gracilibacteria-related CPR
No
"Gribaldobacteria "[ 4] [ 10]
CPR; Parcubacteria; Parcubacteria 2
No
"Handelsmanbacteria "[ 4]
RIF27
No
"Harrisonbacteria "[ 4] [ 10]
RIF43
CPR; Parcubacteria; Parcubacteria 3
No
"Howlettbacteria "[ 10]
CPR; Saccharibacteria-related CPR
No
"Hugbacteria "[ 22]
CPR; Parcubacteria; unclassified Parcubacteria
No
"Hydrogenedentes "
NKB19
No
Ignavibacteriota
"Ignavibacteria", ZB1
FCB group
"Jacksonbacteria "[ 4] [ 10]
RIF38
CPR; Parcubacteria; Parcubacteria 1
No
"Jorgensenbacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 3
No
"Kaiserbacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 4
No
"Katanobacteria "[ 40] [ 10]
WWE3
CPR; Microgenomates-related
No
"Kazanbacteria "[ 10] [ 4]
Kazan
CPR; Saccharibacteria-related CPR
No
"Kerfeldbacteria "[ 4] [ 10]
RIF4
CPR; Parcubacteria; Parcubacteria 1
No
Kiritimatiellota
"Komeilibacteria "[ 4] [ 10]
RIF6
CPR; Parcubacteria; Parcubacteria 1
No
sometimes misspelled as "Komelilbacteria"[ 4]
"Kryptonia "[ 41]
No
KSB1
No
"Krumholzibacteriota "[ 33]
"Kuenenbacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 1
No
"Lambdaproteobacteria "[ 4]
RIF24
Proteobacteria
No
"Latescibacteria "
WS3
FCB group[ 26]
No
LCP-89[ 42]
Lentisphaerota
"Lentisphaerae", vadinBE97
PVC group
"Levybacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Lindowbacteria "[ 4]
RIF2
CPR; Saccharibacteria-related CPR
No
"Liptonbacteria "[ 4] [ 10]
RIF42
CPR; Parcubacteria; Parcubacteria 3
No
"Lloydbacteria "[ 4] [ 10]
RIF45
CPR; Parcubacteria; Parcubacteria 4
No
"Magasanikbacteria "[ 18] [ 43] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 1
No
"Margulisbacteria "[ 4]
RIF30
No
"Marinimicrobia "
SAR406, Marine Group A
FCB group[ 26]
Yes
"Melainabacteria "[ 44]
No
"Microgenomates "[ 45]
OP11
CPR; Patescibacteria
No
Superphylum
"Modulibacteria "[ 39] [ 46]
KSB3, GN06
No
"Moranbacteria "[ 18] [ 10]
OD1-i[ 18]
CPR; Patescibacteria; Parcubacteria; unclassified Parcubacteria
No
"Muproteobacteria "[ 4]
RIF23
Proteobacteria
No
Myxococcota
NC10 [ 47] [ 13]
No
"Nealsonbacteria "[ 4] [ 10]
RIF40
CPR; Parcubacteria; Parcubacteria 2
No
"Niyogibacteria "[ 4]
RIF11
CPR; Parcubacteria; Parcubacteria 4-related
No
Nitrospinota
"Nitrospinae"[ 48]
Yes[ 49] [ 50]
Nitrospirota
"Nitrospirae"
Yes
"Nomurabacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 1
No
"Omnitrophica "[ 15]
OP3
PVC group
No
"Pacebacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Parcubacteria "[ 12]
OD1
CPR
No
Superphylum
"Parcubacteria" 1[ 10]
CPR; Parcubacteria
No
"Parcubacteria" 2[ 10]
CPR; Parcubacteria
No
"Parcubacteria" 3[ 10]
CPR; Parcubacteria
No
"Parcubacteria" 4[ 10]
CPR; Parcubacteria
No
"Parcunitrobacteria "[ 51]
CPR; Parcubacteria; unclassified Parcubacteria[ 52]
No
Superphylum
PAUC34f [ 53]
sponge‐associated unclassified lineage (SAUL)
FCB group
"Peregrinibacteria "[ 54] [ 55] [ 56] [ 57] [ 10]
PER
CPR; Gracilibacteria-related CPR
No
"Peribacteria "[ 10]
CPR; Gracilibacteria-related CPR
No
Planctomycetota
"Planctomycetes"
PVC group
"Poribacteria "[ 58]
PVC group
"Portnoybacteria "[ 4]
RIF22
CPR; Parcubacteria; Parcubacteria 4-related
No
Pseudomonadota
"Proteobacteria"
"Raymondbacteria "[ 4]
RIF7
No
Riflebacteria[ 4]
RIF32
No
Rhodothermota
"Roizmanbacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Rokubacteria "[ 29]
No
"Ryanbacteria "[ 4] [ 10]
RIF10
CPR; Parcubacteria; Parcubacteri 4-related
No
"Saccharibacteria "[ 30] [ 10]
TM7
CPR; Saccharibacteria-related CPR
Yes
"Saltatorellota "[ 59]
"Schekmanbacteria "[ 4]
RIF3
Proteobacteria
No
"Shapirobacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Spechtbacteria "[ 4] [ 10]
RIF19
CPR; Parcubacteria; Parcubacteria 2
No
Spirochaetota
"Spirochaetes"
"Staskawiczbacteria "[ 4] [ 10]
RIF20
CPR; Parcubacteria; Parcubacteria 2
No
"Sumerlaeota "[ 60] [ 61]
BRC1
"Sungbacteria "[ 4] [ 10]
RIF17
CPR; Parcubacteria; Parcubacteria 4-related
No
Synergistota
"Synergistetes"
TA06[ 62]
No
"Tagabacteria "[ 4] [ 10]
RIF12
CPR; Parcubacteria; Parcubacteria 4-related
No
"Taylorbacteria "[ 4] [ 10]
RIF16
CPR; Parcubacteria; Parcubacteria 4
No
"Tectomicrobia "[ 63]
Mycoplasmatota
"Tenericutes"
"Terrybacteria "[ 4] [ 10]
RIF13
CPR; Parcubacteria; Parcubacteria 2
No
Thermodesulfobacteriota
"Thermodesulfobacteria"
Thermomicrobiota
"Thermomicrobia"
Thermotogota
"Thermotogae", OP2, EM3[ 23]
Yes[ 64]
"Torokbacteria "[ 10]
CPR; Parcubacteria; unclssified Parcubacteria
No
UBP-1[ 5]
No
UBP-3[ 5]
No
UBP-4[ 5]
No
UBP-5[ 5]
No
UBP-6[ 5]
No
UBP-7[ 5]
No
UBP-8[ 5]
No
UBP-9[ 5]
No
UBP-10[ 5]
No
UBP-11[ 5]
No
UBP-12[ 5]
No
UBP-13[ 5]
No
UBP-14[ 5]
No
UBP-15[ 5]
No
UBP-16[ 5]
No
UBP-17[ 5]
No
"Uhrbacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 1
No
seem to be polyphyletic: two clades
"Veblenbacteria "[ 4]
RIF39
CPR; Parcubacteria; Parcubacteria 1-related
No
Verrucomicrobiota
"Verrucomicrobia"
PVC group
"Vogelbacteria "[ 4] [ 10]
RIF14
CPR; Parcubacteria; Parcubacteria 4
No
"Wallbacteria "[ 4]
RIF33
No
"Wildermuthbacteria "[ 4] [ 10]
RIF21
CPR; Parcubacteria; Parcubacteria 2
No
"Wirthbacteria "[ 65]
CPR-related bacteria
No
"Woesebacteria "[ 18]
CPR; Patescibacteria; Microgenomates
No
"Wolfebacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; Parcubacteria 3
No
"Woykebacteria "[ 4] [ 22]
RIF34
CPR; Microgenomates
No
WOR-1[ 62]
No
WOR-2[ 62]
No
WOR-3[ 62]
No
"Yanofskybacteria "[ 18] [ 10]
CPR; Patescibacteria; Parcubacteria; unclassified Parcubacteria
No
"Yonathbacteria "[ 4] [ 10]
RIF44
CPR; Parcubacteria; Parcubacteria 4
No
"Zambryskibacteria "[ 4] [ 10]
RIF15
CPR; Parcubacteria; Parcubacteria 4
No
ZB2
OD1-ZB2[ 14]
CPR; Parcubacteria
No
"Zixibacteria "[ 66]
FCB group
No
Supergroups
Despite the unclear branching order for most bacterial phyla, several groups of phyla consistently cluster together and are referred to as supergroups or superphyla. In some instances, bacterial clades clearly consistently cluster together but it is unclear what to call the group. For example, the Candidate Phyla Radiation includes the Patescibacteria group which includes Microgenomates group which includes over 11 bacterial phyla.
Candidate phyla radiation (CPR)
The CPR is a descriptive term referring to a massive monophyletic radiation of candidate phyla that exists within the Bacterial domain.[ 67] It includes two main clades, the Microgenomates and Parcubacteria groups, each containing the eponymous superphyla and a few other phyla.
Patescibacteria
The superphylum Patescibacteria was originally proposed to encompass the phyla Microgenomates (OP11), Parcubacteria (OD1), and Gracilibacteria (GNO2 / BD1-5).[ 26] More recent phylogenetic analyses show that the last common ancestor of these taxa is the same node as that of CPR.[ 68]
Sphingobacteria
The Sphingobacteria (FCB group) includes Bacteroidota, Calditrichota, Chlorobiota, candidate phylum "Cloacimonetes", Fibrobacterota, Gemmatimonadota, Ignavibacteriota, candidate phylum "Latescibacteria", candidate phylum "Marinimicrobia", and candidate phylum "Zixibacteria".[ 26] [ 69]
Microgenomates
Microgenomates was originally thought to be a single phylum although evidence suggests it actually encompasses over 11 bacterial phyla,[ 18] [ 4] including Curtisbacteria, Daviesbacteria, Levybacteria, Gottesmanbacteria, Woesebacteria, Amesbacteria, Shapirobacteria, Roizmanbacteria, Beckwithbacteria, Collierbacteria, Pacebacteria.
Parcubacteria
Parcubacteria was originally described as a single phylum using fewer than 100 16S rRNA sequences. With a greater diversity of 16S rRNA sequences from uncultured organisms now available, it is estimated it may consist of up to 28 bacterial phyla.[ 2] In line with this, over 14 phyla have now been described within the Parcubacteria group,[ 18] [ 4] including Kaiserbacteria, Adlerbacteria, Campbellbacteria, Nomurabacteria, Giovannonibacteria, Wolfebacteria, Jorgensenbacteria, Yanofskybacteria, Azambacteria, Moranbacteria, Uhrbacteria, and Magasanikbacteria.
Proteobacteria
It has been proposed that some classes of the phylum Proteobacteria may be phyla in their own right, which would make Proteobacteria a superphylum.[ 70] For example, the Deltaproteobacteria group does not consistently form a monophyletic lineage with the other Proteobacteria classes.[ 71]
Planctobacteria
The Planctobacteria (PVC group) includes Chlamydiota , Lentisphaerota , candidate phylum "Omnitrophica ", Planctomycetota , candidate phylum "Poribacteria ", and Verrucomicrobiota .[ 26] [ 69]
Terrabacteria
The proposed superphylum, Terrabacteria ,[ 72] includes Actinomycetota , "Cyanobacteria "/"Melainabacteria "-group , Deinococcota , Chloroflexota , Bacillota , and candidate phylum OP10.[ 72] [ 73] [ 26] [ 69]
Cryptic superphyla
Several candidate phyla (Microgenomates , Omnitrophica, Parcubacteria, and Saccharibacteria ) and several accepted phyla (Elusimicrobiota , Caldisericota , and Armatimonadota ) have been suggested to actually be superphyla that were incorrectly described as phyla because rules for defining a bacterial phylum are lacking or due to a lack of sequence diversity in databases when the phylum was first established.[ 2] For example, it is suggested that candidate phylum Parcubacteria is actually a superphylum that encompasses 28 subordinate phyla and that phylum Elusimicrobia is actually a superphylum that encompasses 7 subordinate phyla.[ 70]
Historical perspective
Atomic structure of the 30S ribosomal Subunit from Thermus thermophilus of which 16S makes up a part. Proteins are shown in blue and the single RNA strand in tan.[ 74]
Given the rich history of the field of bacterial taxonomy and the rapidity of changes therein in modern times, it is often useful to have a historical perspective on how the field has progressed in order to understand references to antiquated definitions or concepts.
When bacterial nomenclature was controlled under the Botanical Code , the term division was used, but now that bacterial nomenclature (with the exception of cyanobacteria ) is controlled under the Bacteriological Code , the term phylum is preferred.
In 1987, Carl Woese , regarded as the forerunner of the molecular phylogeny revolution, divided Eubacteria into 11 divisions based on 16S ribosomal RNA (SSU) sequences, listed below.[ 75] [ 76]
Purple Bacteria and their relatives (later renamed Proteobacteria [ 77] )
alpha subdivision (purple non-sulfur bacteria , rhizobacteria , Agrobacterium , Rickettsiae , Nitrobacter )
beta subdivision (Rhodocyclus , (some) Thiobacillus , Alcaligenes , Spirillum , Nitrosovibrio )
gamma subdivision (enterics, fluorescent pseudomonads , purple sulfur bacteria , Legionella , (some) Beggiatoa )
delta subdivision (Sulfur and sulfate reducers (Desulfovibrio ), Myxobacteria , Bdellovibrio )
Gram-positive Eubacteria[ Note 1]
Cyanobacteria and chloroplasts (Aphanocapsa , Oscillatoria , Nostoc , Synechococcus , Gloeobacter , Prochloron )
Spirochetes and relatives
Green sulfur bacteria (Chlorobium , Chloroherpeton )
Bacteroides , Flavobacteria and relatives (later renamed Bacteroidetes
Planctomyces and relatives (later renamed Planctomycetes )
Chlamydiae (Chlamydia psittaci , Chlamydia trachomatis )
Radioresistant micrococci and relatives (later renamed Deinococcus–Thermus [ 82] or Thermi )[ Note 3]
Green non-sulfur bacteria and relatives (later renamed Chloroflexi [ 86] )
Thermotogae (Thermotoga maritima )
Traditionally, phylogeny was inferred and taxonomy established based on studies of morphology. The advent of molecular phylogenetics has allowed for improved elucidation of the evolutionary relationship of species by analyzing their DNA and protein sequences, for example their ribosomal DNA .[ 87] The lack of easily accessible morphological features, such as those present in animals and plants , hampered early efforts of classification and resulted in erroneous, distorted and confused classification, an example of which, noted Carl Woese , is Pseudomonas whose etymology ironically matched its taxonomy, namely "false unit".[ 75] Many bacterial taxa were re-classified or re-defined using molecular phylogenetics.
The advent of molecular sequencing technologies has allowed for the recovery of genomes directly from environmental samples (i.e. bypassing culturing), leading to rapid expansion of our knowledge of the diversity of bacterial phyla. These techniques are genome-resolved metagenomics and single-cell genomics .
See also
References
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^ Cavalier-Smith, T (2002). "The neomuran origin of Archaebacteria, the negibacterial root of the universal tree and bacterial megaclassification" . International Journal of Systematic and Evolutionary Microbiology . 52 (Pt 1): 7–76. doi :10.1099/00207713-52-1-7 . PMID 11837318 .
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Terrabacteria BV1, BV3, BV5
others Cyanoprokaryota
Cyanobacteriota
"Margulisiibacteriota "
"Marinamargulisbacteria"
"Riflemargulisbacteria"
"Saganbacteria"
"Termititenacia"
Firmicutes BV3 Low GC
Bacillota
Bacillota A
"Clostridiia "
"Thermoanaerobacteria"
"Thermosediminibacteria"
Bacillota D
Bacillota E
"Sulfobacillia"
"Symbiobacteriia"
"Thermaerobacteria"
Bacillota G
"Hydrogenisporia"
Limnochordia
"Desulfotomaculota "
"Carboxydocellia"
"Carboxydothermia"
"Dehalobacteriia"
"Desulfitobacteriia"
"Desulfotomaculia"
"Moorellia"
"Peptococcia"
"Syntrophomonadia"
"Thermincolia"
"Halanaerobiaeota"
"Selenobacteria"
CPR group Thermotogida
Fusobacterida Hydrobacteria BV2, BV4
others