International Journal of Systematic and Evolutionary Microbiology (2015), 65, 1070–1074 DOI 10.1099/ijs.0.

000034

Taxonomic Proposal to consistently apply the International

Note Code of Nomenclature of Prokaryotes (ICNP) to
names of the oxygenic photosynthetic bacteria
(cyanobacteria), including those validly published
under the International Code of Botanical
Nomenclature (ICBN)/International Code of
Nomenclature for algae, fungi and plants (ICN), and
proposal to change Principle 2 of the ICNP
Alexander V. Pinevich
Correspondence Department of Microbiology, St. Petersburg State University, 199178 St. Petersburg, Russia
Alexander V. Pinevich
pinevich.A@mail.ru
This taxonomic note was motivated by the recent proposal [Oren & Garrity (2014) Int J Syst Evol
Microbiol 64, 309–310] to exclude the oxygenic photosynthetic bacteria (cyanobacteria) from the
wording of General Consideration 5 of the International Code of Nomenclature of Prokaryotes
(ICNP), which entails unilateral coverage of these prokaryotes by the International Code of
Nomenclature for algae, fungi, and plants (ICN; formerly the International Code of Botanical
Nomenclature, ICBN). On the basis of key viewpoints, approaches and rules in the systematics,
taxonomy and nomenclature of prokaryotes it is reciprocally proposed to apply the ICNP to names
of cyanobacteria including those validly published under the ICBN/ICN. For this purpose, a
change to Principle 2 of the ICNP is proposed to enable validation of cyanobacterial names
published under the ICBN/ICN rules.

Progress towards the dual coverage of Cyanophyceae (the In this connection, the original wording of General
algal class name coined by J. Sachs in 1874)/Cyanophyta (the Consideration 5 of the ICNB (Lapage et al., 1992) should
algal division name coined by F. Steinecke in 1931)/oxygenic be noted: ‘This Code of Nomenclature of Bacteria applies to
photosynthetic bacteria/cyanobacteria (trivial terms intro- all bacteria’. A proposed additional note to General Consider-
duced by R. Stanier in 1974) by the International Code of ation 5 (Tindall, 1999) states that: ‘The term ‘‘bacteria’’
Botanical Nomenclature (ICBN) and the International Code covers those organisms variously recognized as prokaryotes,
of Nomenclature of Bacteria (ICNB) has fundamental Bacteria, Archaea, Eubacteria and Archaebacteria. Due
implications for the International Code of Nomenclature consideration has been given to including cyanobacteria,
of Prokaryotes (ICNP) put forward in 1999 (Labeda, 2000; which are traditionally covered by the International Code of
De Vos & Trüper, 2000). Indeed, Oren & Garrity (2014) Botanical Nomenclature, and has been discussed elsewhere.’
sought to cut the Gordian knot: they proposed to modify Another modified wording of General Consideration 5 as
General Consideration 5 of the ICNP in such a way that the included in the ICNP (Labeda, 2000; De Vos & Trüper,
ICNP no longer covers the nomenclature of cyanobacteria. 2000) also should be noted: ‘This Code of Nomenclature of
Approval of this proposal by the International Committee Prokaryotes applies to all prokaryotes’. The supplementary
on Systematics of Prokaryotes (ICSP) would result in the note to General Consideration 5 of the ICNP states, in
cyanobacteria falling under the unilateral coverage of the particular: ‘Prokaryotes covers those organisms that are
ICN except for several species validly published in 1989– variously recognized as e.g. ... Bacteria’. It is noteworthy that
2008 according to ICNB/ICNP provisions (Oren, 2004). although the terms ‘bacteria’ and ‘prokaryotes’ were adopted
in the ICNB and later the ICNP only by making mention of
Abbreviations: ICBN, International Code of Botanical Nomenclature;
them, their biological essence has been comprehensively
ICNB, International Code of Nomenclature of Bacteria; ICNP,
International Code of Nomenclature of Prokaryotes; ICSB, International confirmed in myriads of publications since the appearance
Committee on Systematics of Bacteria; ICSP, International Committee of the global cytology concept by Stanier & Van Niel (1962)
on Systematics of Prokaryotes. and the global phylogeny concept by Woese & Fox (1977).
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It is well known that living organisms are treated, from
viewpoint of their genotype and phenotype specificity, as
evolutionary biological elements (biosystematics). For this
purpose, classification units are delimited (taxonomy) and
these are named according to codified rules (nomenclature).
A preferred delimitation of taxa is based on phylogeny
reconstruction (Wayne et al., 1987; Klenk & Göker, 2010).
Consequently, the nomenclature, especially of higher taxo-
nomic ranks, should be governed by separate codes (Blackwell,
2008; Komárek, 2011). Specifically, the phylogeny-based
systematics of bacteria is adopted in the second edition of
Bergey’s Manual of Systematic Bacteriology (2001–2012; vols.
1–5) and the bacterial nomenclature is covered by the ICNP.
Therefore, in correspondence with the above-mentioned
wordings of General Consideration 5, cyanobacteria should
be treated under the ICNP (Imhoff & Madigan, 2004).
Since the 1870s, cyanobacteria have been interpreted as
binary dividing plants of the class Schizophyceae (the class
name coined by F. Cohn in 1875) or classified as blue-green
algae (Cyanophyta/Cyanophyceae); consequently, their nomen-
clature was initially regulated by the ICBN. Subsequently,
owing to the first recognition of the bacterial nature
of Schizophyceae/Cyanophyta/Cyanophyceae (Stanier & Van
Niel, 1941), their nomenclature – as that of cyanobacteria –
was proposed to be governed by the ICNB rules (Stanier
et al., 1978). Guided by this proposal, the Judicial Commis-
sion of the International Committee on Systematics of Bacteria
(ICSB) proposed usage of a permissible dual nomenclature
system (the ICBN/ICNB) for Cyanophyta/Cyanophyceae/
cyanobacteria. However, this proposal was never formally
endorsed by the ICSB/ICSP. The main obstacle is the
mutually incompatible provisions of the Botanical and
Bacteriological Codes; in particular, the rules concerning
valid publication of names differ. Details are omitted here
since they are comprehensively discussed elsewhere (Oren,
2004; Oren & Tindall, 2005; Komárek, 2011; Oren, 2011a, b).
The choice of a nomenclatural code to govern cyanobac-
teria has three possible options excluding the proposed
BioCode and CyanoCode (Oren, 2004; Blackwell, 2008;
Komárek, 2011). Namely, these bacteria may be regulated
(i) under the ICN; (ii) under the ICNP; or (iii) under both
the ICN and the ICNP. In the first case, according to the
proposal by Oren & Garrity (2014), cyanobacteria are annexed
by botany. Thus, organisms belonging to one phylogeny-based
system will be described under the nomenclatural principles
adopted for a separate phylogeny-based system. Moreover, the
bacterial system is deprived of its most significant element
taking into account the global role of cyanobacteria in cell
evolution, as well as their diversity, abundance, distribution
and ecology. To prevent the inconsistency, and in contrast to
the proposal by Oren & Garrity (2014), it is proposed to apply
the ICNP to cyanobacterial names including those validly
published under the ICBN.
Operationally, cyanobacteria have already begun to be
described under the ICNB/ICNP. Since 1989, there have
been precedents of valid publication of novel genera and
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Proposal to apply ICNP to the cyanobacteria
species (Oren, 2004; 2011a, b). At the same time, nomen-
clatural obstacles remained, especially for taxa validly
published under the ICBN. Thus, the wording of General
Consideration 5 of the ICNB (Lapage et al., 1992) states: ‘...
The nomenclature of certain other microbial groups is
provided for by other Codes: fungi and algae by the Botanical
Code ...’. In addition, the wording of Principle 2 (ibid.) is:
‘The nomenclature of bacteria is independent of botanical
nomenclature, except for algae and fungi ...’ (a supplement-
ary note provides the following definition: ‘... ‘‘independent’’
means that the same name may be validly used for a taxon
of bacteria as well as a taxon of plants and animals with
the exceptions noted above’). The wording of Rule 10a,
Recommendation 10a (ibid.) is: ‘The following Recommen-
dations apply when forming new generic ... names. 3. Avoid
introducing into bacteriology as generic names such names
as are in use in botany or zoology, in particular well-known
names.’ Finally, the wording of Rule 51b (ibid.) is: ‘Among
the reasons for which a name may be illegitimate ... are the
following. ... 4. If it is a junior homonym (the author’s
remark’: later’ homonym, in newer usage) of a name of a taxon
of ... algae ...’. All these provisions unequivocally implied that
cyanobacteria were considered to be algae, and thus their
exclusion from the ICNB seemed consistent.
However, the proposed wording of General Consideration
5 of the ICNP (De Vos & Trüper, 2000) states: ‘... The
nomenclature of eukaryotic microbial groups is provided
for by other codes: ... algae by the International Code of
Botanical Nomenclature ...’. The wording of the proposed
change to Principle 2 (ibid.) is: ‘The nomenclature of
Prokaryotes is not independent of botanical and zoological
nomenclature. When naming new taxa in the rank of genus
..., due consideration is to be given to avoiding names which
are regulated by ... the International Code of Botanical
Nomenclature.’ In this case, exclusion of cyanobacteria from
the ICNP is counter-logical because they are not eukaryotic
microbes and, correspondingly, algae.
Taking into account the above considerations and code
formulae, it is proposed to change the wording of Principle
2 of the ICNP as follows:
The nomenclature of prokaryotes is not independent of
zoological nomenclature, as well as nomenclature of algae,
fungi and plants. The only exception applies to names of the
oxygenic photosynthetic bacteria (cyanobacteria) validly
published, as names of algae, under the International Code
of Botanical Nomenclature/International Code of Nomen-
clature for algae, fungi and plants.
Supplementary note: ‘independent’ means that the same
name may be validly used for a taxon of bacteria as well as
a taxon of algae, fungi, plants and animals.
The proposed change of the wording of Principle 2 of the
ICNP tries to break, by way of exception, a deadlock in
cyanobacterial nomenclature. This makes an opportunity
to validate under the ICNP the names of cyanobacteria
validly published under the ICBN/ICN.
1071
A. V. Pinevich
The problem of genera names is in contexture with the
problem of combinations at the rank of species or sub-
species. Noteworthy, in the classification of cyanobacteria
elaborated by Rippka et al. (1979) and, with certain modi-
fications adopted in Bergey’s Manual of Systematic Bacte-
riology (Castenholz, 2001), the usage of species epithets is
avoided. Only four binary combinations were validated
under the ICNP: Prochlorothrix hollandica, Halospirulina
tapeticola, Planktotricoides raciborskii and Rubidibacter
lacunae (Oren, 2004; 2011a, b).
‘Operational’ (taxonomic and nomenclatural) speciation of
cyanobacteria is clearly distinct from the problem of
defining their species in accordance within the general
species definition for prokaryotes (Staley, 2006). Impor-
tantly, a genotype-level interspecies threshold has got no
consensus of opinion (Stackebrandt et al., 2002; Doolittle
& Zhaxybayeva, 2009; Tindall et al., 2010; Kim et al., 2014).
Nevertheless and similarly with other prokaryotes, the
systematics of cyanobacteria are currently based on the
polyphasic approach (Colwell, 1970; Tindall et al., 2010)
although the metabolic uniformity of this group confines
phenotypic characteristics to gross morphology and, in rare
cases, ultrastructure. With this approach, a number of
cultured strains could be reliably identified at the species
level (Otsuka et al., 2001; Wright et al., 2001; Komárek,
2006; Palinska & Marquardt, 2008).
Proposal of an ‘operational’ speciation algorithm for
cyanobacteria is beyond the scope of this taxonomic note.
However, one can outline it stepwise: (i) the ‘form-genus’
name is chosen based on matching morphological descrip-
tions of cyanobacteria in hand to those in the ‘Geitlerian’
system (Castenholz, 1992); (ii) under the ‘form-genus’
name, collection strains and database references are
operationally treated (Castenholz, 2001); (iii) the type
strain, as the best characterized one, especially in respect of
phylogeny, is considered an operational ‘form-species’; (iv)
a closely fitting match is scanned for among ‘good’, i.e.
distinct and coherent botanical species, and the opera-
tional ‘form-species’ assimilates the corresponding epithet.
Noteworthy, the samples used in former times for botanical
diagnoses should not be necessarily related to morphologi-
cally matching contemporary strains. In limited cases, this
may be verified by employing molecular biological methods
to historical herbaria (Palinska & Surosz, 2014). Anyway,
given the proposed assimilation of genera names from the
ICBN/ICN, the ICNP formula for validation of combina-
tions including species and subspecies should be sought
later. At the same time, a more radical algorithm is
admissible: species epithets of cyanobacteria are abandoned,
and strain numbers replace the corresponding portion of the
binominal (Castenholz, 1992). However, voluminous devi-
ation from the nomenclature of Linnaeus is hardly
productive.
There are important matters pertaining to cyanobacteria
that have already been pursued and should be gradually
resolved in the future: (i) development of collections of
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pure cultures; (ii) choice of type strains; (iii) removal of
misidentifications (Tindall et al., 2010; Komárek, 2011;
Gupta et al., 2013). These tasks are currently supported, in
informational respect, by the list of bacterial names with
standing in nomenclature (Euzéby, 1997; Parte, 2014).
However, the forthcoming goal is the creation of the
‘Approved Lists of Cyanobacterial Names’ based on valid
publications in the IJSEM according to the ICNB/ICNP
rules. Among the main impediments, besides the current
veto on names with standing in the ICN, is a general
shortage of pure cultures deposited in (at least two)
authoritative collections. The above nomenclatural actions
are endangered with emendation obstacles (Tindall, 1999).
Complex authorship and naming trajectories is exemplified
by Anabaina Bory (de Saint-Vincent) 1822/Anabaena
Bornet et Flahault 1886 (Blackwell, 2008). Duplicate taxa
are also undesirable, as well as proposals of new cyano-
bacterial names (Roldán et al., 2013) published in IJSEM
under the provisions of the ICN. However, as pointed out
by Imhoff & Madigan (2004): ‘There was no doubt and
general agreement that cyanobacteria have to be treated
according to the Bacteriological Code, but that this treat-
ment should, in addition, be in accordance to the Botanical
Code, whenever possible.’
In conclusion, this proposal calls for the nomenclatural
treatment of cyanobacteria along with other bacteria. Alter-
natively, they are sentenced to be tendentiously separate
from other representatives of their group. In other words,
either the ICNP rules remain fast, or the bacterial system is
violated and a long-lasting lacuna in bacterial nomenclat-
ure settled. Unilateral coverage of cyanobacteria by the ICN
would also negatively influence their research, especially
taking into account that this Code has no demand for live
collections, pure cultures and type species.
It should be reminded that bacteria had once been placed in
the plant division Schizophyta (1875; F. Cohn) along with
the fungi of the class Schizomycetes (1857; C. Nägeli), and
cyanobacteria in particular treated as the Cyanophyta/
Cyanophyceae algae. At the same time former ‘ray fungi’
(type genus, Actinomyces Harz 1877) now encompass a wide
range of bacteria belonging to the phylum Actinobacteria
(Ludwig et al., 2012), and there has been no call for
unilateral coverage of these prokaryotes by the ICN.
A résumé of the author’s arguments is as follows. Firstly
and most importantly, a proposal to consistently apply the
ICNP to cyanobacteria conforms formally and logically to
the wording and supplementary note of General Consider-
ation 5. Secondly, it is proposed to legitimize, by way of
exception, the usage of Cyanophyta/Cyanophyceae names
in order to enable descriptions of cyanobacteria under
the ICNP. With valid publications accumulated, a list of
‘Approved Names of Cyanobacteria’ would be compiled.
Thirdly, code restrictions impede the four-decades-long
operational usage of Cyanophyceae/Cyanophyta names by
the global community of (cyano)bacteriologists. Habitual
presence of ‘botanical’ names in literature on cyanobacteria
International Journal of Systematic and Evolutionary Microbiology 65

and databases is hardly possible to eradicate. In fact, an
alternative option to reconsider cyanobacterial genera
under the names coined de novo would have grave or even
chaotic informational consequences.
Acknowledgements
I am grateful to the anonymous reviewer for criticism and offering
many valuable suggestions that improved the manuscript consistency.
References
Blackwell, W. H. (2008). ‘‘Anabaena’’, ‘‘Anabaina’’, and codes of
nomenclature: a review of the feasibility of name correction, and a
possible direction for the future. Phytologia 90, 324–354.
Castenholz, R. W. (1992). Species usage, concept, and evolution in
the cyanobacteria (blue-green algae). J Phycol 28, 737–745.
Castenholz, R. W. (2001). General characteristics of the cyanobac-
teria. In Bergey’s Manual of Systematic Bacteriology, 2nd edn, vol. 1,
pp. 474–478. Edited by D. R. Boone, R. W. Castenholz & G. M.
Garrity. New York: Springer.
Colwell, R. R. (1970). Polyphasic taxonomy of the genus vibrio:
numerical taxonomy of Vibrio cholerae, Vibrio parahaemolyticus, and
related Vibrio species. J Bacteriol 104, 410–433.
De Vos, P. & Trüper, H. G. (2000). Judicial Commission of
the International Committee on Systematic Bacteriology. IXth
International (IUMS) Congress of Bacteriology and Applied
Microbiology. Minutes of the meetings, 14, 15 and 18 August 1999,
Sydney, Australia. Int J Syst Evol Microbiol 50, 2239–2244.
Doolittle, W. F. & Zhaxybayeva, O. (2009). On the origin of
prokaryotic species. Genome Res 19, 744–756.
Euzéby, J. P. (1997). List of bacterial names with standing in
nomenclature: a folder available on the internet. Int J Syst Bacteriol 47,
590–592.
Gupta, V., Ratha, S. K., Sood, A., Chaudhary, V. & Prasanna, R.
(2013). New insights into the biodiversity and applications of
cyanobacteria (blue-green algae) – prospects and challenges. Algal
Research 2, 79–97.
Imhoff, J. F. & Madigan, M. T. (2004). International Committee on
Systematics of Prokaryotes. Subcommittee on the taxonomy of
phototrophic bacteria. Minutes of the meetings, 27 August 2003,
Tokyo, Japan. Int J Syst Evol Microbiol 54, 1001–1003.
Kim, M., Oh, H.-S., Park, S.-C. & Chun, J. (2014). Towards a
taxonomic coherence between average nucleotide identity and 16S
rRNA gene sequence similarity for species demarcation of prokar-
yotes. Int J Syst Evol Microbiol 64, 346–351.
Klenk, H.-P. & Göker, M. (2010). En route to a genome-based
classification of Archaea and Bacteria? Syst Appl Microbiol 33, 175–182.
Komárek, J. (2006). Cyanobacterial taxonomy: current problems and
prospects for the integration of traditional and molecular approaches.
Algae 21, 349–375.
Komárek, J. (2011). Introduction to the 18th IAC Symposium in
České Budějovice 2010, Czech Republic. Some current problems of
modern cyanobacterial taxonomy. Fottea 11, 1–7.
Labeda, D. P. (2000). International Committee on Systematic
Bacteriology. IXth International (IUMS) Congress of Bacteriology
and Applied Microbiology. Minutes of the meetings, 14 and 17 August
1999, Sydney, Australia. Int J Syst Evol Microbiol 50, 2245–2247.
Lapage, S. P., Sneath, P. H. A., Lessel, E. F., Skerman, V. B. D.,
Seeliger, H. P. R. & Clark, W. A. (editors) (1992). International Code of
Downloaded from www.microbiologyresearch.org by
http://ijs.sgmjournals.org
IP: 112.215.175.96
On: Fri, 05 Oct 2018 15:28:48
Proposal to apply ICNP to the cyanobacteria
Nomenclature of Bacteria (1990 Revision). Bacteriological Code.
Washington, DC: American Society for Microbiology.
Ludwig, W., Euzéby, J., Schumann, P., Busse, H.-J., Trujillo, M. E.,
Kämpfer, P. & Whitman, B. W. (2012). Road map of the phylum
Actinobacteria. In Bergey’s Manual of Systematic Bacteriology, 2nd edn,
vol 5, part A, pp. 1–28. Edited by M Goodfellow, P Kämpfer, H.-J.
Busse, M. E. Trujillo, K. Suzuki, W. Ludwig & W. B. Whitman. New
York: Springer.
Oren, A. (2004). A proposal for further integration of the
cyanobacteria under the Bacteriological Code. Int J Syst Evol
Microbiol 54, 1895–1902.
Oren, A. (2011a). Cyanobacterial systematics and nomenclature as
featured in the International Bulletin of Bacteriological Nomenclature
and Taxonomy/International Journal of Systematic Bacteriology/
International Journal of Systematic and Evolutionary Microbiology.
Int J Syst Evol Microbiol 61, 10–15.
Oren, A. (2011b). Naming Cyanophyta/Cyanobacteria – a bacteriol-
ogist’s view. Fottea 11, 9–16.
Oren, A. & Garrity, G. M. (2014). Proposal to change General
Consideration 5 and Principle 2 of the International Code of
Nomenclature of Prokaryotes. Int J Syst Evol Microbiol 64, 309–310.
Oren, A. & Tindall, B. J. (2005). Nomenclature of the cyanophyta/
cyanobacteria/cyanoprokaryotes under the International Code of
Nomenclature of Prokaryotes. Algol Stud 117, 39–52.
Otsuka, S., Suda, S., Shibata, S., Oyaizu, H., Matsumoto, S. &
Watanabe, M. M. (2001). A proposal for the unification of five species
of the cyanobacterial genus Microcystis Kützing ex Lemmermann 1907
under the rules of the Bacteriological Code. Int J Syst Evol Microbiol
51, 873–879.
Palinska, K. A. & Marquardt, J. (2008). Genotypic and phenotypic
analysis of strains assigned to the widespread cyanobacterial
morphospecies Phormidium autumnale (Oscillatoriales). Arch
Microbiol 189, 325–335.
Palinska, K. A. & Surosz, W. (2014). Taxonomy of cyanobacteria: a
contribution to consensus approach. Hydrobiologia 740, 1–11.
Parte, A. C. (2014). LPSN–list of prokaryotic names with standing in
nomenclature. Nucleic Acids Res 42 (D1), D613–D616.
Rippka, R., Deruelles, J., Waterbury, J. B., Herdman, M. & Stanier,
R. Y. (1979). Generic assignments, strain histories and properties of
pure cultures of cyanobacteria. J Gen Microbiol 111, 1–61.
Roldán, M., Ramı́rez, M., del Campo, J., Hernández-Mariné, M. &
Komárek, J. (2013). Chalicogloea cavernicola gen. nov., sp. nov.
(Chroococcales, Cyanobacteria), from low-light aerophytic environ-
ments: combined molecular, phenotypic and ecological criteria. Int J
Syst Evol Microbiol 63, 2326–2333.
Stackebrandt, E., Frederiksen, W., Garrity, G. M., Grimont, P. A.,
Kämpfer, P., Maiden, M. C., Nesme, X., Rosselló-Mora, R., Swings, J.
& other authors (2002). Report of the ad hoc committee for the re-
evaluation of the species definition in bacteriology. Int J Syst Evol
Microbiol 52, 1043–1047.
Staley, J. T. (2006). The bacterial species dilemma and the genomic-
phylogenetic species concept. Philos Trans R Soc Lond B Biol Sci 361,
1899–1909.
Stanier, R. Y. & Van Niel, C. B. (1941). The main outlines of bacterial
classification. J Bacteriol 42, 437–466.
Stanier, R. Y. & Van Niel, C. B. (1962). The concept of a bacterium.
Arch Mikrobiol 42, 17–35.
Stanier, R. Y., Sistrom, W. R., Hansen, T. A., Whitton, B. A.,
Castenholz, R. W., Pfennig, N., Gorlenko, V. N., Kondratieva, E. N.,
Eimhjellen, K. E. & other authors (1978). Proposal to place the
nomenclature of cyanobacteria (blue-green algae) under the rules of
1073
A. V. Pinevich

the International Code of Nomenclature of Bacteria. Int J Syst
Bacteriol 28, 335–336.
Tindall, B. J. (1999). Proposals to update and make changes to the
Bacteriological Code. Int J Syst Bacteriol 49, 1309–1312.
Tindall, B. J., Rosselló-Móra, R., Busse, H.-J., Ludwig, W. & Kämpfer,
P. (2010). Notes on the characterization of prokaryote strains for
taxonomic purposes. Int J Syst Evol Microbiol 60, 249–266.
Wayne, L. G., Brenner, D. J., Colwell, R. R., Grimont, P. A. D., Kandler,
O., Krichevsky, M. I., Moore, L. H., Moore, W. E. C., Murray, R. G. E. &
other authors (1987). International Committee on Systematic
Bacteriology. Report of the ad hoc committee on reconciliation of
approaches to bacterial systematics. Int J Syst Bacteriol 37, 463–464.
Woese, C. R. & Fox, G. E. (1977). Phylogenetic structure of the
prokaryotic domain: the primary kingdoms. Proc Natl Acad Sci U S A
74, 5088–5090.
Wright, D., Prickett, T., Helm, R. F. & Potts, M. (2001). Form species
Nostoc commune (Cyanobacteria). Int J Syst Evol Microbiol 51, 1839–
1852.

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