Prokaryote Nomenclature

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Prokaryote NomenclatureAharon Oren, The Hebrew University of Jerusalem, Jerusalem, Israel

Nomenclature of prokaryotes is regulated by the provisions of the International Code

of Nomenclature of Prokaryotes. To obtain standing in the nomenclature, names ofnew taxa must be validly published according to the rules of that Code.

Introduction

As of 4 October 2007, the names of 7255 different species ofprokaryotes have been validly published in accordance

with the internationally accepted rules of nomenclature, as

fixed in the International Code of Nomenclature of Pro-karyotes (ICNP, The Bacteriological Code) (Lapageet al., 1992).

Nomenclature is the assignment of names to the taxo-nomic groups defined during classification, and this articleexplores the question how species, genera, families andhigher taxa within the prokaryote world (Bacteria andArchaea combined) are named. Issues of prokaryote no-menclature are tightly linked to issues of their systematics/

taxonomy and classification, but these terms do not haveidentical meanings. Systematics is the study of organismswith the ultimate object of characterizing and arrangingthemin an orderlymanner. The term taxonomy, often used

synonymously with systematics can be defined as the the-ory and practice of classifying organisms into groups (taxa)on the basis of similarities and relationships. Systematicsincludes the study of the evolutionary and phylogenetic re-lationships of the organisms. Such studies lead to the es-tablishment of schemes of classification, i.e. the orderlyarrangement of units into groups. Nomenclature is thenused to properly name the taxonomic units thus established,from species and subspecies to classes and kingdoms.

The basic unit of classification of all living organisms isthe species. Although over 7000 different species of pro-karyotes have been described and named according to theestablished rules discussed later, there still is no universallyrecognized definition of what a prokaryote species is. To thebotanist and the zoologist the definition of the conceptspecies presents relatively few problems. In the plant andthe animal world a species is generally defined as a popu-lation of individuals that can interbreed under natural

conditions, produce fertile offspring and that is reproduc-tively isolated from other populations. Such a definition isuseless in the case of the prokaryotes that show no sexualreproduction. One may circumscribe the species as a dis-tinct group of strains that have certain distinguishing fea-tures and that generally bear a close resemblance to oneanother in the more essential features of organization, oran assemblage of clonal populations that share a high deg-ree of phenotypic similarity, coupled with an appreciabledissimilarity from other assemblages of the same generalkind. Such definitions provide little practical informationon how close that resemblance and similarity should be fortwo strains to be classified in thesamespecies, what featuresof organization should be considered essential, and whatdegree of dissimilarity is required for two strains to warrantclassification in different species. Seealso: Species Concepts

Based on the experience of the last 20 years, a pragmaticdefinition of the prokaryote species has emerged, based on

the recommendations published in 1987 by a committee ofexperts, and updated a few years ago (Stackebrandt et al.,2002). The species concept is based on a polyphasicapproach, which includes description of diagnostic phe-notypic features combined with genomic properties. Indi-vidually many of the phenotypic and chemotaxonomiccharacteristics used as diagnostic properties are insufficientto delineate species, but together they provide sufficientdescriptive information. The most widely used genomicproperty included in descriptions of new species is the se-quence of the small subunit ribosomal ribonucleic acid(RNA) gene, and many classification schemes are based onthis gene. However, species delineation should not be based

on the sequence of a single gene only. A widely acceptedcriterion, based on the complete genome, defines a pro-karyotic species as a group of strains, including the typestrain (see later), that share at least 70% total genomeDNADNA hybridization and have less than 58C DT

m

(=the difference in the melting temperature between thehomologous and the heterologous hybrids formed understandard conditions). The delineation value of 70% is arti-ficial, but has proven satisfactory in most cases. Deoxyri-bonucleic acid (DNA) relatedness values between 30% and70% indicate a moderate degree of relationship, oftenparallel to the extent of the genus. There are cases in theliterature of species and even genera that share more than70% DNADNA similarity. Thus, the genera of enteric

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Article Contents

. Introduction

. Prokaryote Nomenclature: A Historical Overview

.

The International Code of Nomenclature of Prokaryotes. How Prokaryote Names are Formed

. Effective and Valid Publication of Names of Prokaryotes

. The Special Case of Cyanobacteria Nomenclature

Online posting date: 15th July 2008

ELS subject area: Microbiology

How to cite:

Oren, Aharon (July 2008) Prokaryote Nomenclature. In: Encyclopedia ofLife Sciences (ELS). John Wiley & Sons, Ltd: Chichester.

DOI: 10.1002/9780470015902.a0021150

ENCYCLOPEDIA OF LIFE SCIENCES & 2008, John Wiley & Sons, Ltd. www.els.net 1
http://dx.doi.org/10.1038/npg.els.0001744http://dx.doi.org/10.1038/npg.els.0001744


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bacteria Escherichia and Shigella share more than 85%similarity. However, for pragmatic reasons the separationinto two species is maintained. In spite of the great ad-vances made in recent years in genomic analysis and thedevelopment of other modern approaches to characterizeprokaryote isolates, DNADNA hybridization combined

with a thorough phenotypic characterization is still the bestcriterion for species delineation (Stackebrandt et al., 2002).See also: Phylogeny Based on 16S rRNA/DNA

Description of a species should ideally be based on acomparative study of a large number of isolates to definethe degree of variation of certain properties within theboundaries of the species. In practice, however, more thanhalf of the new species descriptions published in recentyears were based on the study of single isolates.

Prokaryote Nomenclature: A Historical

Overview

When Antoni van Leeuwenhoek firstsaw prokaryotes in thelast decades of the seventeenth century, he did not name thetiny organisms that he observed in his primitive mi-croscopes. The first attempts towards a nomenclature ofthe prokaryotes probably stem from the work of ChristianEhrenberg (17951877) in the third decade of the nineteenthcentury. Ehrenberg divided the prokaryotes known tohim on the basis of morphological criteria into fourgenera: Bacterium (straight rigid filaments; three species),Vibrio (twisted filaments; nine species), Spirochaeta (non-

rigid spiral filaments; one species) and Spirillum (rigid spi-ral filaments; three species). A more elaborate classificationof the bacteria proposed in 1875 by Ferdinand Cohn(18281898) which recognized six genera, classified in fourtribes: the tribe Sphaerobacteria with the genus Micrococ-cus, the tribe Microbacteria with the genus Bacterium, thetribe Desmobacteria with generaBacillus and Vibrio and thetribe Spirobacteria with genera Spirillum and Spirochaeta.See also: Cohn, Ferdinand Julius; Leeuwenhoek, Antonivan; Systematics: Historical Overview

In the first half of the twentieth century techniques forcultivation of different types of prokaryotes, aerobes as wellas anaerobes, were developed, and many new prokaryotes

were isolated, described and named. In an attempt to keeptrack of the ever-increasing number of published names ofprokaryotes, a handbook was established named BergeysManual of Determinative Bacteriology, published in eighteditions between 1923 and 1974. It rapidly became clear thatdifferent names had often been given to identical or verysimilar bacteria.Synonyms abounded, and the nomenclaturebecame ever more confusing. The publication of the IndexBergeyanaAn Annotated AlphabeticListing of Names of theTaxa of the Bacteria (Buchanan et al., 1966), which con-tained approximately 28 900 entries, shows the urgent needfelt at the time to establish order in the increasing chaos.

An entirely new start was made in prokaryote nomen-clature in 1980 with the publication of the Approved listsof

bacterial names (Skerman et al., 1980). Rule 24a of theBacteriological Code (see later) states: Priority of publi-cation dates from 1 January 1980. On that date all namespublished prior to 1 January 1980 and included in the Ap-proved Lists of Bacterial Names of the ICSB [the Interna-tional Committee on Systematics of Bacteria] are treatedfor

all nomenclatural purposes as though they had been validlypublished for the first time on that date. With the publi-

cation of the Approved Lists, which contained approxi-mately 2500 speciesnames,all other earlierpublishednameslost their validity. Central registration and indexing of newnames validly published after 1 January 1980, according tothe rules of the Bacteriological Code (Lapage et al., 1992;see later), ensured that the number of species, genera, fam-ilies and orders of prokaryotes whose names have been val-idly published is known at anytime (Table1). For each taxona nomenclatural type has to be designated; the type is thatelement of the taxon with which the name is permanentlyassociated. All relevant information is published in theInternational Journal of Systematic and Evolutionary Micro-biology (IJSEM) (until 1999: the International Journal of

Systematic Bacteriology, IJSB). The information is alsoavailable online at http://www.bacterio.cict.fr, a web siteestablished and maintained by Dr. Jean Euze by of theUniversity of Toulouse, France (Euze by, 1997). This site isupdated monthly with the publication of the latest issue ofIJSEM. This makes it easy for the bacteriologist to check

whether a certain organism has been described and namedin accordance to the internationally agreed criteria. In con-trast, a botanist who wants to establish whether a certainplant has been recorded before in theliterature has to search

the scientific journals andbooksas far back as1753, the yearin which Linnaeus published his Species Plantarum, andthere exists no central registration of plant names validlypublished under the International Code of BotanicalNomenclature (ICBN, the Botanical Code). Central reg-istration of prokaryote names has also enabled the estab-

lishment of a quality control mechanism to ensure thatnames of taxa newly added to the list would be properlyformed. See also: Codes of Nomenclature

Table 1 The number of different prokaryote taxa described

with names with standing in the nomenclature, as of 4 October2007. Derived from http://www.bacterio.cict.fr

Number of phylaa 28

Number of classesb 67

Number of orders 105

Number of families 233

Number of genera 1616c

Number of species 7255d

aNomenclature of phyla is not covered by the Bacteriological Code.bNomenclature of classes is not covered by the Bacteriological Code.cOf which approximately 85 are considered as synonyms.dThisnumberrepresents thetotal number of names of differentspecies

validly published: 8441 validly published names, minus 1108 newcombinations,minus 11 nominanova, minus 67 homotypicsynonyms.

Prokaryote Nomenclature

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The International Code ofNomenclature of Prokaryotes

Nomenclature of the prokaryotes is regulated by the rules ofthe Bacteriological Code. The first edition was published

in 1948 as The International Code of Nomenclature ofBacteria and Viruses. The last published version of theInternational Code of Nomenclature of Bacteria, as ap-proved at the Ninth International Congress of Microbio-logy, Moscow, 1966 and revised in 1990 (Lapage et al.,1992) has since been amended at subsequent meetings of theInternational Committee on Systematics of Prokaryotes(ICSP, see later). A new revised version of the Code is cur-rently in preparation to be named International Code ofNomenclature of Prokaryotes. The Bacteriological Codepresents the formal framework according to which pro-karyotes are named and according to which existing names

can be changed or rejected. It covers therules forthe namingof species and subspecies, genera, families and ordersof prokaryotes. No provisions are made by the Code forthe naming of the higher taxa: classes, phyla and kingdoms.See also: Codes of Nomenclature

The rules that regulate the nomenclature of prokaryotes,as published in the Bacteriological Code, are set by theICSP (before 2000: the ICSB). This committee, a constit-uent part of the International Union of MicrobiologicalSocieties, discusses nomenclatural problems that havearisen in different groups of prokaryotes and proposeschanges and amendmentsto the rules of the BacteriologicalCode. The Judicial Commission of the ICSP deals with

problematic cases in prokaryote nomenclature and rendersjudicial decisions in instances of controversy about thevalidity of a name, identity of type strains and cases ofemerging problems with the interpretation of the rules of

the Bacteriological Code. It also may propose amend-ments to the Code and consider exceptions that may beneeded to certain rules. The ICSP has also establishedtaxonomic subcommittees (currently 28) that discussnomenclatural problems of specific groups of prokaryotes.The minutes of the meetings of the ICSP, its Judicial Com-mission and its taxonomic subcommittees are published inIJSEM. Informationon theICSP, itscurrent officers and itssubcommittees can be found at: http://www.the-icsp.org.

It must be stressed that prokaryote nomenclature is thusgoverned by internationally approved rules and regula-tions, but that there is no official classification of pro-karyotes. The classification provided in Bergeys Manual ofSystematic Bacteriology is widely used among microbiolo-gists, but was never intended to obtain official status. Gen-eral Consideration 4 of the Bacteriological Code clearlystates: Rules of nomenclature do not govern the delimita-tion of taxa nor determine their relations. The rules areprimarily for assessing the correctness of the names appliedto defined taxa; they also prescribe the procedures for cre-ating and proposing new names. The Code deals with allprokaryotes: the nomenclature of the Archaea and the Bac-teria are governed by the same rules. See also: Archaea

How Prokaryote Names are Formed

Microbiologists who have occasion to use the scientificnames of the microorganisms with which they deal gener-ally prefer to use correct names and to use them correctly.Thus opens the foreword to the first edition (1948) of the

Bacteriological Code. The Code contains many rules andrecommendations to guide authors describing new species,genera, families, etc. of prokaryotes and how to properlyname the new taxa. Similar to the eukaryotic organisms thespecies of prokaryotes have generic names and specific ep-ithets derived from Latin, latinized Greek or latinized mod-ern words or names. Principle 3 of the Code states: Thescientific names of all taxa are Latin words treated as Latinregardless of their origin. They are usually taken from Latinor Greek. In the binomial system, as introduced by CarolusLinnaeus for the plant world in the 1750s, each organism isknown by a combination consisting of the name of the ge-nus followed by a single specific epithet. Another importantstatement in the Code is Principle 4: The primary purposeof giving a name to a taxon is to supply a means of referringto it rather than to indicate the characters or the history ofthe taxon. See also: Linnaeus, Carl (Linne )

The main rules for the formation of correct genus andspecies names for prokaryotes are found in Rules 10a and12c of the Bacteriological Code:

(Rule 10a): The name of a genus_ is a substantive, oran adjective used as a substantive, in the singularnumber and written with an initial capital letter. Thename may be taken from any source and may even be

composed in an arbitrary manner. It is treated as a Latinsubstantive.(Rule 12c): A specific epithet may be taken from anysource and may even be composed arbitrarily.A specific epithet must be treated in one of the three

following ways:As an adjective that must agree in gender with the ge-neric name.As a substantive (noun) in apposition in the nominativecase.As a substantive (noun) in the genitive case.

Each description of a new species should include a pro-

posal for its name and explain the etymology of that name.Practical recommendations on how to name a new pro-karyote can be found in the Orthography appendix to theBacteriological Code and in a review paper by Tru per1999. To illustrate theimplementation of Rules 10aand 12cof the Code, here are a few examples of how genus andspecies names are formed.

Some genus names are derived from a single Latin stem:Vibrio, Bacillus or from a single Greek stem: Clostridium,Thermus; others combine two or more Latin stems: Lacto-bacillus, Sulfolobus, twoor more Greek stems: Haemophilus,Halococcus, Ectothiorhodospira (the red spiral with sulfuroutside), or arebased on a combination of Latin and Greek

stems: Flavobacterium, Halorubrum, Halolactibacillus, etc.


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Quite a few genus names are latinized personal names:Escherichia, Shigella, Pfennigia, Pasteuria, etc. named tohonour the contributions of Theodor Escherich, KiyoshiShiga, Norbert Pfennig and Louis Pasteur to the science ofmicrobiology. Not always are such names derived from thefamily name of the investigator: the genus name Erwinia

honours Erwin F Smith (18541929), and Simkania wasformed from the personal name of Simona Kahane. It is

recommended to refrain from naming genera after personsunconnected with bacteriology or at least natural sciences(recommendation 10a(1) of the Bacteriological Code). Anexample of a genus name formed in an arbitrary manner isDesemzia, based on the initials of the DSMZ, the DeutscheSammlung von Mikroorganismen und Zellkulturen, theGerman culture collection of microorganisms. Another in-teresting arbitrary name is the cyanobacterial genus Nostoc,named in the fifteenth century by Paracelsus based on theold English Nosthryl (nostril) and the German equivalent

Nasenloch (about special problems connected with namingcyanobacteria, see later).

In the past, the nomenclature of prokaryotes was inde-pendent of botanical and zoological nomenclature. As aresult, there are a number of names of genera that havehomonyms in eukaryote world. Thus, Bacillus is not onlythe name of a bacterium (Firmicutes) but also of an insect,and Proteus is not only a memberof theEnterobacteriaceaebut also an amphibian inhabiting caves. The genus namesBacillus and Proteus appeared on the Approved Lists of1980, and therefore have standing in the prokaryote no-menclature. With the modification of Principle 2 of theBacteriological Code in 1999, nomenclature of pro-

karyotes is no longer independent of the nomenclature ofother groups of organisms. Therefore, when in 2005 it wasproposed to name a new genus in honour of Antoni vanLeeuwenhoek, the discoverer of the prokaryote world, thename Leeuwenhoekia could not be used as it was alreadyused for a mite; the name Leeuwenhoekiella was chosen

instead. See also: Leeuwenhoek, Antoni vanMany specific epithets are formed from Latin or latinized

adjectives: albus (white), aureus (golden), subtilis (slender),halophilus (salt-loving), etc. (masculine forms; in accord-ance to Rule 12(c)1 the gender should agree with thegender of the generic name). Participles are also treated asadjectives: ferrooxidans (iron-oxidizing), fermentans (fer-

menting), natans (floating), etc. In most cases, these adjec-tives reflect some evident phenotypic property of thespecies. Another category of specific epithets based on

Latin or latinized adjectives is the geographical names re-flecting the location where the species was first isolated.These adjectives generally carry the suffix -ensis (masculine,feminine) or -ense (neutre): Halorubrum sodomense for thebiblical city of Sodom, Haloferax lucentense for Alicante,Spain, named Lucentum in Roman times and Thermo-anaerobacterium aotearoense (one of the longest names everdevised for a prokaryote!) named after Aotearoa the landof the long white cloud, being the Maori name for NewZealand. In recent years, the number of such geographical

names has strongly increased, and in some cases the

creation of such names has been misused for geopoliti-cal or other purposes not related to the science of micro-biology. It can only be hoped that the recent call to reducethe creation of such geographical names and to eradi-cate the phenomenon of localimania (Tru per, 2005) willhave the desired result.

Substantives in apposition in the nominative case,sensu Rule 12(c)2 are relatively rare in prokaryote nomen-

clature. Examples are Desulfovibrio gigas (the giant) andPaenibacillus fonticola (inhabitant of a fountain). The ar-bitrary specific epithed in Bacteroides thetaiotaomicron alsobelongs to this category. Far more abundant is the use of asubstantive in the genitive case (Rule 12(c)3), examples be-ing Streptococcus suis (of the hog), Corynebacterium diph-

theriae (of diphtheria), Pediococcus acidilactici(of lacticacid) and Clostridium carnis (of flesh). Famous microbi-ologists of the past and the present can also be honouredwith similarly formed specific epithets: Thiobaca trueperi

(of Hans Tru per), Methanosphaera stadtmanae (ofTherese Stadtman), etc.

Although the Bacteriological Code aims at stability ofnames (Principle 1(1) of the Code), and the useless creationof names is to be avoided (Principle 1(3)), changes in no-menclature do occur regularly as our understanding ofprokaryote taxonomy, phylogeny, etc. advances. Rules 34and 41 of the Code describe the procedure of transferringspecies to another genus while retaining the original specificepithet as a combinatio nova (comb. nov.). Thus, the bac-terium originally described as Vibrio harveyi in 1936was renamed Beneckea harveyi in 1973, to be restored toV. harveyi in 1981. When the genus Ralstonia was estab-

lished in 1996 to accommodate two species ofBurkholderiaand one species of Alcaligenes, the species Alcaligeneseutrophus named in 1969 was renamed Ralstonia eutropha.When in view ofthe novel insights into the taxonomy of thegroup the genus Ralstonia was split in 2004, R. eutropha

became Wautersia eutropha. Similarly, Bacillus stearother-mophilus became Geobacillus stearothermophilus, and thereare many more such cases.

More information on prokaryote nomenclature can be

found in handbookssuch as Bergeys Manual of SystematicBacteriology, published since 1984 as the successor ofBergeys Manual of Determinative Bacteriology and TheProkaryotes published in three editions since 1981, in the

original articles describing new taxa in IJSEM, as well as inthe already mentioned web site http://www.bacterio.cict.fr.

Effective and Valid Publication ofNames of Prokaryotes

Since the publication of the Approved Lists in 1980(Skerman et al., 1980), IJSEM (prior to 1980: IJSB) is theonly framework in which new names of taxa can be pub-lished to obtain standing in the nomenclature of pro-

karyotes (valid publication according to Rules 2732 ofthe Code). One way to achieve valid publication of the new




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name of species, genera, etc. is by publication of thedescription of the new taxa in that journal, while meetingdifferent criteria as determined by the Code. For new spe-cies a nomenclatural type has to be designated, which is thename-bearer of that species and is the reference specimenfor the name. Whenever a new prokaryote species is

described, the authors are obliged to deposit the type strainas a viable and pure culture in at least two publicly acces-

sible culture collections located in different countries forsafekeeping, and make subcultures available to any inter-ested scientist for further study. For valid publication ofthe name, the description should fulfil further criteria asdetermined by the ICSP.

Descriptions of new taxa and proposals for new namescan also be published in other journals. This is known aseffective publication of the names (Rule 25 of the Code).Names thus published can only obtain standing in thenomenclature when they are included in the Validation

Lists (Lists of new names and new combinations previ-ously effectively, but not validly, published) that appearperiodically in IJSEM. Authors of papers in which nameshave been effectively published and who wish to validatethe names must submit copies of the original publication,together with proof that all criteria for valid publicationhave been met, such as deposition of the type strain in twoculture collections, availability of the type strain withoutrestrictions to the scientific community, etc. The request for

validation of the name will then be handled by one of theassociate editors of the journal together with the list editor(Tindall et al., 2006).

Under the current rules of the Bacteriological Code,

valid publication of a species name is possible only forprokaryotes that can be cultured and maintained in pureculture. To provide the opportunity to name speciesthat have been relatively well characterized but cannotat present be maintained in pure culture, the categoryCandidatus has been instituted, a provisional status that

can be established for a putative taxon of an incompletelydescribed prokaryote, awaiting its future cultivation andvalid description of the name (Murray and Stackebrandt,1995). As of 4 October 2007, 84 such Candidati have beendescribedin IJSEM/IJSB. The category Candidatus is notcovered by rules of the Bacteriological Code, and there-fore a name included in the category Candidatus cannot

be validly published.

The Special Case of CyanobacteriaNomenclature

The phylum Cyanobacteria presents special nomenclatureproblems, as the group is also included by the botanistsunder the rules of the Botanical Code as Cyanophyta orblue-green algae. Historically, the group has always beencovered by the provisions of the Botanical Code. In 1962,

Roger Stanier and Cornelis van Niel, in their classic paperon The concept of a bacterium wrote: The distinctive

property of bacteria and bluegreen algae is the prokaryo-tic nature of their cells. Soon after, bacteriologistsproposed to include the group under the rules of the Bac-teriological Code. However, the two Codes are very differ-ent and are incompatible in a number of essential aspects:

1. One of the key provisions under the BacteriologicalCode is central registration/indexing of names validlypublished, centralized in IJSEM. Under the BotanicalCode few restrictions exist on the journal in which newnames may be validly published, and as a result, theexisting botanical nomenclature information is widelyscattered. Central registration/indexing does not yetexist. In most cases priority of publication dates back to

1753, but for two specific groups of Cyanophyta laterdates are in use: 1886 and 1892.

2. Under the rules of the Bacteriological Code, thenomenclatural type of a species is a living type strain,maintained in pure culture. In contrast, type specimens

of species under the Botanical Code may not be livingplants or cultures but herbarium specimens, illustra-tions, or in some cases cultures preserved in a metabo-lically inactive state.

3. The Bacteriological Code is not independent of theBotanical Code. Therefore, it is currently impossibleto describe under the Bacteriological Code a new spe-cies of a genus whose name was earlier validly publishedunder the Botanical Code. Such genus names have novalid status under the rules of the BacteriologicalCode, making the new combination illegitimate.

The 1980 approved list of bacterial names (Skermanet al., 1980) did not contain any names of Cyanobacteria,and only a few genus and species names of cyanobacteriahave since been validly published under the Bacteriologi-cal Code. Attempts are underway to solve the nomencla-ture problems of this group and to find a solution thatwill satisfy bacteriologists and botanists alike (Oren andTindall, 2005). See also: Cyanobacteria

References

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the Bacteria. Baltimore: Williams and Wilkins.

Euze by J (1997) List of bacterial names with standing in nomen-

clature: a folder available on the internet. International Journal

of Systematic Bacteriology 47: 590592.

Lapage SP, Sneath PHA, Lessel EF Jr et al. (1992) International

Code of Nomenclature of Bacteria (1990 revision). Washington,

DC: American Society of Microbiology.

Murray RGE and Stackebrandt E (1995) Taxonomic note: im-

plementation of the provisional status Candidatus for incom-

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Systematic Bacteriology 45: 186187.

Oren A and Tindall BJ (2005) Nomenclature of the cyanophyta/

cyanobacteria/cyanoprokaryotes under the International Code




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of Nomenclature of Prokaryotes. Algological Studies 117:

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Skerman VBD, McGowan V and Sneath PHA (1980) Approved

lists of bacterial names. International Journal of Systematic

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Stackebrandt E, Frederiksen W, Garrity GM et al. (2002) Report

of the ad hoc committee for the re-evaluation of the speciesdefinition in bacteriology. International Journal of Systematic

and Evolutionary Microbiology 52: 10451047.

Tindall BJ, Ka mpfer P, Euze by J etal. (2006) Valid publication of

names of prokaryotes according to the rules of nomenclature:

past history and current practice. International Journal of Sys-

tematic and Evolutionary Microbiology 56: 27152720.

Tru per HG (1999) How to name a prokaryote? Etymological

considerations, proposals and practical advice in prokaryotic

nomenclature. FEMS Microbiology Reviews 23: 231249.

Tru per HG (2005) Is localimania becoming a fashion for pro-

karyote taxonomists? International Journal of Systematic and

Evolutionary Microbiology 55: 1753.

Further Reading

Brenner DJ, Staley JT and Krieg NR (2001) Classification of

procaryotic organisms and the concept of bacterial speciation.

In: Boone DR and Castenholz RW (eds) Garrity, GM (editor-

in-chief) Bergeys Manual of Systematic Bacteriology, 2nd edn,

vol. 1. The Archaea and the Deeply Branching and Phototrophic

Bacteria, pp 27 31. New York: Springer.

Dworkin M, Falkow S, Rosenberg E et al. (eds) (2006) The Pro-

karyotes. A Handbookon the Biology of Bacteria: Ecophysiology

and Biochemistry, 7 vols. New York: Springer.

Garrity GM (editor-in-chief) (5 vols; vols 1 and 2 were published

in 2001 and) (2005) Bergeys Manual of Systematic Bacteriol-ogy, 2nd edn. New York: Springer.

Oren A (2004) Prokaryote diversity and taxonomy: present status

and future challenges. Philosophical Transactions of the Royal

Society. Series B 559: 623638.

Rossello -Mora R and Amann R (2001) The species concept for

prokaryotes. FEMS Microbiology Reviews 25: 3967.

Sneath PHA (2001) Bacterial nomenclature. In: Boone DR,

Castenholz RW and Garrity GM (eds) Bergeys Manual of

Systematic Bacteriology, 2nd edn, vol. 1. The Archaea and the

Deeply Branching and Phototrophic Bacteria, pp. 8388. New

York: Springer.

Tru per HG (2001) Etymology in nomenclature of prokaryotes.

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Documents

Prokaryote Nomenclature