Proposals Concerning the Higher Taxa of Bacteria
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0020-7713/78/0028-0001$02.00/0 INTERNATIONALJO~JRNAL OF SYSTEMATIC BACTERIOLOGY,Jan. 1978, p. 1-6 Vol. 28, No. 1 Copyright 0 1978 International Association of Microbiological Societies Printed in U.S. A. Proposals Concerning the Higher Taxa of Bacteria N. E. GIBBONSt AND R. G. E. MURRAY Department of Bacteriology and Immunology, University of Western Ontario, London, Ontario, Canada N6A 5C1 Names are proposed for divisions, classes, and some orders of bacteria. The kingdom Procaryotae Murray 1968 is divided into three divisions: Gracilicutes divisio nov., Firmacutes divisio nov., and Mollicutes Edward and Freundt 1967, for organisms having, respectively, a gram-negative cell wall, a gram-positive cell wall, and no cell wall. Gracilicutes comprises the class Photobacteria classis nov. for organisms having a phototrophic metabolism and Scoto bacteria classis nov. for those having a nonphototrophic metabolism. Photobacteria is divided into two subclasses: Oxyphotobacteriae subclassis nov. for organisms that carry out an aerobic phototrophic metabolism, using water as an electron donor and producing oxygen, and Anoxyphotobacteriae subclassis nov. for those that have an anaerobic phototrophic metabolism. Oxyphotobacteriae comprises the orders Cyano bacteriales ordo nov. (formerly blue-green algae) and Prochlorales Lewin 1977. Anoxyphoto bacteriae contains the order Rhodospirillales Pfennig and Triiper 1971 and Chlorobiales ordo nov. The possible need of a fourth division to include organisms that have a rigid cell wall but do not contain peptidoglycan is noted. The name proposed for such a division is Mendocutes, that is, organisms having a cell wall with faults and blemishes. In the 8th edition of Bergey’s Manual of De- (even though Haeckel later accepted Schizo- terminative Bacteriology (4), the Editorial phyta Cohn) for a miscellaneous group of orga- Board of the Manual accepted the kingdom nisms under Protista. Copeland (7) accepted name Procaryotae Murray 1968, 252 (20) but Monera as one of his four kingdoms, preferring was not prepared to support formal names for this name to Schizophyta Cohn 1875, which subdivisions of the kingdom and thus used the implies an association with plants. Bacteriolo- vernacular designations “The Cyanobacteria” gists mostly ignored the kingdom name Monera, and “The Bacteria.” However, Murray (21), in although Stanier and van Niel (28) accepted it an introductory chapter to the Manual, made in their critique of the 5th edition of Bergey’s some suggestions, again mostly in the form of Manual. But even in the 7th edition of the vernacular names, as a basis for the further Manual (3), Protophyta Sachs 1874 was ac- separation of the taxa of bacteria. These have cepted as a division of the plant kingdom, with been discussed further, and a formal nomencla- Schizophyta Cohn as a synonym. ture is now proposed. Margulis (18) has proposed Prokaryota and The division of the living world (on the basis Eukaryota as superkingdoms. Although such a of cellular organization) into procaryotes and rank may be useful for the eucaryotes, so that eucaryotes was first proposed by Chatton (5). the time-honored kingdoms Plantae and Ani- Since the recognition by Stanier (27) and Stanier malia may be retained, the need for such a and van Niel (29) of the utility of this funda- super-rank for Procaryotae is debatable. Al- mental approach, there has been wide accept- though Murray (20) did suggest the taxon Pro- ance of it by those interested in general system- caryotae “at the highest level” and appropri- atics, e.g., Murray (20), Allsopp (l), Whittaker ately at the same level as Eucaryotae, there (33), and Margulis (18). Allsopp (1) stated “. seems no need for the superkingdom Procary- all organisms can be separated into two major otae, the kingdom Monera, and subkingdoms Kingdoms, the Procaryota, which comprise the for the bacteria and aerobic photosynthesizers Bacteria and Cyanophyta, and the Eucaryota as proposed by Margulis. Whether both a su- which includes all other organisms whether perkingdom and a kingdom are required for har- plants, animals or protists.” Whittaker (33), mony with Eucaryotae is another matter and while accepting the fundamental dichotomy, still should be of concern to botanical and zoological included the procaryotes in the kingdom Mo- taxonomists. Lewin (15) sees no point in making nera, a name first proposed by Haeckel in 1866 “prokaryota” synonymous with “bacteria” and seemingly prefers to consider the blue-green pro- t Deceased 10 December 1977. caryotes as algae. If this conservative point of 1 2 GIBBONS AND MURRAY INT. J. SYST.BACTERIOI,. view is generally shared by cyanophycologists, bled conveniently into a fourth division for a subkingdom for the blue-greens may appeal which we propose the name Mendocutes (from to them. We still consider procaryotes and bac- the Latin adjective mendosus, having faults, teria as synonymous. blemishes), This division logically would precede The 8th edition of Bergey’s Manual has re- Mollicutes in a hierarchical classification. cently been criticized by Sonea and Panisset Division I Gracilicutes contains two classes: (26) as ignoring the distinct and unifying aspect (i) Photobacteria classis nov. (type class) and of bacterial genetics. These authors present the (ii) SCOtObQCteriQclassis nov. for the photosyn- view that “instead of accepting the existence of thetic and nonphotosynthetic bacteria, respec- species, genera or families (and presumably tively, The Bacteriological Code (14) is not higher taxa) in the bacterial world, we consider altogether clear on the naming of higher taxa. that the planetary clone consists of many dis- Rule 7 (Names of Taxa above the Rank of tinctly differentiated cells, as in the case of an Genus) could apply to all higher taxa. It states individual tree or an individual animal.” How- quite specifically that the name of a taxon “is ever, they admit that “these currently used in the feminine gender, plural number, . .” names should be kept for practical purposes, However, Rule 8 (Names of Taxa above the even if we know that no speciation exists among Rank of Order) seems to negate Rule 7 at this bacteria.” (Quotations are from English sum- rank and above, since it simply indicates that mary.) Studies of deoxyribonucleic acid and ri- the name is a Latin or Latinized word. Further- bonucleic acid and hybridization experiments more, one of the examples given, Schizomycetes, wiIl presumably provide fundamental data to is a masculine, plural word. We, therefore, feel illuminate such problems. There is close to a justified in using neuter, plural nouns for these century of biological study indicating that many classes. elements of the planetary clone are of sufficient Class Photobacteria comprises two sub- stability to be recognized, whenever they are classes: (i) Oxyphotobacteriae su bclassis nov. isolated, by the characters they exhibit and that (type subclass) and (ii) Anoxyphotobacteriae recognition is a practical necessity. Actually the subclassis nov. organisms seem more stable than either the The subclass Oxyphotobacteriae includes or- names or the bacteriologists who coin them. So ganisms which, by using water as an electron for the present, we prefer to think of a bacterial donor, produce oxygen in the light. For some kingdom rather than a planetary clone, and di- time, all of the blue-green bacteria have been visions and classes seem more practical than considered as belonging to a single order. Re- subclones for the varied and complex group cently, Lewin (16) suggested a new division, considered below. Prochlorophyta, for unicellular, bright green, Formal names are proposed for the higher spherical procaryotes that contain chlorophylls taxa of bacteria, based on the vernacular names a and b, do not contain any detectable bilin pig- proposed by Murray (21, p. 9) but including ments, and show no evidence of phycobilisomes suggestions derived from discussions with col- on the thylakoids. He has now (17) suggested leagues. that these organisms belong to the genus Pro- The kingdom Procaryotue Murray 1968, 252 chloron in the family Prochloroaceae in the is divided on the basis of cell wall types into order Prochlorates. As procaryotes, these un- three divisions: (i) Gracilicutes divisio nov., doubtedly belong in the subclass Oxyphotobac- (type division) which possesses the gram-nega- teriae, and we propose that this subclass com- tive type of cell wall; (ii) Firmacutes divisio prise the order Cyanobacteriales Stanier ordo nov., which possesses the gram-positive type of nov. (type order) to include all blue-green bac- cell wall (which is usually thicker and less dif- teria other than those in the order Prochlorales ferentiated than the gram-negative type); and Lewin 1977, 217. (iii) Mollicutes (Edward and Freundt) divisio The subclass Anoxyphotobacteriae circum- nov., which lacks a true cell wall. These three scribes organisms that carry out a phototrophic groups, suggested by Murray in 1962 (19), have metabolism under anaerobic conditions. It is determinative value. They conserve, for the proposed that the subclass comprise the orders most part, the general arrangement used in the (i) Rhodospirillales Pfennig and Truper 1971, 8th edition of Bergey’s Manual, although, ex- 17 (type order) and (ii) Chlorobiales ordo nov.; cept for Mollicutes, formal ranks were not as- this dichotomy is thus made at the ordinal level signed. A fourth division may be needed for rather than the subordinal level as proposed by organisms that do not have a clear-cut type of Truper (30). cell wall containing peptidoglycan, such as some, No attempt is made to subdivide class I1 Sco- if not all, of the methane bacteria, halophilic tobacteria. Names could be assigned based on bacteria, and Sulfolobus. These can be assem- parts 2 to 12 and 18 of the 8th edition of the VOL. 28,1978 PROPOSALS CONCERNING HIGHER TAM OF BACTERIA 3 Manual, but it is interesting that the authors some of the filaments may be sheathed. Repro- of these parts and sections provided only five duction is mainly by binary fission; in some names at the ordinal level.