INTERNATIONAL JOURNAL OF SYSTEMATIC BACTERIOLOGY VOL. 21, No. 2 April 1971, pp. 197-206 Printed in U.S.A. Copyright 0 1971 International Association of Microbiological Societies Nomenclature of with Special Reference to the Order Actinomycetales'

THOMAS G. PRIDHAM

Northern Regional Research Laboratory,z Peoria, Illinois 61604

The number of names for streptomycetes that is in the scientific literature now is exceeded only by those for organisms placed in the genus Bacillus Cohn 1872. The genus Streptomyces Waksman and Henrici 1943 may well rank in first place if names in the patent and quasiscientific literature are included. The overwhelming number of names and the lack of a precise definition of a particular species or subspecies, of type or neotype strains, and of certain essential details have brought about problems in assessing the status of many names. The major problems encountered in a 2-year study are discussed, and a simple format is suggested, use of which may help to clarify future nomenclature.

Twelve years ago, I presented (29) before ture of Bacteria (20); type strains, where these the First Latin-American Congress for Micro- can be located and obtained, are being as- biology held at Mexico, D.F., some suggestions sembled and recharacterized (35 -38) through on establishing a logical order in streptomycete the International Streptomyces Project, and a classification. minumum set of substrata and tests have been (i) Compilation and evaluation of available recommended for description of A ctino- literature on nomenclature and characterization mycetales in patents (1 1, 12). of streptomycetes. One item upon which insufficient attention (ii) Decision on the proper code of nomen- has been focused is nomenclature. For the past clature to follow and on a common language to several years I have been engaged in a practical describe new taxa. application of the International Code of (iii) Agreement on generic names to be used. Nomenclature of Bacteria (hereinafter referred (iv) Check of type strains and their recharac- to as the Code). My results have been informa- t erization by contemporary met hods. tive, yet disappointing, frustrating, and men- (v) Assignment of numbers to each im- tally exhausting. My experience is an indict- portant named or unnamed strain reported in ment against departments of bacteriology, the literature. microbiology, and biology the world over. (vi) Agreement on designation of a minimum Although I speak primarily to actinomycetolo- set of substrata and tests for characterization of gists, unquestionably my remarks apply to strains. other subdisciplines within microbiology. Too (vii) Determination of ranges of variation many of us either do not know about the Code, within particular taxa, if possible. do not understand it, or simply choose to (viii) Provision for adequate preservation of ignore it. On the other hand, those concerned cultures of type strains. with the development of the Code may not In the ensuing years, considerable advance- have fully realized the impact of applied micro- ment has been made on some of these sug- biology or of the teamwork approach in solving gestions. Actinomycetologists have agreed ( 1) microbiological problems. Applied micro- on using the International Code of Nomencla- biology and the research team present many new problems to the nomenclature of micro- Presented at a round table on Nomenclature of organisms, particularly the A ctinomycetales Actinornycetales at the 10th International Congress and other taxa of industrial-economic interest. I for Microbiology, Mexico, D.F., 9-1 5 August 1970. daresay that a majority of the new taxa which This is a laboratory of the Northern Marketing and Nutrition Research Division, Agricultural Re- confront us now have come from applied search Service, US. Department of Agriculture. microbiology. Therefore its stake in nomencla- 197 198 PRIDHAM INT. J. SYST. BACTERIOL. ture is high. I am particularly concerned be- Only by taking certain liberties can one arrive cause classification and nomenclature seem to at any reasonably accurate contemporary pic- move along separate pathways. Taxonomists ture of the Order as it is known from a should apply the correct names to their taxa. taxonomic viewpoint. The taxonomic informa- Hence, the relationship between the two discip- tion is there, but there are problems associated lines should be closer. with communicating this information through As shown in Table 1, the number of names nomenclature. of streptomycetes in the scientific literature Perhaps we have lost sight of the principal now is exceeded only by the number of names reason for naming a microorganism, i.e., to of organisms placed in the genera Bacillus Cohn facilitate communication. In virtually every 1872 and Bacterium Ehrenberg. (The name of microbiological operation one ultimate ques- the latter genus has been placed on the list of tion that arises is, “What is the name of the rejected names.) organism?” The accuracy of this determination The genus Streptomyces Waksman and eventually reflects the state of the discipline Henrici 1943, when combined with the genus and its progress. Despite the fundamental dis- Harz 1877, most of which are coveries that have come out of pure streptomycetes or streptoverticillia anyway, and despite the excellent chemical work result- may well rank in first place as far as number of ing from studies of secondary metabolites, our named species and subspecies is concerned if means of communicating in terms of the those in the patent and quasiscientific literature microorganisms involved is in a sad state be- are included. The overwhelming number of cause of failures to follow the Code. Or, names, lack of a precise definition of a species perhaps from another viewpoint, the Code may or subspecies, lack of type or neotype strains, be too involved and restrictive. Because of its and lack of essential details associated with complex nature, it may present too many particular names have brought about questions obstacles for practical application. Only in in assessing the status of many names. Principle 8 of the Code do I find reference to My task was a frustrating one complicated an explanation of why microorganisms are initially by translation, the Actinomyces versus named: “. ..to supply a means of referring to Streptomyces, and patent problems. I will taxa.” Savory (34) gives a fuller discussion in discuss some of these and present a simple his interesting little book on nomenclature. Of format that may help to alleviate the situation course, to aid communication any kind of in the future. Unless more attention is paid to system or supplement would suffice, provided the Code, I predict continuing confusion and it is acceptable. Suggestions toward this end much busy work in the years to come, have been made (13, 39), but the binary system especially if one considers the multitude of new supplemented with strain designations still names that might be proposed through applica- appears to be the most acceptable, at least to tion of a typological-nominalistic kind of me, for the Actinomycetales. species concept. The information I have Scientists require that a name be correct gathered suggests that literal interpretation of according to certain rules. However, because of the Code would result in a rather short list of the impact of applied microbiology, we are in a names of organisms belonging to the order position where scientific law and the much Actinomycetales that are scientifically correct. older juridicial law (10) are opposed in certain

No. of pages

Genus IBa IB-1 IB-2 IB-3 Total

Bacillus ...... 188-112 2 11-112 1-114 203-114 Bacterium (illegal) ...... 109 1-1/2 12-1/2 314 123-314 Actinomyces-Streptomyces ...... 77 2-112 8 2-112 90 Salmonella ...... 8 1 0 0 0 81 Pseudomonas ...... 37-314 2-114 0 0 40 Micrococcus ...... 36 112 0 0 36-112

Q IB, reference 8; IB-1, reference 15; IB-2, reference 43; IB-3, reference 44. VOL. 21,1971 A CTINOMYCETALES NOMENCLATURE 199 areas, e.g., Rule 11 of the Code, wherein names taxonomic reports on nomenclature. This of bacteria published in patents are not recog- consequence can be fully appreciated only nized as valid. through study of the Code and implementation Of the three initial broad problems I en- of its “Principles, Rules, and Recommenda- countered, that of translation, though difficult, tions.” Resolving taxonomic problems also will is perhaps the simplest to solve. Lack of ultimately clarify some of the nomenclatural adequate services or capabilities for translating problems. a variety of languages leads to a considerable lag The third major problem I encountered or breach in communication simply because too revolves around Rule 11 of the Code and few people are adept in all foreign languages, concerns names of organisms published in taxonomic papers may not have high priority patents. I am fearful that this Rule may have for translation services, and translations are placed nomenclatural specialists in a position expensive. I am beginning to look with jaun- worse than before the Rule was promulgated. I diced eye on the time, effort, and expense of still am not certain as to the reason(s) for its obtaining translations when much of the infor- proposal. In the review of this manuscript, mation turns out to be of little or no real value. three reviewers offered the following as their Only through much perseverance and the help ideas of the reasons for promulgation of Rule of many colleagues was I able to get reasonable 11: The patent literature is not sufficiently translations of much of the material needed to available unless one knows of its existence, and evaluate adequately the taxonomy and nomen- published patents are not taken automatically clature involved in my study. During this work by even large institutions; they are not edited I pondered with some nostalgia on an older by microbiologists; new names are in most cases requirement for descriptions of microorganisms difficult to find in the patent literature, the in Latin, a language most of us might bungle titles ordinarily do not contain the names, the through with some ease. Lack of a universally names are not indexed, and in many cases the acceptable language will continue to hamper titles do not even give a clue that the patent our efforts, and increased international com- contains a new name; it is impossible to gain munication will amplify the problem. access to the culture upon which the patent The A ctin o my ces-Strep tomyces problem has description is based. In my view, the taxonomy been and is particularly frustrating because of contained in most patents has as muchvalidity the requirement of the Code with respect to as in any other kind of printed matter. The proposal of new combinations [Rule 12c (3)]. same questions would arise in interpreting Lessel’s (24) remarks about this point are worth patent taxonomy as in scientific publications. noting. As it now stands, one must give a very Patents hold a better psoition than most scien- flexible interpretation of the literature where tific printed matter from the standpoint of date many new combinations are implied but not of publication, the date being precisely speci- definitely proposed, as Rule 12c (3) requires. fied to the month and day, whereas publication Unless we resolve the Actinomyces- dates for journals, etc., are not always so Streptomyces controversy, we can look forward precisely specified. Rule 12a further confuses to a double nomenclature for these organisms the issue by allowing the date of patent and much unnecessary documentation of that issuance to take priority, if and when the nomenclature. So far as I am concerned, the patent name is validated in a later-appearing genera Actinomyces and Streptomyces are dif- scientific publication. ferent, and many taxa placed in the genus The only major argument I have with A ctinomyces are, in fact, streptomycetes and patents from a nomenclatural viewpoint occurs streptoverticillia (30). when a new taxon is proposed. Sometimes the Another aspect of the problem of proposing only cited “authority” for the name is a new names and combinations concerns sub- business concern or research organization. How- species. Occasionally an author will propose a ever, the authority problem is in no greater subspecific name without the species name ever question in the patent literature than it is in the having been independently proposed. For scientific literature. As a result of Rule 11, example, Leimgruber et al. (23) proposed the many newly proposed names for organisms in name Streptomyces refuineus var. thermo- the order Actinomycetales, as well as for other tolerans without proposal of the species name organisms, are “in limbo” nomenclaturally. Streptomyces refuineus, which to my know- Such names are used continually by the scien- ledge has not been previously published else- tific and nonscientific community and serve a where. There certainly is a need for tax- useful purpose despite their nonvalidity. But onomists to consider the impact of their there are more problems involved. For one 200 PRIDH AM INT. J. SYST. BACTERIOL. reason or another, the person or persons re- effort, plus occasionally some money, are re- sponsible for coining the name cannot or will quired for the general public or microbiological not validate the, name in a scientific publica- institutions to secure the material in hand. tion. Consequently, there is a real bar to From a list of the kinds of printed matter, a progress in taxonomy, dependent, in final decision could be made as to which of these analysis, upon scientifically acceptable names. constitute effective publication: mimeo- Had Rule 11 not been enacted, the nomen- graphed, multilithed, or reproduced material; clature specialist would simply ferret out the newspapers; descriptive brochures; catalogues; first patent or other publication concerned with abstracts of papers and patents; theses; micro- the new name and resolve the authority prob- film copies of printed matter; dictionaries; lem. Now he not only has to do this task, but encyclopedias; books; monographs; patents; also for proper documentation he should record patent applications; annual reports of scientific all patents and other material issued prior to and industrial organizations; advertisements; the first valid publication of the name. Also advertising brochures; nonscientific, scientific, Rule 11 has brought about the embarrassing and trade magazines and journals; and isolated situation for some taxonomists who use the publications from business and research organi- nonvalid names in scientific publications and zations among others. One solution, of course, then find themselves set down (incorrectly) as would be for scientists to ensure that the first authorities for the name even though they had publication of a new name be documented in nothing to do with isolation of the organism in print in an acceptable scientific journal. Listing question, with its characterization (other than an evaluation of the name Actinomyces olivo- providing additional information obtained in a viridis is an example of the kind of problem taxonomic study), or with coinage of the name. now encountered. Accordingly, I suggest some consideration be A ctin omy ces olivoviridis n. sp. (sic) given to resolving the problem of names in Krasil’nikov, Kuchaeva and Skryabin 1959. patents with the idea of amending Rule 11 so (Supplement to Program for Symposium on that these names can be properly removed from , 18-23 May 1959, Prague, Czechos- limbo. lovakia; Multilithed abstract carrying no pagina- At the start of my work, I was faced with tion). Not Val. Pub. the problem of selecting only validly published Actinomyces olivoviridis n. sp. (sic) and legitimate names requiring, of course, the Kutchaeyeva, Krasil’nikov and Skriabin 1960. evaluation of published material in terms of the (Proceedings of the Symposium on Antibiotics, Code. Some nomenclatural evaluations were 18-23 May 1959, Prague, Czechoslovakia: available in Index Bergeyana (8) and two of the 58-59). Not Val. Pub. Inadequate description. supplements thereto (15, 43). I was not in Actinomyces oliuovirilis (sic) n. sp. (sic) complete agreement with some of the con- K u c ha ev a, Kr asil’ nik ov, Skryabin and clusions drawn therein. My master list still Taptykova, In Rautenshtein (ed.) 1960. (Tr. contained an exceedingly long list of names Inst. Mikrobiol., Akad. Nauk, SSSR 8:248). A from the scientific, quasiscientific, and patent lapsus calami. literature. Also, I was quite aware of the A ctinomyces-strep tomy ces problem. Actinomyces olivoviridis n. sp. (sic) Thus, the first problem was to seek out the Kuchaeva, Krasil’nikov, Skryabin and original source of the name and to track Taptykova In Rautenshtein (ed.) 1960. (Tr. through subsequent references to first valid Inst. Mikrobiol., Akad. Nauk, SSSR 8:25 1 ; publication. It was no easy task to compile and English translation 1966, 249). Val. Pub., Leg. evaluate such lists of references, particularly Aside from the question of validity of when many were in foreign languages and in publication of the name, another problem of patents. An immediate question arose. “What real concern was determination of the proper constitutes an effective publication?” Because authorities for names. One might imply from Rule 11 of the Code singles out patents in rules 14a and 14b of the Code that, for valid particular, one might raise questions about publication of names of species and subspecies, other organs of communication, e.g., abstracts, there is no requirement for citation of the theses, microfilms of theses, and annual reports author and date of publication of the name. of scientific research organizations. It may be Rule 15a states that it is necessary to cite the that we need a list of acceptable publications. author who first published the name in ques- In these days of widespread dissemination of tion in order for the indication of the name of a virtually all kinds of printed matter, with or taxon to be accurate and complete. Possibly as without charge to the recipient, only time and a consequence of the separation of Rules 14 VOL. 21,1971 ACTINOMYCETALESNOMENCLATURE 20 1 and 15, and possibly because tackling the then might be raised with respect to a defini- authority problem is just too much for the tion for the term “authority.” The person who average microbiologist, particular combinations first coins a name might be an attorney, a of generic names and specific or subspecific chemist, a Latin or Greek scholar, or the epithets are accompanied by many different microbiologist who isolated and characterized citations of authority in the literature. The job the organism. “Published” may be the key of determining the proper authority thus be- word in Rule 15a, but it needs clarification. comes one of considerable magnitude. For Another example in point is Streptomyces example, Table 2 lists some “complete” names hydrogenans. So far as I have determined, the containing the epithet “albus” I have found in name was first proposed in a German “Aus- the literature. gelegeschrift” by Lindner et al. (26) and hence The same situation can be found with other is not validly published. A valid publication of common specific epithets used in the names of the name is cited in Index Bergeyana with the organisms placed in the Order A ctinomycetales, authorities Lindner et al. (25). However, a e. g., “aureus, ” “griseus, ” “roseus, ” and notation in this paper states that K. Wallhausser “viridis. ” is credited with the investigation and charac- Obviously there is much confusing informa- terization of the taxon. In the old days of tion in the literature with respect to authorities bacteriology, lone scientists often isolated or- for names. When only one person is listed as the ganisms, characterized them, conducted taxo- author of a publication there is no problem, but nomic evaluations, and coined the names. Now, in these days of the research team, single with the teamwork approach to many problems authorship is an exception rather than the rule. in microbiology, each of these operations may When more than one author is listed, many be carried out by different individuals. We need questions arise. In some papers five or more a precise definition of what constitutes an authors are listed, for example: authority for a name. The problem is further A ctinomyces grisinus Krasil’nikov, complicated by the practice on the part of Belozersky, Rautenstein, Korenyako, Nikitina, some microbiologists of using personal names 3s Sokolova and Uryson 1957. strain designations. These have been construed S t r e p tomy c es c i n n a moneus Benedict, by some as the names of the authors of the Dvonch, Shotwell, Pridham and Lindenfelser name of the species or subspecies. One major 1952. collection used this device for many years. Rule 15a of the Code states that these could Microbiologists should exert all possible be reduced to the senior author followed by the pressure to see that the proper authority, once abbreviation et al. Obviously, not all the we have established exactly what that is, is authors worked on characterization of the connected with the name of a new taxon organism or in coining its name. The question somewhere in the body of the printed matter. Insertion of a simple statement such as, “The TABLE 2. Authorities cited for authors decided to assign this strain to a new Actinomyces albus as taken from various species S. kanamyceticus n. sp. Okami et literature sources Umezawa.” would solve the problem. This form was used by the Japanese workers in this A. albus Gasperini, 1894. particular example (41). A more formal state- A. albus (Rossi Doria) Gasperini, 1894. ment might have been: “We propose that the A. albus (Rossi Doria) Krassil’nikov, 1949. new taxon be named Streptomyces kanamy- A. albus Krainsky, 1914. ceticus Okami and Umezawa.” Comparable A. albus Waksman et Curtis, 1916. statements would allow continuity in authority A. albus Lehmann et Neumann. designation from the first printed material A. albus Gasperini, 1890. distributed. They also are proper for inclusion A. albus (Krainsky) Waksman et Curtis. in patents and published patent applications, A. albus (Krainsky, A., 1914) emend. Waksman S.A. provided no juridicial barriers exist, though et R. E. Curtis, 1916. these sources are nonvalid publications now. In A. albus (Rossi-Doria, 1891) Krainsky, A., 1914. my view, the problem of authorities for names requires reexamination with reference to the A. albus (Krainsky) Waksman et Curtis, 1919, A. albus Tarozzi, 1909. Code. Another question related to valid publi- A. albus Waksman et Curtis, 1916 vide etiam Waksman, 1919. cation concerns Rule 12a, wherein it is stated that a name to be validly published must be A. albus Krasil’nikov, 1941. accompanied by a description of the taxon or 202 PRIDHAM INT. J. SYST. BACTERIOL. by a reference to a previously and effectively sideration be given to this adequacy require- published description of it. The question of ment in establishing valid publication. As a whether the organism is described sufficiently matter of fact, a contemporary taxonomist so that the name applied to the organism is not could reevaluate the entire Order Actinomyce- a nomen dubium (a name whose application is tales and eliminate many older names based on uncertain) is left to the individual. Often adequacy of description alone. descriptions leave much to be desired insofar as A minor problem, but one for which some adequacy is concerned, and, with the kinds of note should be taken, concerns the final entry organisms with which we are dealing, today’s in the complete name for a taxon, i.e., the page adequate description may be inadequate 5 years on which the proposal for the new name hence. Examples of names accompanied by appears. In some publications, the new name is inadequate descriptions are: referred to several times in various parts of the Streptomyces floridae-a raspberry to violet- manuscript. For example A. rectiviozaceus colored actinomycete; produces viomycin (3). Artamonova in Krasil’nikov 1965 is specified as Streptomyces colombiensis-produces vita- “n. sp.” in the title of the paper (2); in the min BI2 (31). introduction on page 234; in the title for an Streptomy ces natalensis - produces pimaricin illustration on page 235; in a table on page 236; (40). in a section of physiological and biochemical Actinomyces diastatochromogenes-cultural characteristics on page 237; in a table of carbon and physiological characteristics only (22). utilization results on page 238; on pages Strepto my ces sub trop,icus -produces alb o- 239-241, 245-247, and 249; and finally, in mycin complex (6). No such name in publica- boldface type heading a general summary of the tion. characteristics on page 250. Zvirbulis and Hatt Streptomyces toyocaensis- produces toyoca- cited page 250 in their evaluation of the name mycin (28). (43). However, what is the rule here? Person- Streptomyces jamaicensis-produces mona- ally, I believe the pagination should refer to the mycin (14). first page on which the name occurs, regardless Each of these names is evaluated as being of whether it is in the title or of how close it is validly published in Index Bergeyana. None had to a description. In the paper by Artamonova an adequate description in my opinion. I (2), the description is scattered throughout the submit that Rule 12a of the Code might pages and is summarized on page 250, whereas contain the statement, “. . . must be ac- the implied proposal of a new taxon lies in the companied by an adequate description of the title of the article. taxon or by a reference to a previously and Another major problem encountered during effectively published adequate description of my work lay in documenting the designation of it.” Otherwise, the only requirement is for the type strain of a given taxon. In the great effective publication, i.e., the determination of majority of published descriptions of strepto- whether printed matter containing the name mycetes and streptoverticillia, there is no pre- represents an acceptable publication, i.e., cise designation of a type strain. Despite some whether it is a scientific publication. objections to the type-strain concept, it appears Requiring an adequate description, of to be the only practical approach to take when course, raises the question, “How does one confronted with a multispecies genus such as determine what constitutes an adequate des- Streptornyces. It really does not matter what cription of a taxon, particularly when the position a type strain may occupy in the species taxonomy is continually subject to the addition sphere, and certainly one could never hope to of new criteria?’’ It would seem that the various cover the entire range of variation exhibited by international subcommittees on taxonomy a particular taxon. Thus, from a systematic could answer this question. For example, our viewpoint, there is need for a bench mark of International Subcommittee on Taxonomy of some kind, i.e., the type strain. Some descrip- A ctinornycetes already has recommended cer- tions obviously were based on study of a single tain minimal criteria for use in descriptions strain. These can be assumed to be the type appearing in patents (1 1, 12). These same strain (single isolate), Rule 9d (2). Unfortu- minimal criteria could be recommended for nately, sometimes the type strain was not characterization and description of any assigned a strain designation; cultures dis- organism in the order Actinomycetales in any tributed as belonging to the type strain bore printed matter and could be revised at periodic designations different from the published desig- intervals. I therefore suggest that some con- nation for the type strain; different individuals VOL. 21,1971 ACTINOMYCETALES NOMENCLATURE 203 have received different strains on request for indicating the particular strain number. Use of the type strain; and so on. All sorts of the names of individuals or localtities as strain variations of this theme were encountered, and designations may result in confusion with the the search for the correct original strain desig- names of authorities for the taxon in question nation often was tedious. Also, where no strain and are not recommended.” Thus, NRRL of several listed or characterized is designated as B-150, NIHJ-236, ATCC 17560, CBS 123.39, the type strain in the original publication, and INMI 18-18 are appropriate designations anyone may designate a type strain for that that adequately pinpoint particular strains and taxon, but it has to be one of the strains on can serve effectively in communication either which the original description of the species or with or without the formal name of the subspecies was based and presumably would be organism. This method for strain designation listed andlor characterized in the original publi- has been used by the major culture collections cation. Any other strains are simply reference for many years. strains or potential neotype strains. This lack of Actinomycetologists should pay particular formal designation of the type strain was a attention to the selection of the type strain and major obstacle in getting at the facts. Inter- to its appropriate designation in the initial estingly, I had to refer to some papers which abstracts, papers, or patents wherein the name were not effectively published in ferreting out is proposed. There is no question in my mind the original strain designations for some strep- that with the organisms with which we are t omycetes. concerned, there is need for a type strain Another point of concern has been the appropriately designated, regardless of its par- misconception that the authority also refers to ticular taxonomic position. All other strains the strain designation, i.e., that S. griseus represent reference strains and potential neo- (Krainsky) Waksman and Henrici, with no types should the type strain be lost. strain designation as listed in culture collection Another major problem I encountered con- catalogues or other publications, refers in fact cerns derivation or coining of specific and to progeny of Krainsky’s original strain isolated subspecific epithets. Frequently, allusions are ca. 1914. This concept is not true. Progeny of made in the text as to the derivation of a most, if not all, of Krainsky’s original isolates particular name. In other instances, for ex- no longer exist. Names abcompanied by their ample, “kanamyceticus, ’’ there is no indication. proper authority citation also should be ac- The seventh edition of Bergey’s Manual of companied by the correct strain designation(s) Determinative BacterioZogy (7) neatly presents in taxonomic works. It is beyond my compre- this information, i.e., the syllabication of the hension why a systematist cannot, on first epithet followed by its etymology. For ex- isolation and purification of members of a ample, for S trep t o myces violaceoniger particular taxon, assign to the members strain (Waksman and Curtis, 1916) Waksman and designations and, moreover, select one of a Henrici 1948, 947 one finds: vi.o.la.ce.o’ni.ger. series of similar strains as the type strain for the L. adj. violaceus violet; L. adj. niger black; M.L. new taxon he intends to name. The particular adj. uiolaceoniger violet-black. strain may represent the central point or either I attempted to work out many of the names end of a range of variation. This problem would included in my exercise for which no syllabica- have to be worked out later and actually tions or etymologies were given. It took about represents one part of a lumping process or 30 min per name using publications such as population study. Collin’s Latin Gem Dictionary (2 l), Marchant Beyond the selection of the type strain and and Charles’ revision of Case113 Latin Diction- its designation, its documentation could be set ary (no date), Jaeger’s A Source-Book of’ forth in patents or other publications by Biological Names and Terms (18), Cash’s A insertion in the text of a simple statement such Mycological English-Latin Glossary (9), and as, “The type strain of this taxon is (2- to Webster’s New Unabridged Dictionary (42), as 5-letter abbreviation for research organization well as many other foreign language dictionaries or culture collection)-( 1- to 6-number desig- and sources of information. There are, of nation),” e.g., NRRL B-1699. Accordingly, I course, later editions of some of the references suggest modification of Rule 8a (1) to include a cited. An unabridged Latin dictionary should statement such as, “It may be designated in any also be used for such work. manner, preferably by two to five letters One complication occurred because Rule 6 indicating the collection or laboratory housing of the Code states that a specific epithet may the organism, followed by one to six numerals be taken from any source whatever and may 204 PRIDHAM INT. J. SYST. BACTERIOL. even be composed arbitrarily. As a result, I still Index Bergeyana. Any strains (if they are have some names that defy solution. If names available) carrying such questionable names are composed arbitrarily, such a notation should be considered unnamed until taxono- should be made and some explanation of the mists either rename them more appropriately or reason for the epithet should be given. assign new, correct names to them. Table 3 lists There now is a formal list of names in Index the epithet “citreus” and its feminine and Bergeyana and its several supplements, and I neuter forms as now represented in the order recommend that the epithets listed therein not Actinomycetales. In my opinion these deserve be used in coining names of new taxa in the the same fate as “aureus. ” A ctinomycetales. I encountered several ex- Further, I believe the practice of correcting amples where the same name was proposed for the original orthography of an epithet when the different taxa. Accordingly, I suggest that an epithet is misspelled leads to even greater “absolute epithet priority rule” for the order confusion despite good intentions. There are A ctinomycetales be considered and that evalua- examples now of accepted names where a slight tion of the nomenclature and taxonomy of a difference in spelling exists for two completely particular species or subspecies in the order d iff e re n t taxa-strep tomy ces can us (sic) Actinomycetales Buchanan 1917 should be Heinemann, Kaplan, Muir and Hooper, 1953, based on the earliest history associated with a 1239 and Actinomyces canis (sic) Baudet 1934, specific or subspecific epithet. This action 301. Canus is the Latin adjective meaning white would broaden Rules 6 and 7. For example, as or gray; canis is the Latin noun meaning dog or outlined in Pridham et al. (32), the first use of bitch. On the other hand, there are examples of the epithet “aureus, ” or its feminine and neuter epithets differently spelled that might be con- forms “aurea ” and “aureum ” in referring to an strued as representing different taxa but are actinomycete, was by Johan-Olsen ( 19). Sub- referring to one and the same taxon and, in sequently other investigators, apparently un- fact, to the same strain; e.g., Streptomyces aware of his proposal, coined presumably new canescens (17) Waksman 1957, 968, and Strep- names with one or the other of the epithets tomyces canescus Hickey, Corum, Hidy, Cohen, cited. These probably referred to entirely dif- Nager and Kropp 1952, 473. I, myself, un- ferent organisms. Application of an “absolute wittingly perpetrated similar confusion in con- priority’’ concept would help to clear the air nection with the nomenclature of the cinna- with respect to the taxonomy of some of the mycin-producing organism. In attempting to older “species.” There should be only one ” ” or and that with “aureus, “aureum, “aurea” TABLE 3. Names of Actinomycetales that the basionym A ctinomyces aureus Johan-Olsen contain “citreus,’’ “citrea, ’’ or “citreum ” 1893, 79. Furthermore, the taxonomy of that as specific or subspecific epithets taxon should be based on the characteristics cited in his publication. As pointed out earlier Date of (32), study of the reported characteristics for Name proposal strains with the specific epithets “aureus, ” ” or ” assigned in the literature “aurea, “aureum Actinomyces citreus ...... 1894,1914 on especially in connection Actinomycetales, Streptothrix citrea ...... 1896 with the genera A ctinomyces, Streptothrix, Nocardia citrea ...... 1916,1948 and (and I Nocardia, Discomyces, Oospora Proactinomyces citreus ...... 1938 might add suggests that the Streptomyces), Actinomyces jlavus subsp. citreus ... 1941 species (Actinomyces aureus, Streptothrix Mycobacteriu m citreu m ...... 194 1 and aurea, Nocardia aurea, Oospora aurea, M. citreum subsp. flavoviride ...... 1941 Streptomyces aureus) are taxonomically ques- M. citreum subsp. liquefaciens ..... 1941 tionable and should not be recognized. Any M. citreum subsp. paulotrophum ... 1941 names other than A ctinomyces aurcus Johan- M. citreum subsp. putrificum ...... 1941 Olsen 1893, 79 which contain the specific Streptomy ces citreus ...... 1948 epithet aureux or its orthographic variants Proactinomyces citreus subsp. simply represent misidentifications based on marinae ...... 1949 lack of knowledge of the history associated M. citreum subsp. mucosum ...... 1949 with the epithets ” ” or “aureus, “aurea, M. citreum subsp. lacticum ...... 1949 as they relate to the order “aureum” Actino- Promicromonospora citrea ...... 1961 mycetales. Many names such as these already P. citrea niger ...... 1961 have been rejected by taxonomists through Actinomyces candidus subsp. citreus . 1964 omission. Others have been rejected formally in VOL. 21,1971 ACTINOMYCETALESNOMENCLATURE 205 correct what I believed to be an error in spelling tempted simply to ignore the Code. Hopefully of the epithet, I changed the name from they will not. Simplification of the Code might Streptomyces cinnamoneus (sic) Benedict, help, lest we become so embroiled in promul- Dvonch, Shotwell, Pridham and Lindenfelser gation of additional new rules and regulations 1952, 591, to Streptomyces cinnamomeus (33). as to exert a stifling effect on progress in the It would have been better had I left the original com bined area of systematics-nomenclature. I spelling of the epithet intact since now there purposely have selected examples to illustrate exist about a half dozen different names for the international aspects of the problems. this taxon. I have presented this material to illustrate LITERATURE CITED the problems that arise when an average micro- 1. Anonymous. 1963. News and Notes. Action of the biologist attempts to coordinate taxonomic Subcommittee on Taxonomy of the Actino- studies with nomenclature. Such coordination mycetes. Int. Bull. Bacteriol. Nomencl. Taxon. should be the obligation of any systematist. 13(3):175. However, it is not so easy as it may seem. Some 2. Artamonova, 0. E. 1965. New species of violet of the problems I have discussed can be handled colored actinomycetes Actinomyces rectivio- by following a simple format (30) in proposing laceus n. sp., p. 234-251. In N. A. Krasil’nikov a new name or new combination as illustrated (ed.), Biology of individual groups of Actino- in the following example concerning mycetes. (In Russian) Moscow. Actino- 3. Bartz, Q. R., J. Ehrlich, J. D. Mold, M. A. Penner, myces a1 bohe1 vat us. and R. M. Smith. 1951. Viomycin, a new For a new name. “We propose that this new tuberculostatic . Amer. Rev. Tubercl. taxon be named Actinomyces albohelvatus 63( 1):4-6. Krasil’nikov, Korenyako and Nikitina. 4. Baudet, E. 1934. Actinomycose du chien. Ann. The type strain of Actinomyces albohelvatus Parasit. Hum. Comp. 12:296-308. is INMI 1349 by original designation. 5. Benedict, R. G., W. Dvonch, 0. L. Shotwell, T. G. Etymology: al.bo.hel.va’tus L. adj. albus Pridham, and L. A. Lindenfelser. 1952. Cinna- white; L. adj. honey-yellow; M.L. adj. mycin, an antibiotic from Streptomyces cinna- hebatus moneus nov. sp. Antibiot. Chemother. albohelvatus light honey-yellow, referring to 2591 -594. the color of the vegetative mycelium of the 6. Brazhnikova, M. G., N. N. Lomakina, and L. I. organism. ’’ Murav’eva. 1954. Albomycin, its characteristics For a new combination. “Streptomyces and chemical nature. Dokl. Akad. Nauk. SSSR albohelvatus [ Krasil’nikov, Korenyako and 99:827-830. 7. Breed, R. S., E. G. D. Murray, and N. R. Smith. Nikitina in Krasil’nikov (ed.), 19651 comb. 1957. Bergey’s manual of determinative bac- nov. teriology, 7th ed. The Williams & Wilkins Co., Type strain: (none by original designation) is Baltimore. INMI 1349 (by subsequent designation by 8. Buchanan, R. E,, J. G. Holt, and E. F. Lessel, Jr. (ed.). 1966. Index Bergeyana. An annotated Krasil’nikov in Shirling and Gottlieb (38) for alphabetic listing of names of the taxa of the International Strep tomy ces Project ). bacteria. The Williams & Wilkins Co., Baltimore. S y n o n y m y : A c t ino myces alb o h el va t us 9. Cash, E. K. 1965. Mycological memoir no. 1. K rasil’nikov, Korenyako and Nikitina in A mycological English-Latin glossary. Hafner Krasil’nikov (ed.) 1965, 224. Publishing Co., New York. Etymology: al.bo.hel.va’tus L. adj. 10. Cranberg, L. 1968. Law-scientific and juridicidl. afbus Amer. Sci. 56(3):244-253. white; L. adj. helvatus honey-yellow; M.L. adj. 11. Gottlieb, D. 1963. Recommendations for descrip- albohelvatus light honey-yellow, referring to tions of some Actinomycetales appearing in the color of the vegetative mycelium of the patent applications. Int. Bull. Bacteriol. organism.” Nomencl. Taxon. 13(3):169-170. If our present approach to taxonomy and 12. Gottlieb, D. 1964. Recommendations for descrip- nomenclature of the Actinomycetales is to tions of’ some Actinomycetales appearing in continue, the simple formats outlined above patent applications. Amer. SOC.Microbiol. News would do much to lessen the labors of whoever 30(2):13- 14. might be involved in collating information on 13. Gould, S. W. 1954. Permanent numbers to supple- the large numbers of new names and new ment the binomial system of nomenclature. Amer. Sci. 42:269-274. combinations that are bound to appear. 14. Hassall, C. H., and K. E. Magnus. 1959. Mona- I hope that my remarks might bring about mycin: a new antibiotic. Nature (London) some dialogue and alert systematists to the 184: 1223- 1224. consequences of their handling of nomenclature 15. Hatt, H. D., and E. Zvirbulis. 1967. Status of in any kind of printed matter. As it now stands, names of bacterial taxa not evaluated in Index I can see why some taxonomists might be Bergeyana (1 966). 1. Names published circa 206 PRIDHAM INT. J. 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